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PRIT 


NOTA 


+ lepidopterologica 


Vol. 18 No.1 1995 ISSN 0342-7536 


ci Vier Preset: | Miclinet Fibiger 
U : Manfred Sommerer — 
rH ee. 


for members] ip, changes of se and orders for Nota ip 
volumes and other. er should: be sent to the = seien 


mé 


mini may order Nota lepidopterologica through 6 our 
o Books Kirkeby Sand 9 DK- 771 Stenstrup, De a 


Nota lepidopterologica 


Vol.18 No. 1 Basel, 31.1.1996 ISSN 0342-7536 


Editor : Steven E. Whitebread, Maispracherstrasse 51, P.O. Box 27, CH-4312 Magden, 
Switzerland. FAX : +41-61-8412238. 


Assistant Editors: Emmanuel de Bros (Binningen, CH) 
Dr. Roger Dennis (Wilmslow, GB) 
PD Dr. Andreas Erhardt (Binningen, CH) 
Dr. Yves Gonseth (Neuchätel, CH) 
Dr. Alexander Pelzer (Wennigsen, D) 


Contents — Inhalt — Sommaire 


Dennis, R. L. H.: Oviposition in Zerynthia cretica (Rebel, 1904) : 
loading on leaves, shoots and plant patches (Papilionidae) .................. 3 
Kozıov, M. V.: Subalpine and alpine assemblages of Lepidoptera in 
the surroundings of a powerful smelter on the Kola Peninsula, NW 
RS STE EN SL EN ARR Rise MN (7 
Kozıov, M. V. & Rosinson, G. S.: Identity and distribution of two 
dimorphic oriental fairy moths — Nemophora decisella (Walker, 1863) 
and Nemophara cantharites (Meyrick, 1928) (Adelidae)...................... 39 
SANETRA, M. & FIEDLER, K. : Behaviour and morphology of an aphyto- 
phagous lycaenid caterpillar : Cigaritis (Apharitis) acamas Klug, 1834 
(Lyeaenidae) an na a LU ke A A 57 
SHREEVE, I. G., Dennis, R. L. H. & WırLıams, W. KR. : Uniformity 
of wing spotting of Maniola jurtina (L.) in relation to environmental 


hessnoesneiy (SAN EMA) I... nen. V7 
Book reviews — Buchbesprechungen — Analyses 38 
Xth European Congress of Lepidopterology ...................n... 2 
Butterfly Conservation’s 2nd International Symposium ........ 16 


Nota lepid. 18 (1) : 2 ; 31.1.1996 ISSN 0342-7536 


Xth European Congress of Lepidopterology 


Miraflores (Madrid, Spain) 
3-7th May 1996 


The Council of the Societas Europaea Lepidopterologica (SEL) kindly 
invites all lepidopterists to attend the Xth European Congress of 
Lepidopterology to be held in Miraflores (Madrid, Spain) from 3rd 
to 7th May 1996. The site of the Congress is the University Hostel 
“La Cristalera”, at Miraflores de la Sierra, which is situated in the 
Sierra de Guadarrama, about 50 km north of the city of Madrid. 


Main topics will be : 


General and specific problems of taxonomy and evolution 
Ecology, endangered species and species protection 
Holarctic zoogeography and local faunistics 

New discoveries in Microlepidoptera 

Tropical Lepidoptera 

Applied lepidopterology 


For more details please contact : 


Xth European Congress of Lepidopterology, 
Department of Biology, 

Universidad Autonoma, 

E-28049 Madrid, 

Spain 


Tel. : +34/ 1-3978281 
Fax : +34/ 1-3978344 


Nota lepid. 18 (1) : 3-15 ; 31.1.1996 ISSN 0342-7536 


Oviposition in Zerynthia cretica (Rebel, 1904) : 
loading on leaves, shoots and plant patches 
(Lepidoptera, Papilionidae) 


Roger L. H. DENNIS 


Department of Biological Sciences, Keele University, Keele, Staffs. STS 5BG, U.K. 
(Address for correspondence : 4 Fairfax Drive, Wilmslow, Cheshire SK9 6EY, U.K.) 


Summary 


Zerynthia cretica females tend not to lay eggs on small plant patches with 
small leaves, unless there are few or distant neighbouring plants. On plants 
chosen for oviposition, females lay proportionately more eggs on large plant 
patches with large leaves, typically at the plant patch margin, and tend to 
load eggs on fewer than the expected number of shoots and leaves. In Z. 
cretica, egg loading on plants may be facilitated by variable clutch size and 
the tendency of females to add to egg batches laid by other females on the 
same leaves and shoots ; direct observations are required on both these aspects 
of behaviour. Some of the potential consequences of egg loading on hostplant 
patches for the population dynamics of the insect are discussed, particularly 
the influences of drought and parasitism. 


Résumé 


Les femelles de Zerynthia cretica ont tendance a ne pas pondre leurs ceufs 
sur de petites touffes de plantes avec de petites feuilles, à moins que les plantes 
des alentours soient peu nombreuses ou éloignées. Sur les plantes choisies 
pour la ponte, les femelles pondent proportionnellement plus d’ceufs sur les 
grosses touffes de plante avec de grandes feuilles, typiquement a la limite 
de la touffe de plante. Pour la femelle de Z. cretica, la ponte des ceufs sur 
la plante peut étre facilitée par la dimension variable des touffes et la penchant 
des femelles a ajouter leurs ceufs aux paquets pondus par d’autres femelles 
sur les mémes feuilles et pousses ; il faut arriver a des observations directes 
sur ces deux aspects du comportement. L’auteur discute des conséquences 
possibles de la ponte des œufs sur les touffes de la plante nourricière pour 
la dynamique de population de Z. cretica, en particulier les influences de la 
sécheresse et du parasitisme. 


Zusammenfassung 


Zerynthia cretica-Weibchen legen Eier tendenziell nicht auf kleine Gruppen 
kleinblättriger Pflanzen, außer wenn es insgesamt nur wenige oder nur weit 
entfernte Nachbarpflanzen gibt. Auf den für die Ablage ausgewählten Pflanzen 
legen die Weibchen verhältnismäßig mehr Eier auf große Gruppen großblätt- 
riger Pflanzen, insbesondere am Rand der Gruppen, und sie neigen dazu die 
Eier auf weniger Triebe und Blätter zu legen, als zu erwarten wäre. Bei Z. 
cretica wird eine Anhäufung von Eiern auf einzelnen Pflanzen möglicherweise 
dadurch gefördert, daß die Gelegegröße variabel ist und daß Weibchen neue 
Eier zu bereits vorhandenen Gelegen hinzufügen. Direkte Beobachtungen 
beider Verhaltensaspekte sind wünschenswert. Einige der potentiellen Kon- | 
sequenzen des Ablageverhaltens auf Pflanzengruppen für die Populationsdy- 
namik des Insekts werden diskutiert, insbesondere der Einfluß von Trockenheit 
und Parasitismus. 


Introduction 


In Crete, the endemic butterfly Zerynthia cretica oviposits on the 
endemic plant Aristolochia cretica (Lam) (Aristolochiaceae). Two other 
Aristolochia species are present in the island group (Aristolochia 
sempervirens L. and A. parvifolia Sm.) but the butterfly only occurs 
near A. cretica (L. Chilton, pers. comm.). A. cretica occurs in two 
distinct habitats: (1) by shady, cool limestone and dolomitic rocks, 
typically with a northern aspect or screened by the slopes of cliffs and 
gorges ; (1) on dry soil under cultivated olives. Z. cretica occurs in 
both habitats in southern Crete, but in the vicinity of Plakias is more 
commonly associated with the former habitat (N = 5) than the latter 
(N = 1). A. cretica is also on the island of Karpathos, but Z. cretica 
has not been recorded there. Elsewhere, close relatives of Z. cretica 
use other Aristolochia species. On Rhodes, the supposed sibling species 
to Z. cretica, Zerynthia cerisy (Godart) must use one or both of 
Aristolochia guichardii (Davis & Khan) or A. parvifolia (Sm.) (Olivier, 
1993). Between April 11-23, 1990, some data were collected on egg 
distributions of Z. cretica on A. cretica plant patches at Plakias in 
southern Crete. During that period, few adults were still evident, but 
the flight season was nearing an end. Thus, the bulk of the eggs had 
been laid and observations were restricted to determining egg distri- 
butions on the hostplant. 


Study area and methods 


Observations were made in a small area (70x 40m; Fig. 1) on 
Kakomouri headland, Plakias. The area was mapped, divided into a 


4 


Fig. 1. Study site on Kakomouri headland, Plakias, for Zerynthia cretica. The hostplant, 
Aristolochia cretica, is also found on the surrounding cliffs. 


number of topographically distinct zones and the number of Aristolochia 
cretica plant patches enumerated. Some 110 plant patches were 
identified in 11 zones. Of these, 33 patches (30%) marked for 
measurement, were selected so as to represent an unbiased sample of 
the plants accessible for survey on this part of the headland. The mean 
aspect of the plant patches was north-north-west (339° + 29°), A 
number of measurements were made on each plant patch : maximum 
length and breadth (PATCH SIZE = length x breadth, cm?) ; maximum 
patch height (PLANT HEIGHT, cm) ; the number of shoots (SHOOT 
NUMBER) ; the number of leaves (LEAF NUMBER, those on large 
plant patches were estimated from a sample of shoots along the axes 
taken for plant patch length and breadth); the maximum leaf size 
(LEAF SIZE, = 14146 cm?, where / and b are respectively the length 
and maximum breadth of the largest leaf) ; the mean distance to the 
three nearest neighbouring patches (MEAN DISTANCE, cm) and the 
number of patches within a 5m radius (DENSITY). The degree of 
exposure to sunlight (EXPOSURE, length of time, h) and the toughness 
of leaves (LEAF TOUGHNESS) of each plant were determined on 
a three and four-point ranked scale respectively. Each shoot and leaf 
was examined for the number of juvenile Z. cretica (eggs, cases, larvae), 


5 


providing data on the number of Z. cretica for individual leaves, shoots 
and plant patches. Also observed were egg colour, the position of eggs 
on each shoot (leaf number from shoot tip) and the position of eggs 
on leaves (upper- or undersurface). 


The distributions of all variables were tested for normality (Kolmogorov 
Smirnov test ; Shapiro-Wilks’ W test) and transformed as necessary. 
Some variables could not be normalised (EXPOSURE, LEAF TOUGH- 
NESS) and thus distribution-free tests were generally used. Correlations 
are Spearman’s r,. Multiple regression applied stepwise entry criteria 
controlled by significant F values and limited to four variables in any 
analysis. The data reduction technique applied principal factors analysis 
with VARIMAX orthogonal rotation of the factor axes (Nie et al, 
1975); 


Results 


Between April 11-23, all juvenile stages of Z. cretica were found from 
recently laid eggs up to 4th instar larvae. Adults were sparse. In the 
previous year, the peak adult flight period occurred in late March 
(Dennis, pers. obs.). The eggs are large, spherical and prominent on 
the hostplant (see Hensle, 1993). At first, they are yellow, but become 
brown. On hatching, the larvae do not consume the egg case, which 
remains attached to the plant (Fig. 2). Larvae typically eat a hole 
through the leaf to produce characteristic feeding damage (Fig. 2) to 
continue feeding on the upper surface. A number of larvae can occur 
exposed on the hostplant simultaneously and are highly apparent (Fig. 
3) though there is no obvious gregarious behaviour as in the case of 
Pieris brassicae L. (Pieridae) or Aglais urticae L. (Nymphalidae) (see 
Dennis, 1992 : 119). Defence mechanisms include rows of spines and 
an orange forked osmaterium (Fig. 4). 


Some 4696 leaves were examined. Of these, 4353 leaves had no eggs 
and 343 had one egg or more (mean, 0.125; variance 0.154). The 
distribution of eggs on leaves has been compared with a poisson 
distribution ; this reveals that there is a significant excess of leaves 
without eggs (observed 4353 vs expected 4142) and with two eggs or 
more (observed 133 vs expected 37) and a deficit of leaves with just 
one egg (observed 210 vs expected 517) (x72 = 442.1, P< 0.001). As 
there is a possibility that these results may be biased by including data 
from plant patches which received no eggs at all, the comparison was 
repeated excluding leaves from eight patches apparently lacking 
oviposition events. The previous observation was confirmed 
(Xe = 437.7, P < 0.001). As many as nine eggs were found on a single 


6 


Fig. 2. Five egg cases of Zerynthia cretica and early larval feeding damage on a leaf 
of Aristolochia cretica. 


leaf, but it was evident that when large numbers (N > 3) were found 
on leaves, they were probably the result of separate oviposition events 
by the same female on different occasions or by different females : 
(i) varying egg colour, cases and larvae indicated that they had been 
laid at different times, days if not weeks apart ; (11) eggs also occurred, 
loosely, in separate “batches”, occasionally on different leaf surfaces. 
However, as it is possible that eggs of the same or different colour 
could have been laid by different females and that eggs laid apart on 
the same leaf could have been laid by the same female at much the 
same time, it is not possible to make an accurate assessment of “brood” 
size, not without following females and directly observing oviposition 
events. 


The vast majority of eggs (N = 586) were laid on the underside of 
leaves (90%) ; a small fraction were laid on the upper surface of leaves 


7 


Fig. 3. Apparency of Zerynthia cretica larvae on the hostplant : loading of individuals 
on a hostplant patch, exposed feeding and thermoregulation on the leaf upper surfaces, 
obvious feeding damage of leaves and frass accumulation. 


(9%), and occasionally eggs were found on stems and on adjacent plants 
(1%). Even the test difference between leaf surfaces for leaves with 
eggs, rather than for number of eggs, indicates that the leaf undersurface 
is highly favoured compared to the leaf uppersurface for oviposition 
GE = 207.852 20.000): 


Oviposition on shoots reveals much the same bias as for individual 
leaves. Some 655 shoots were examined. Eggs were found on 217 shoots 
(mean 0.533 ; variance 0.856). Compared to a poisson distribution, 
significantly more shoots than expected had two leaves or more with 
eggs (observed 87 vs expected 65), and fewer shoots than expected 
had a single leaf with eggs (observed 130 vs expected 205) (42, = 56.0, 
P<0.001). This difference remained when the eight plant patches 
without oviposition records were excluded (#21, 19.2, P < 0.001). Such 
biased loading of eggs on a smaller than expected number of shoots 
and leaves could be the result of female preference for any differences 
in host quality, but it may also be influenced by differences among 
shoots and leaves in accessibility to females. Moreover, any difference 
would be compounded by ’clutch’ laying. Thus, it is notable, that on 
one circular-shaped hostplant patch, although central plants were not 
entirely avoided, the eggs were nevertheless loaded around the perimeter 


8 


Fig. 4. Mature larva of Zerynthia cretica, illustrating spinal defences and osmaterium. 


of the patch. A comparison based on the pooled area of plants within 
three inner annuli to that in the outer annulus produced #24, = 6.72, 
P < 0.001). The distribution corresponds with a characteristic hostplant 
edge-effect (see Porter, 1992: 69). There was no bias for aspect in 
egg distribution around this patch (423, = 2.41, P > 0.3). 


Hostplant patches surveyed varied in area from 25 cm? to 2400 cm?, 
and comprise from one to 85 shoots. The nine variables measuring 
hostplant characteristics boil down into three factors using VARIMAX 
factor analysis, accounting jointly for some 81% of the total variance. 
The first factor, in particular, is dominated by the close relationship 
between the three patch size variables (39%), whereas the second and 
third factors focus on PLANT HEIGHT and LEAF SIZE, and LEAF 
TOUGHNESS respectively. PATCH SIZE, SHOOT NUMBER and 
LEAF NUMBER are very highly correlated (r, minimum, 0.88, 
P< 0.001). These three patch size variables are not significantly 


) 


correlated with plant height (r, maximum, 0.33, P = 0.06), leaf size 
(r, maximum, 0.32, P = 0.077) and leaf toughness (r, maximum, 0.14, 
P = 0.42) but covary significantly with exposure to sunlight (r,, 0.44 
to 0.63, P — 0.011 to 0.001). Patches with taller plants tend to have 
larger leaves (r,, 0.55, P < 0.001) and somewhat softer leaves (LEAF 
TOUGHNESS r,, -0.31, P= 0.08) but are not greatly affected by 
degree of EXPOSURE to sunlight as measured in the study area (r,, 
—0.09, P=0.63). Correlations between LEAF SIZE and LEAF 
TOUGHNESS (r, -0.17, P = 0.39), LEAF SIZE and EXPOSURE 
(rs, —0.26, P= 0.15) and between LEAF TOUGHNESS and EXPO- 
SURE (r,, 0.45, P = 0.009) are as expected in sign, but only the latter 
relationship is significant. Altogether, hostplant patches that are less 
shaded from direct sunlight are larger and have more shoots and leaves, 
but tend to be slightly shorter and have smaller and tougher leaves. 
In this study area, there is a weak (though not significant) tendency 
for larger plant patches to have more distant neighbours (r,, 0.26, 
P=0.15), but all hostplant patches surveyed have at least one 
neighbour each within 5m radius, and the correlation with DENSITY 
is weaker still (r,, -0.12, P = 0.49). There is also a tendency for hostplant 
patches with taller shoots (though not significantly with larger leaves) 
to be less isolated (PLANT HEIGHT with MEAN DISTANCE r, 
—0.38, P = 0.03 ; with DENSITY r, 0.49, P = 0.003 ; LEAF SIZE with 
MEAN DISTANCE r, 0.16, P =0.38; with DENSITY r,, -0.18, 
P = 0.32). 


The data collected indicate that Z. cretica lays more eggs on larger 
plant patches (PATCH SIZE r,, 0.69, P< 0.001) with more shoots 
(SHOOT NUMBER r,, 0.63, P < 0.001) and leaves (LEAF NUMBER 
r,, 0.65, P < 0.001). Plant patches with larger leaves also tend to receive 
more eggs (LEAF SIZE r,, 0.37, P = 0.033), but hostplant shoot height 
(PLANT HEIGHT r, 0.24, p = 0.19), hostplant exposure to sunlight 
(EXPOSURE rf, -0.19, P = 0.29) and hostplant leaf condition (LEAF 
TOUGHNESS r,, 0.06, P = 0.76) seem to have little influence on the 
number of eggs laid on hostplant patches. In part, this could be owing 
to the dominating influences of hostplant patch size. Egg numbers on 
hostplant patches are not greatly influenced by the number and distance 
of nearest neighbours (DENSITY r,, -0.21, P = 0.23; MEAN DIS- 
TANCE r,, 0.19, P = 0.29). The inference from the correlations is that 
more isolated patches receive more eggs, but this could be an artefact 
of larger patches tending to be more isolated in the survey area. 


Applying multiple regression to the set of transformed (normalised) 
variables, some 43% of the variation in egg load can be explained 


10 


by the two variables, SHOOT NUMBER (one of three highly 
correlated patch size measures) and LEAF SIZE, that correlate 
significantly with numbers of eggs on plant patches (Table 1). These 
two variables are effectively the key variables (with highest loadings) 
on the first two factors of a VARIMAX factor analysis of the nine 
measurements on hostplant condition (see earlier). It 1s possible that 
some of the residual variation in egg numbers on hostplant patches 
is accounted for by the length of time that hostplant patches have 
been accessible to Z. cretica females, particularly as the butterfly had 
been on the wing for at least a month prior to the survey. Those 
hostplant patches available or accessible for oviposition for longer 
periods are more likely to have more larvae. It was found that in a 
simple linear regression of numbers of eggs on hostplant patches against 
hostplant patch size (SHOOT NUMBER or PATCH SIZE), hostplant 
patches with an excess of eggs tended to have larvae compared to 
hostplant patches with a deficit of eggs (Fisher exact test, P = 0.022). 
The total number of eggs on hostplant patches also correlates signif- 
icantly with the percent of larvae on patches (r,, 0.595, P = 0.001) as 
does the number of unhatched eggs with the number of larvae on 
hostplant patches with oviposition records (r,, 0.635, P = 0.0007). In 
multivariate mode, the PERCENT OF LARVAE account for some 
11% of the residual variation from the regression of numbers of eggs 
on hostplant patches against patch size (SHOOT NUMBER) and 
LEAF SIZE. When all three independent variables are entered 
simultaneously, then some 49% of the variation in the numbers of eggs 
on hostplant patches can minimally be “explained” (Table 1). 


Table | 


Multiple regression parameters for the distribution of Zerynthia cretica 
on hostplant patches 


Dependent variable : Number of Z. cretica 


(i) Independent variables : NUMBER OF SHOOTS ; LEAF SIZE 


R = 0.68 ; R? = 0.46 ; Adj R? = 0.43 

F = 12.96 ; df 2,30 ; P = 0.00009 ; SE estimate = 0.50 
Number of shoots, beta = 0.56, P = 0.0003 

Leaf size, beta = 0.31, P = 0.027 


(ii) Independent variables : NUMBER OF SHOOTS ; LEAF SIZE ; 
PERCENT LARVAE 


R = 0.73 ; R? = 0.54 ; Adj R? = 0.49 

F = 11.17; df 3,29 ; P = 0.00005 ; SE estimate = 0.47 
Number of shoots, beta = 0.49, P = 0.0008 

Leaf size, beta = 0.27, P = 0.046 

Percent larvae, beta = 0.28, P = 0.041 


I] 


Eight of the plant patches received no eggs at all. When coded as 
present or absent on hostplant patches, Z. cretica correlates significantly 
with the same set of variables as does the number of Z. cretica eggs 
(1.e., SHOOT NUMBER, Gamma 0.51 ; PATCH SIZE, Gamma 0.54 ; 
LEAF NUMBER, Gamma 0.50; z= 2.49 to 2.52, P< 0.01 ; LEAF 
SIZE, Gamma 0.52, z = 2.58, P = 0.009), but also correlates signif- 
icantly with hostplant patch isolation measures (DENSITY, Gamma 
-0.48 ; MEAN DISTANCE, Gamma, 0.42, z > + 2.09, P < 0.04). The 
more apparent are hostplant patches and the fewer are neighbouring 
patches, the more likely it is that Z. cretica will lay eggs on those 
plant patches. Application of logistic regression (LOGIT model) 
indicates that some 29% of the variance in presence or absence of Z. 
cretica on hostplant patches can be accounted for by the size of 
hostplant patches (1.e., SHOOT NUMBER, LEAF SIZE) and isolation 
(1.e., MEAN DISTANCE) from neighbours. Whether plants are used 
as a resource or not may also depend on how long they have been 
available or accessible, but the percent of larvae on hostplant patches, 
which may indicate the length of time they have been available to 
females, accounts for little of the residual variance (r, 0.243, P = 0.17) 
and has little explanatory power (c. 6%). 


Discussion 


Observations from this brief survey indicate a significant loading on 
Aristolochia cretica hostplants by Z. cretica females, (i) on leaves; (ii) 
on shoots ; and (iii) on the edges of patches. Egg loading on hostplant 
patches has several components in Z. cretica: (i) plant patch char- 
acteristics of size, quality and spacing that influence selectivity ; (ü) 
indications of potential for variable clutch egg-laying by females that 
affords flexibility in their response to plant characteristics ; and (ii) 
the possibility of the deposition of multiple batches by different females 
on the same hostplant patch. 


Although there may be benefits to group living (e.g., aposematism ; 
Sillén-Tullberg & Leimar, 1988 ; Guilford, 1988) of the kind noted in 
Z. cretica (see Fig. 3), egg loading has at least two potential consequences 
for the population dynamics of a butterfly such as Z. cretica, both 
tending towards density-dependent ‘regulation’ of the upper bound to 
population size. First, excessive egg loading on hostplant patches could 
result in defoliation of the host and force mature larvae to search for 
new hosts to complete development. Such searching behaviour could 
lead to increased mortality among Z. cretica larvae. This has been 
observed to occur in Moroccan Zerynthia rumina (L.) (S. P. Courtney, 


12 


pers. comm.). Secondly, concentrations of eggs and larvae may enhance 
host-location by parasitoids (Godfray, 1994). 


Although large hostplant patches acquire significantly more eggs than 
small patches, this in itself does not lead to excessive loading of eggs 
on large patches compared to small patches. In fact, the correlation 
of hostplant patch egg :shoot ratio (Z. cretica eggs/shoot number) with 
patch size variables is low and negative (r, max, -0.148, P = 0.48) and, 
if anything, points to relatively more eggs being loaded on small 
hostplant patches. Some hostplant patches seem to have much higher 
egg :shoot ratios than others (range : 0.06 — 5.8), and larvae developing 
on these patches may be forced to search for complementary resources. 
The deterioration of hostplant quality (toughening of leaves) in the 
arid climate of southern Crete, as summer progresses, may also act 
as a further stimulus for larvae to seek out new resources. It is 
noteworthy, then, that Z. cretica females tend to avoid laying eggs 
on the smallest patches, but may be induced to do so when neighbouring 
plants are few or distant. The second consequence of egg loading on 
hostplant patches may well have more impact on Z. cretica located 
on large plant patches. This will depend on how parasitoids are cued 
into their Z. cretica hosts. The larger the hostplant patch, the more 
apparent will be both hostplant cues and host cues (direct and indirect) 
to parasitoids, especially as the numbers of Z. cretica increase much 
in proportion to patch size. There is the possibility that parasitism 
will be greater on large plant patches, since larger plants may be more 
likely (though not invariably) to have larvae at any time (r, min, 0.42, 
P = 0.014), and the presence of larvae could substantially enhance cues 
for parasitoids seeking early stadia (Fig. 3). 


These potential outcomes of female behaviour are worth more detailed 
study, especially as they invoke questions about negative feedbacks 
associated with population size and density. Additional survey work 
is needed over a wider area to test the limited observations made here. 
The present survey was carried out, of necessity, in a small relatively 
accessible area. Thus, the effects of plant patch isolation have not been 
fully accounted ; for instance, more isolated plants may be more heavily 
burdened with egg loads regardless of patch size. 


Although a substantial amount of the variance in the distribution of 
Z. cretica has been determined (49%), still much remains unaccounted. 
It is feasible that hostplant chemistry plays an important part in host 
choice ; much as Zerynthia polyxena (D & S), slow flying Zerynthia 
cretica is probably a chemically defended aposematic butterfly (Brower, 
1984). Other parameters (e.g. predation) may also be at work ; eggs 


13 


and larvae may already have been lost before or during the survey 
dates and such mortality may be uneven over plant patches. Some 
eggs may have been lost from rainwash ; a torrential storm occurred 
overnight midway through the survey. Also, without information on 
adult numbers, their distribution and movement, it is not possible to 
determine how egg distribution relates to population parameters or 
chance influences. Nevertheless, the present brief survey indicates that 
there is much to be gained from a more detailed examination of Z. 
cretica ecology and of the comparative biology of different Zerynthia 
species and populations which use a variety of hostplants in different 
terrains (De Freina, 1979 ; Hensle, 1993 ; Jordano & Gomariz, 1994). 
However, this work will need to incorporate data from directly tracing 
female behaviour. 


Acknowledgements 


My thanks to Lance Chilton and to Steven Courtney for information on 
Aristolochia cretica and Zerynthia rumina respectively, and to an anonymous 
referee for helpful comments on the manuscript. 


References 


Brower, L. P., 1984. Chemical defence in butterflies. Im Vane-Wright, R. 
I. & Ackery, P. R. (Eds): The Biology of Butterflies. Symposium of 
the Royal Entomological Society of London, Number 11. London: 
Academic Press. | 

DENNIS, R. L. H. (Ed.), 1992. The Ecology of Butterflies in Britain. Oxford : 
University Press. 

DE FREINA, J. J., 1979. Zur Kenntnis der Gattung Allancastria unter 
Berücksichtigung der A. cerisyi und A. deyrollei. Ent. Z. Essen 89 : 129- 
142. 

Goprray, H. C. J., 1994. Parasitoids. Behavioural and Evolutionary Ecology. 
Princeton : University Press. 

GUILFORD, T., 1988. The evolution of conspicuous coloration. Am. Nat. 131: 
S7-S21. 

HENSLE, J., 1993. Beobachtungen bei westanatolischen Osterluzeifaltern 
(Lepidoptera : Papilionidae). Nachr. ent. Ver. Apollo 14 : 289-299. 
JORDANO, D. & Gomariz, G., 1994. Variation in phenology and nutritional 
quality between host plants and its effect on larval performance in a 
specialist butterfly, Zerynthia rumina. Entomologia exp. appl. 71 : 271- 

DT 

Nig, N. H., Hurı, €. H., Jenkins, J. G., STEINBRENNER, K. & BENT, D. 
H., 1975. SPSS Statistical Package for the Social Sciences. New York : 
McGraw Hill Book Company. 


14 


OLIVIER, A., 1993. The Butterflies of the Greek island of Rodés : Taxonomy, 
Faunistics, Ecology and Phenology. Antwerpen : Vlaamse Vereniging 
voor Entomologie. 

Porter, K., 1992. Eggs and egglaying. Jn Dennis, R. L. H. (Ed.) : The Ecology 
of Butterflies in Britain. Oxford : University Press. 

SILLEN-TULLBERG, B. & LEIMAR, O., 1988. The evolution of gregariousness 
in distasteful insects as a defence against predators. Am. Nat. 132 : 723- 
734. 


15 


Nota lepid. 18 (1) : 16 ; 31.1.1996 ISSN 0342-7536 


Butterfly Conservation’ 2nd International Symposium 


Conserving Lepidoptera in a Changing Environment 


University of Warwick, UK 
6-8th September 1996 


Following the successful Ist Symposium at Keele University in 
September 1993, Butterfly Conservation is to hold its 2nd International 
Symposium in 1996. 


The Symposium will be organised as a series of thematic sessions 
covering a wide range of subjects involved in the conservation of 
Lepidoptera, with particular emphasis on the problems of environmental 
change, including both climate and land use. 


Some proposed topics are : 


The Future of Lepidoptera in Europe 
Semi-Natural Habitats 

Lepidoptera as Indicators of Habitat Quality 
Local Conservation Initiatives 

Lepidoptera and Climate Change 
Conservation and Landscape 

Management of Woodlands 

Restoration Ecology 

Conservation of Tropical Lepidoptera 


Although there will be a number of keynote speakers, the majority 
of the Symposium will be composed of submitted papers and posters. 
Details of session topics and instructions for submission of abstracts 
will be detailed in the second announcement. If you wish to receive 
further information (Butterfly Conservation members will receive this 
information automatically) please contact : 


Dr. Andrew S. Pullin, 
Department of Biological Sciences, 
Keele University, 

Staffs STS 5BG, UK 


Fax : 01782 630007 
e-mail : bia0S@keele.ac.uk. 


16 


Nota lepid. 18 (1) : 17-37 ; 31.1.1996 ISSN 0342-7536 


Subalpine and alpine assemblages of Lepidoptera 
in the surroundings of a powerful smelter 
on the Kola Peninsula, NW Russia 


Mikhail V. Kozrov 


Laboratory of Ecological Zoology, University of Turku, FIN-20500 Turku, Finland 


Summary 


Subalpine and alpine Lepidoptera were collected in 1993-1994 in the Monche 
and Tshuna Mts. (Kola Peninsula, NW Russia) in ten localities situated 4 to 
32 km SSW of a powerful smelter in Monchegorsk. No differences in species 
composition were discovered either between subalpine and alpine habitats or 
among localities. Ordination analysis revealed no clear environmental gradients, 
but mean ambient concentration of sulphur dioxide (estimated by manganese 
concentration in birch leaves) explained a significant part of variation in the 
abundance of Lepidoptera, whereas no relationships were found between abun- 
dance and foliar concentration of the main metal pollutant (nickel). The 
number of specimens collected during a fixed time (30 min) peaked in localities 
with moderate pollution loads, while high pollution cause populations to 
decline. However, the rarefaction-corrected species richness on three plots 
closest to the smelter was 1.3 times higher than on more distant plots, while 
diversity (measured by Shannon entropy) did not change. In contrast to the 
transect counts of butterflies, the method used in the present study is good 
for environmental assessment programmes even in the areas with low species 
richness and harsh climatic conditions. 


Résumé 


L’auteur a récolté des Lépidoptères subalpins et alpins en 1993-1994 dans 
les Monts Monche et Tshuna (Péninsule de Kola, Russie du Nord-Ouest) 
dans dix localités situées à 4-32 km au Sud-Ouest/Sud d’une puissante usine 
de fonderie à Monchegorsk. Il n’a pas trouvé de différences dans le cortège 
des espèces, soit entre biotopes alpins et subalpins, soit entre les localités. 
L'analyse n’a pas révélé de gradients environnementaux nets; mais la 
concentration ambiante moyenne de dioxyde de soufre (estimée d’après la 
concentration du manganèse dans les feuilles de bouleau) expliquait une partie 
significative de la variation quant à l’abondance des Lépidoptères. On n’a 
en revanche pas trouvé de rapports entre cette abondance et la concentration 
du principal métal polluant (nickel) dans les feuilles. Le nombre de spécimens 
récoltés durant un temps fixe (30 minutes) atteignit son maximum dans les 
localités modérément polluées, tandis qu’une forte pollution entraîne une 


It) 


diminution des populations. Cependant, la richesse en espèces, corrigée selon 
la raréfaction, sur trois terrains très proches de la fonderie était 1,3 fois plus 
élevée que sur les terrains plus éloignés, tandis que la diversité (mesurée par 
entropie de Shannon) n’a pas changé. Comparativement au comptage des 
papillons par transect, la méthode utilisée pour la présente recherche convient 
pour les programmes d'évaluation de l’environnement même dans les régions 
pauvres en espèces et/ou dont le climat est vraiment très rude. 


Introduction 


Subalpine and alpine habitats of the Kola Peninsula, NW Russia, form 
relatively small patches within the boreal forest zone. However, these 
mountain habitats display a distinctive fauna of moths and butterflies 
(Kozlov & Jalava, 1994). This fauna includes some rare and local 
species, which should be considered in conservation programmes in 
northern Europe. 


In contrast to boreal forest lepidopterans, the data on habitat asso- 
ciations and abundance of moths and butterflies in subalpine birch 
woodlands and alpine tundra in Fennoscandia are quite fragmentary. 
The community ecology of mountain Lepidoptera was studied by 
Fridolin (1935, 1936), and after his pioneer work, only little attention 
has been paid to this subject. The importance of ecological investigations 
in subalpine and alpine communities of the Kola Peninsula is empha- 
sised by the large-scale environmental deterioration caused by aerial 
emissions of the nickel smelter in Monchegorsk (Kozlov & Haukioja, 
1995). Recently, the north-eastern parts of the Monche Mts. have 
become surrounded by industrial barrenlands, but the extent of polluted 
territory in the mountains and the environmental effects of pollutants 
on subalpine and alpine ecosystems are poorly documented. Therefore 
the main objective of this investigation was to quantify the impact 
of aerial emissions on the abundance and species richness of the 
mountain Lepidoptera. 


Material and methods 
Study area and emission source 


The Kola Peninsula, situated in the north-west corner of Russia, 
belongs geographically to northern Fennoscandia. The highest peaks 
are in the centre of the Peninsula: Chibiny mountains (1190 m), 
Lovozerskie tundra (1120 m), Tshuna-tundra (1072 m) and Monche- 
tundra (965 m). These mountains are formed of alkaline phosphate 
minerals and have a very rich alpine flora (Ramenskaja, 1983). 


18 


The Severonikel smelter complex situated in Monchegorsk (68° N, 
33°E) produces aerial emissions consisting mostly of SO, and heavy 
metals (Ni, Cu, Co). The total amount of pollutants emitted in 1990 
was 2.64 X 108 kg, including 2.33 X 108 kg of SO, and 1.58 X 107 
kg of dusts containing heavy metals (2.7 X 106 kg of nickel, 1.8 X 
10° kg of copper) (Berlyand, 1991). This level of pollution was 
characteristic at least for the years 1986-1990, but during the past two 
years emissions of sulphur dioxide have been reduced (1.82 X 108 kg 
in 1992, 1.31 X 108 kg in 1993) (“Gipronikel” Institute, official data). 


Sampling procedure 


Ten pairs of 200 X 200 m size plots were established at the timber- 
line (formed by Betula pubescens tortuosa) on the north-eastern and 
eastern slopes of the Monche and Tshuna Mts., 4 to 32 km of the 
smelter (Fig. 1). Within each locality, the two plots representing birch 


(av) 
_ 
xe) 
S 
=) 
~ 
t 
qd 
(= 
=) 
Le 
(02) 
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Fig. 1. Study area. Localities (dots) are numbered ; the smelter is marked by an asterisk. 
Mountain relief is shown by contours of 400 and 800 m. Inserted : position of the 
study area within the Kola Peninsula. 


19 


woodlands and mountain tundra were situated 200 to 800 m apart 
(altitudinal difference 20 to 100 m). 


The plots were chosen with as similar vegetation as possible. In the 
subalpine woodlands, birch trees were sparse, 2 to 4 m tall, with an 
understorey formed by Betula nana and Juniperus sibirica and a surface 
cover dominated by dwarf shrubs (Empetrum nigrum, Vaccinium 
myrtillus) and lichens (Cladina, Cetraria). The tundra plots belong 
to Betula nana communities, with a predominance of E. nigrum in 
dwarf-shrub layer, below which are sparse mosses (Pleurozium schreberi, 
Pohlia nutans, Orthocaulis kunzeanus, Barbilophozia hatcheri) and 
usually a dense lichen cover (Cetraria nivalis, C. islandica, Cladina 
stellaris, Cladonia spp.). 


Samples were taken during the periods 12-23 July 1993 and 15-22 July 
1994 by collectors walking from Monchegorsk to the southernmost 
plot through the roadless mountains. It was planned to apply a sımilar 
sampling effort to all plots, but weather conditions and some technical 
problems interfered with the sampling design. Moreover, the field 
processing of samples taken in 1993 clearly demonstrated that the total 
number of specimens collected in the two localities closest to the smelter 
was too low for reasonable estimation of diversity characteristics. 
Therefore, additional sampling was arranged on these two plots. 


Sampling was conducted when the temperature was above 10°C, the 
wind did not exceed 3 on the Beaufort scale, and not earlier than 
2 hours after the last rain. The collector hunted for all moths and 
butterflies within the plot during 30 min. The sampled specimens were 
sorted according to species, and voucher specimens were preserved for 
determination when necessary. Time of sampling sessions was rando- 
mised among localities ; weather conditions during sessions (air tem- 
perature, wind speed, cloudiness) were recorded. In total, 1599 specimens 
representing 107 species were collected during 175 sessions (Table 1). 


The following variables were recorded at each plot (Table 1) : habitat 
(HAB: subalpine woodland or alpine tundra), altitude (ALT, to the 
nearest 10-20 m), slope aspect (Asp, to the nearest 45°), total plant 
cover (cov), tree cover (TRE), dwarf-shrub cover (SHR), number of 
vascular plant species (vPL), total species number for vascular plants, 
mosses and lichens (TPL). Within each of 20 plots, 14 subplots were 
randomly chosen to measure characteristics of the vegetation. TRE, 
VPL and TPL were accounted in four plots of 10 X 10 m size, whereas 
cov and sHR were measured in ten plots of 1 X 1 m size ; complete 
results of this work will be published elsewhere (Koroleva, pers. comm.). 


20 


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26 


To assess pollution load, at each site we sampled about 50 leaves from 
each of five trees of mountain birch, Betula pubescens tortuosa (dia- 
meter 40-80 mm, height 3-5 m). The trees were randomly chosen at 
the upper tree limit, approximately between the subalpine and alpine 
plots. Leaves were taken both in 1993 and 1994 at a height of 1.2- 
1.4 m and stored in paper bags. Unwashed leaf samples were dried 
for 12h at 80°C, then ground with a mill and analysed separately. 
Concentrations of nickel and manganese were determined by X-ray 
fluorescence (Spectrace 5000 spectrometer, Tracor X-ray, Holland). 
Nickel is the main metal pollutant of the smelter, whereas foliar 
concentration of manganese decreases with an increase of SO, concen- 
tration in the ambient air (Kozlov ef al., 1995). The locality-specific 
means based on two years of data are used in the analyses (variables 
NIC and MAN). 


The quality of the analytical data was checked by replicate analyses 
of the same samples, representing different contamination levels, both 
by blind tests with the same analytical procedure in the original 
laboratory, and by inductively-coupled plasma atomic emission spec- 
troscopy after wet digestion (HNO;-H,O,). The latter analyses were 
conducted in the Finnish Forest Research Institute, a laboratory 
belonging to the intercalibration network organised by the International 
Union of Forest Research Organisations and the European Community. 
In both cases, the results of the independent analyses showed a very 
high correlation (r = 0.935-0.994, n = 30, p < 0.0001). The mean 
ratios (based on 30 samples) between concentrations of metals estimated 
by X-ray fluorescence and by the atomic emission spectroscopy were : 
1.07 for Ni and 1.21 for Mn. Since we were interested in intra-plot 
differences more than in absolute values, the X-ray fluorescence was 
accurate enough to meet the goals of our study. 


Statistical analysis 


The number of specimens, number of species and Shannon diversity 
index (H) were analysed in respect to sampling conditions by the SAS 
GLM procedure ; Type III Sum of Squares was used to assess the 
significance of effects. Distributions of the numbers of specimens and 
species, as well as of their log-transformed values, significantly deviated 
from the normal distribution ; therefore these characteristics were 
square-root transformed before analyses. Since sampling design was 
unbalanced in respect of time, collector, and weather conditions, least- 
Squares means (LSMs) were computed for each locality. LSMs are 
simply estimators of the plot marginal means that would be expected 
if the design had been balanced (SAS Institute, 1990). 


27 


The relationships between lepidopteran assemblages and the environ- 
mental variables listed above were analysed by canonical correspondence 
analysis (CCA ; CANOCO statistical program) (Ter Braak, 1987). The 
pooled list of species recorded at each study plot (Appendix) was used 
to estimate rarefaction corrected values for species richness (Krebs, 
1989) and to calculate the plot-specific values of the Shannon diversity 
index (H). Comparisons between plots or plot groups were conducted 
by analysis of variance (SAS ANOVA procedure). Correlation coef- 
ficients were calculated by SAS CORR procedure (SAS Institute, 1990). 


Results 


Subalpine and alpine lepidopteran assemblages differ in respect of their 
sensitivity to sampling conditions. The numbers of both specimens and 
species in alpine samples were mostly influenced by air temperature ; 
sampling time, wind speed and cloudiness also showed significant effect 
on abundance. In contrast, numbers of specimens in subalpine samples 
depended mostly on the collector (Table 3). 


CCA of samples from 10 alpine and 10 subalpine plots explained only 
30.2% of variation and revealed no clear pattern in plot ordination. 
The Monte Carlo permutation test showed that even the first axis was 
not significant (99 random permutations, F = 1.12, P < 0.22). Thus, 
CCA analysis revealed no differences in species composition between 


Table 3 


Effects of sampling conditions on basic characteristics of individual 30-min samples. 


Habitat Source | F values for dependent variables (2) 
of variation df Specimens Species Diversity 


2221 
0.60 
0.00 
3:89 
3.88 
2 


SOUS 
14.10*** 
20:85 

I SD FE 

0.00 


(2) Numbers of species and specimens were square-root transformed. Significance levels : 
**# _ P = 0.001, ** — P=0.01, * — P=0.05. 


Subalpine | Plot 
Time 
Temperature 
Wind speed 
Cloudiness 
Collector 


Plot 

Time 
Temperature 
Wind speed 
Cloudiness 
Collector 


28 


subalpine and alpine lepidopteran assemblages, and therefore the data 
collected from subalpine and alpine plots in the same locality were 
pooled for further analysis. 


Abundance of Lepidoptera did not depend on foliar nickel concen- 
trations (Fig. 2), but demonstrated a clear dome-shaped pattern in 
relation to foliar manganese (Fig. 3). Three groups of plots were 
distinguished on the basis of both Mn concentration in birch leaves 
and the abundance of Lepidoptera: slightly polluted plots with low 
abundance (localities 7, 8 and 9), moderately polluted plots with high 
abundance (localities 5, 6 and 10), and heavily polluted plots with low 
abundance (localities 1-4). Within each group, intra-plot variation in 
abundance was low=(s5 4) — Oly P< 016058 Fo 4) = I-11, P 
< 0.3403 ; and Fa, 7, = 1.67, P < 0.1811, respectively), whereas intra- 
group variation was highly significant Fo, 172) = 31.03, P < 0.0001). 
The decrease in density close to the smelter was more pronounced 
in tundra (37% of the abundance in moderately polluted plots) than 
. in woodlands (51%). 


Among other environmental variables, abundance correlated only with 
the total number of plant species recorded in study plots (Fig. 4). Since 
numbers of specimens per 30-min sample were low, both the numbers 
of species and the diversity index (H) calculated for individual samples 
simply follow changes in abundance (r = 0.96, n = 175, P < 0.0001, 
and r = 0.92, n= 175, P < 0.0001). 


The rarefaction-corrected locality-specific values of species richness as 
well as the diversity index (H) calculated for the pooled samples 
correlated neither with foliar manganese nor nickel concentrations. 
However, rarefaction-corrected species richness on the three plots 
closest to the smelter was 1.3 times higher than on more distant plots 
(Fig. 5), while the diversity index did not change with the pollution 
load (Fig. 6). Correlation between diversity and the number of plant 
species (TPL) was not significant. 


- Discussion 


Suitability of the sampling procedure 


In past years, transect counts have been used for collecting quantitative 
information on day-active Lepidoptera. This method, originally deve- 
loped for butterflies (Pollard, 1977), was later applied to all macro- 
lepidopterans which can be distinguished in the field by the experienced 
observer (Väisänen, 1992). The drawbacks of transect counts are: 


29 


Specimens / 30 min sample 


10 20 30 40 50 60 


Specimens / 30 min sample 
oa oo 


100 200 300 400 500 600 700 800 900 
Concentration of manganese in birch leaves, ug/mg 


18 
16 
14 
12 
10 


Specimens / 30 min sample 


12 14 16 18 20 22 24 26 28 30 
Total number of plant species in study plot 


Figs 2-4. Locality-specific corrected means (LSMs) and standard errors of total 
abundance of Lepidoptera (specimens collected during 30 min) in relation to 
environmental variables: 2 — foliar nickel concentration; 3 — foliar manganese 
concentration (weighted lowest approximation shown) ; 4 — total number of plants 
in study plots (regression is significant at p = 0.01). 


30 


= x : : ; 

= 20 ous si 

= 15 e 

g 10 

zs 5 


y 100 200 300 400 500 600 700 800 900 
Concentration of manganese in birch leaves, 1g/mg 


Diversity index (H) 


100 200 300 400 500 600 700 800 900 
Concentration of manganese in birch leaves, ug/mg 


Figs 5-6. Locality-specific diversity (means and standard errors) calculated for the 
pooled samples plotted against foliar manganese concentration: 5 — rarefaction- 
corrected species richness (number of species expected in the sample of 50 individuals) ; 
6 — Shannon diversity index (H). 


(1) impossibility of counting ‘microlepidoptera’, (2) the importance of 
an observer’s experience in species recognition, (3) the possible over- 
estimation of population densities by repeated counts of the same 
individuals and (4) the impossibility of confirming the species observed. 
The sampling method used in the present study lacked these short- 
comings. However, the numbers of several moths may be underestimated 
due to their fast long-distant flight and camouflage ; this conclusion 
especially relates to the noctuids of the genus Sympistis. 


Both the number of species and diversity index (H) calculated for the 
individual samples did not depend on the collector (Table 3) indicating 
that personal experience was not of critical importance for obtaining 


31 


a representative sample. However, the number of collected specimens 
depended on the collector, but only in subalpine habitats (Table 3). 
This conclusion coincides with the general impression from collecting 
in the mountains : in alpine tundra, the number of sampled Lepidoptera 
almost completely depends on the frequency of their spontaneous 
appearance, while in more protected habitats of birch woodlands one 
can increase the catch by active searching and by disturbing moths 
from the vegetation. But since the samples were not selective in respect 
to the lepidopteran species, and the same collectors sampled in all plots, 
problems were not encountered when analysing the pooled data. 


Density pattern along pollution gradient 


Herbivorous insects usually respond to moderate pollution loads by 
an increase in population density, while high concentrations of pollutants 
have an adverse effect (Riemer & Whittaker, 1989; Kozlov, 1990). 
Numerous insect groups such as Eriocraniidae (Kozlov & Haukioja, 
1993), Tortricidae (Kozlov, unpubl. data), Noctuidae (Kozlov, Jalava 
et al., in press), chloropid flies (Zvereva, 1993) and the leaf beetle, 
Melasoma lapponica L. (Zvereva et al., 1995) show the dome-shaped 
density patterns around the Severonikel smelter, when the lowland 
habitats are considered. These changes in densities are mostly related 
to sulphur dioxide, not heavy metals (Riemer & Whittaker, 1989 ; 
Zvereva et al., 1995), although correlations with metal pollutants may 
also exist. 


Background concentration of nickel in birch leaves in the Monchegorsk 
region is 16 + 1 ug/mg, whereas the maximum site-specific value 
observed in the industrial barrenland near the Severonikel smelter was 
586 ug/mg. In the lowlands, foliar Ni concentrations of 20 to 60 ug/ 
mg were recorded in plots situated 20 to 50 km from the smelter (Kozlov 
et al., 1995), where abundance of Lepidoptera was slightly higher than 
in the unpolluted locality (Kozlov, unpubl. data). Thus, the decrease 
in total abundance of Lepidoptera in the four montane plots proximate 
to the smelter is hardly related to metal pollutants. 


In unpolluted regions of the Kola Peninsula, the mean concentration 
of manganese in birch leaves is 1032 + 62 ug/mg (Kozlov et al., 1995). 
Decrease of foliar manganese resulting from both the direct impact 
of sulphur dioxide on plant cells and the decreased availability of Mn 
due to increased pH of soils has repeatedly been observed (Hutchinson 
& Whitby, 1974; Lobersli & Steinnes, 1988), and therefore the Mn 
concentration in birch leaves can satisfactorily describe the impact of 
sulphur dioxide (Fig. 7). 


32 


oe 
1 
H 


I 1 1 


= | 
100 200 BO) 700, S00 1110052211300 
Concentration of manganese in birch leaves, ug/mg 


SO, concentration in ambient air, ug/m® 


Fig. 7. Locality-specific ambient concentrations of sulphur dioxide in forest plots 
around Severonikel smelter (after Baklanov & Rodjushkina, 1993; Barkan, 1993 ; 
Kryuchkov, 1993) plotted against the corresponding concentrations of manganese in 
birch leaves (after Kozlov et al., 1995). 


The minimum site-specific Mn concentration observed in mountains 
(locality 4 : 120 ug/mg) was the same as recorded in industrial barren- 
lands near the Severonikel smelter (130 ug/mg ; Kozlov et al., 1995) ; 
the maximum observed in mountains (848 ug/mg) was still lower than 
the background value estimated in the lowlands. However, the direct 
comparison of foliar manganese concentrations measured in forests and 
in subalpine woodlands might well be inappropriate, just because 
different soil properties affect the mobility of Mn?* (Lobersh & 
Steinnes, 1988). However, the changes in abundance of Lepidoptera 
both in lowland forests (Kozlov, 1994 and unpubl. data) and in 
subalpine habitats follow a similar dome-shaped pattern, with maximum 
density attained at plots with estimated mean ambient SO, concen- 
trations around 100 ug/m?. 


The important difference between forest and mountain habitats concerns 
the effects of the smelter on vegetation, which has resulted in forest 
decline within distances of 5 to 15 km (Kozlov & Haukioja, 1995) 
but has caused almost no structural changes in subalpine and alpine 
plant communities (Koroleva, unpubl. data). Therefore the changes in 
lepidopteran assemblages in subalpine and alpine habitats can be more 
closely related to the pollution impact on insects and/or their host 
plants, rather than to pollution-induced habitat deterioration. 


35 


Environmental assessment using data on Lepidoptera 


Among Lepidoptera, only butterflies and skippers were generally 
considered as ecological indicators (Gilbert, 1984 ; Murphy er al., 1990). 
Monitored over large areas during decades (Pollard, 1991), they 
contribute much to conservation biology in both a population and 
community context (Ehrlich & Murphy, 1987). Butterfly assemblages 
were found to be excellent indicators of heterogeneity due to topo- 
graphic/ moisture gradients, limited indicators of heterogeneity due to 
anthropogenic disturbance, and poor indicators of plant diversity 
(Kremen, 1992). However, in some areas such as northern boreal 
forests, subalpine woodlands and alpine tundra, both the diversity and 
abundance of butterflies are too low to use them in environmental 
assessment programs. 


The present study has demonstrated that assemblages of Lepidoptera 
can be used for environmental assessment even in the areas with low 
species richness and relatively harsh climatic conditions. Consistent with 
the results of Kremen (1992), our data show that the pollution-induced 
disturbances make a low although significant contribution to the total 
variation in abundance of Lepidoptera. The number of vascular plant 
species on the plot (variable vpL) have no explanatory value, but the 
addition of moss and lichen species (variable TPL) improves the model 
suggesting that the richness of mosses and lichens correlates with some 
of the environmental variables not accounted in the present study 
(moisture conditions for example). 


Although there is a general consensus that polluted habitats display 
a reduction of diversity (Magurran, 1988), the Shannon diversity index 
based on samples of Lepidoptera showed no decrease even in the two 
localities closest to a powerful smelter. The increase of rarefaction- 
corrected species richness on plots closest to the smelter is probably 
explained by an increase of vegetation heterogeneity caused by pollution. 
The typical arcto-alpine species (Olethreutes aquilonanus, O. noricanus, 
Oeneis bore) as well as species endangered in Finland (Clossiana freija, 
Sympistis zetterstedtii) are recorded even in the two plots closest to 
the smelter, confirming that the existing levels of pollution had caused 
only a decrease in density, not decline in species. 


Acknowledgements 


The field work in the Lapland Biosphere reserve became possible due to kind 
permission of V. Zolotukhin and A. Paraketsov. I am extremely grateful to 
E. Melnikov, V. Zverev and A. Lvovsky who collected most of the data in 


34 


harsh environmental conditions of roadless mountains. N. Koroleva provided 
me with unpublished data on vegetation at the study plots. J. Jalava and 
L. Kaila helped me with moth determination. E. Haukioja, K. Mikkola and 
E. Zvereva made invaluable comments to an earlier draft of the manuscript. 
Suggestions by two anonymous referees prompted me to reconsider the data, 
and contributed much to the revised version of the paper. Special thanks 
to A. Bachtiarov who conducted heavy metal analyses. The work was 
supported by the Academy of Finland through grants to E. Haukioja and 
K. Mikkola. 


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Khibines au point de vue biocenotique. Ent. Obozr. 26 : 165-182. 

FRIDOLIN, V. Ju., 1936 Community of animals and plants of Chibiny mountain 
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& Leningrad. 295 pp. [in Russian]. 

GILBERT, L. E., 1984. The biology of butterfly communities. Jn Vane-Wright, 
R.I. & Ackery, P.R. (Eds): The biology of butterflies, pp. 41-54. 
Princeton Univ. Press, Princeton, New Jersey. 

HUTCHINSON, T. C. & Wuirsy, L. M., 1974. Heavy metal pollution in the 
Sudbury mining and smelting region of Canada. I. Soil and vegetation 
contamination by nickel, copper and other metals. Environ. Conserv. 
1 : 123-132. 

Kozıov, M. V., 1990. Impact of anthropogene factors on the populations of 
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mation, Moscow. 189 pp. [in Russian]. 

Koziov, M. V., 1994. Structure of lepidopteran communities in an air 
pollution gradient: do emissions affect biodiversity ? 5th European 
Congress of Entomology, 29 August - 2 September 1994, University of 
York, UK. Abstracts p. 152. 


35 


Kozıov, M. V. & HAukioJa, E. 1993. Pollution-induced changes in populations 
of Eriocraniidae (Lepidoptera) in surroundings of Monchegorsk. In 
Kozlov, M. V., Haukioja, E. & Yarmishko, V. T. (Eds) : Aerial pollution 
in Kola Peninsula: Proc. Intern. Workshop, April 14-16 1992, St. 
Petersburg. Apatity, p. 369. 

Kozıov, M. V. & Hauxiosa, E., 1995. Pollution-related environmental 
gradients around the “Severonikel” smelter complex at the Kola Penin- 
sula, Northwestern Russia. Jn Munawar, M. & Luotola, M. (Eds) : The 
contaminants in the nordic ecosystem : the dynamics, processes and fate. 
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lands, pp. 59-69. 

Kozıov, M. V., HAuxkiosa, E., BAKHTIAROV, A. V. & STROGANOV, D.N., 
1995. Heavy metals in birch leaves arround a nickel-copper smelter at 
Monchegorsk, Northwestern Russia. Environmental Pollution 90 : 291- 
299. 

Kozıov, M. V. & JALAVA, J., 1994. Lepidoptera of the Kola Peninsula, north- 
western Russia. Entomologica Fenn. 5 : 65-85. 

Kozıov, M. V., JALAVA, J., Lvovsxy, A. L. & MikkorA, K., in press. Popu- 
lation densities and diversity of Noctuidae (Lepidoptera) along an air 
pollution gradient in Kola Peninsula, Russia. Entomologica Fenn. 

Kress, C. J., 1989. Ecological methodology. Harper & Row, New York. 654 


Pp. 

KREMEN, C., 1992. Assessing the indicator properties of species assemblages 
for natural areas monitoring. Ecol. Applications 2 : 203-217. 

KRYUCHKOV, V. V., 1993. Degradation of ecosystems around the “Severonikel” 
smelter complex. In Kozlov, M. V., Haukioja, E. & Yarmishko, V. T. 
(Eds): Aerial pollution in Kola Peninsula: Proc. Intern. Workshop, 
April 14-16 1992, St. Petersburg. Apatity, pp. 35-46. 

LoBERSLI, E. M. & STEINNES, E., 1988. Metal uptake in plants from a birch 
forest area near a copper smelter in Norway. Water air soil pollut. 37 : 
25-39. 

MAGURRAN, A. E., 1988. Ecological diversity and its measurement. Chapman 
& Hall, London. 179 pp. 

MurPHY, D. D., Freas, K. E. & Weiss, S. B., 1990. An environment- 
metapopulation approach to population viability analysis for a threatened 
invertebrate. Conserv. Biol. 4 : 41-51. 

POLLARD, E., 1977. A method for assessing changes in the abundance of 
butterflies. Biol. Conserv. 12 : 115-134. 

POLLARD, E., 1991. Monitoring butterfly numbers. /n Goldsmith, B. (Ed.) : 
Monitoring for conservation and ecology, pp. 87-111. Chapman & Hall, 
London. 

RAMENSKAJA, M.L., 1983. The analysis of the vegetation of the Murmansk 
region and Karelia. Nauka, Leningrad, 215 pp. [in Russian]. 

Riemer, J. & WHITTAKER, J. B., 1989. Air pollution and insect herbivores : 
observed interactions and possible mechanisms. Jn Bernays, E. A. (Ed.), 
Insect-plant interactions, Vol. 1, pp. 73-105. CRC-press, Florida. 


36 


SAS Institute, 1990. SAS/Stat. User’s guide, version 6.0. SAS Institute, Cary, 
N.C. 

TER Braak, C. J. F., 1987. The analysis of vegetation-environment relation- 
ships by canonical correspondence analysis. Vegetatio 69 : 69-77. 

VAISANEN, R., 1992. Distribution and abundance of diurnal Lepidoptera on 
a raised bog in southern Finland. Annis zool. fenn. 29 : 75-92. 

ZVEREVA, E. L., 1993. Effects of industrial pollution on fly communities 
(Diptera Brachycera). Ent. Obozr. 72 : 558-569 [in Russian ; English 
translation : Ent. Rev. Wash. ]. 

ZVEREVA, E. L., Kozıov, M. V. & NEUVONEN, S., 1995. Population density 
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in surroundings of smelter complex. Environ. Ent. 24 : 707-715. 


37 


Nota lepid. 18 (1) : 38 ; 31.1.1996 ISSN 0342-7536 


Book reviews — Buchbesprechungen — Analyses 


Die Zünslerfalter (Pyraloidea) Mitteleuropas. Bestimmen - Verbreitung 
- Fluggebiet - Lebensweise der Raupen. F. SLAMKA. 112 pp., 12 arb- 
afeln. 1995. Bestellungen an: F. Slamka, Racianska 61, SK-83102 
Bratislava, Slowakei. Preis : 30,- DM zzgl. Porto. 


Zünslerfalter (Pyralidae und Crambidae) zählen zu den wenigen Schmetter- 
lingsfamilien, die praktisch allen Lepidopterologen, also auch den ausschließlich 
auf Großschmetterlinge konzentrierten, durchaus geläufig sind. Dies liegt einer- 
seits an vielfach auffallend gefärbten oder gezeichneten Tieren mit manchmal 
für Kleinsehmetterlinge erstaunlichen Größen, andererseits aber auch in der 
relativ leichten Registrierbarkeit zahlreicher Arten, sowohl tagsüber als auch 
am Licht. 


Während für Großbritannien oder Skandinavien seit den 80er Jahren bereits 
gut ausgestattete feldführerartige Bestimmungsliteratur existiert, machte sich 
in Mitteleuropa ein eklatanter diesbezüglicher Mangel bemerkbar. Der Autor 
des nunmehr vorliegenden Bestimmungsbuches hat die große Aufgabe über- 
nommen erstmals ein Werk zu schaffen, das mit relativ geringem Aufwand 
eine Determination der zentraleuropäischen Arten ermöglicht. Mitteleuropa 
wird dabei unter Ausschluß der Schweiz oder auch Norditaliens definiert, 
allerdings werden folgende Länder berücksiehtigt : Deutschland, Österreich, 
Polen, Tschechische Republik, Slowakei und Ungarn. Auch Vorkommen in 
Rumänien und der Westukraine finden Berücksichtigung. Das Buch ermöglicht 
somit eine zuverlässige Bestimmung fast aller Arten dieser Staaten. Rezensent 
konnte nur wenige Taxa finden, die nicht behandelt wurden wie z.B. Dioryc- 
tria simplicella, Spoladea recurvalis oder eingeschleppte Arten wie Vitula 
edmandsii und Agassiziella bilinealis. 


Alle behandelten 377 Arten werden farbig, in ansprechender Qualität, auf 
insgesamt 12 Tafeln abgebildet. Zusätzlich werden für sämtliche Taxa Geni- 
talskizzen beider Geschlechter sowie Strichzeichnungen wichtiger habitueller 
Unterscheidungsmerkmale gegeben. Die Textinformationen selbst sind - dem 
Zweck des Buches als Feldführer entsprechend - eher knapp gehalten und 
umfassen u.a. wichtigste Synonyma, Informationen zur Verbreitung, biologische 
und phänologische Angaben zu jeder Art sowie ein abschließendes Litera- 
turverzeichnis. 


Mit diesem empfehlenswerten Buch wird der Einstieg in die interessanten 
Zünsler, ohne die extremen Anfangsschwierigkeiten bei vielen anderen Klein- 
schmetterlingsfamilien, nunmehr sehr leicht gemacht und man kann mit 
Interesse weitere Arbeiten des Autors, wie z.B. ein geplantes Werk über die 
Großschmetterlinge Mitteleuropas, entgegensehen. Der Preis des Buches ist 
extrem günstig und die Arbeit wird somit zu einem Muß für alle an 


Lepidopteren Interessierte. 
I Peter HUEMER 


38 


Nota lepid. 18 (1) : 39-56 ; 31.1.1996 ISSN 0342-7536 


Identity and distribution of two dimorphic oriental 
fairy moths — Nemophora decisella (Walker, 1863) 
and Nemophora cantharites (Meyrick, 1928) 
(Lepidoptera, Adelidae) 


Mikhail V. Kozıov* & Gaden S. RoBınson** 


* Laboratory of Ecological Zoology, University of Turku, FIN-20500 Turku, Finland 
** Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 
5BD, UK 


Summary 


Sexual dimorphism in wing pattern has, historically, caused confusion in the 
association of males and females of both Nemophora decisella (Walker) and 
N. cantharites (Meyrick). The wide distribution of N. decisella, from the 
Russian Primorye to Java, combined with geographical variation in its external 
characteristics, has provoked repeated descriptions of this species for which 
eight junior synonyms are established here. One new synonym is established 
of N. cantharites. Data on all primary types are provided, and the species 
are redescribed. Variation with latitude in eye size, wing length, and the width 
and position of the forewing medial band is discussed for N. decisella. 


Resume 


Le dimorphisme sexuel révélé par le dessin des ailes est à l’origine historique 
de la confusion qui règne dans le groupe des mâles et des femelles de 
Nemophora decisella (Walker) et de N. cantharites (Meyrick). La distribution 
géographique très étendue de N. decisella, soit de Primorye (Russie) à Java, 
s’ajoutant a la variation géographique des caractères extérieurs de ce taxon, 
en a déclenché des descriptions répétées, contraignant les auteurs à créer huit 
junior-synonymes dans le présent travail. Pour N. cantharites ils établissent 
aussi un nouveau synonyme. Ils fournissent des renseignements sur les types 
primaires et redécrivent les deux espèces. Pour N. decisella, ils traitent des 
variations dans les caractères suivants : dimension de l’œil (largeur), longueur 
des ailes, largeur et position de la bande médiane de l’aile antérieure. 


Introduction 


Adelidae are small to medium-sized, colourful, frequently diurnal moths 
with conspicuously elongated antennae. The genus Nemophora Hoff- 
mannsegg, 1798 (type species : Phalaena (Tinea) degeerella Linnaeus, 


39 


1758, by subsequent designation of Hampson, 1918, Novit. Zool. 25 : 
388 ; for the authorship of the generic name, see Nye & Fletcher, 1991) 
is one of the most taxonomically complicated groups of Adelidae, and 
even the identity of the western Palaearctic species is sometimes 
confused. The eastern Palaearctic and Oriental species of this genus 
have been neither reviewed nor even listed since Meyrick’s (1912) 
catalogue in Genera Insectorum. However, the Japanese species have 
been considered by Okano (1959) and Moriuti (1983), and the fauna 
of the Malay archipelago and the Philippines has been revised by 
Diakonoff (1951, [1968}). 


In the course of taxonomic revision of the genus Nemophora we 
recognized that females of two oriental species, N. decisella (Walker) 
and N. cantharites (Meyrick), differed in wing pattern from conspecific 
males. This type of sexual dimorphism has not previously been recorded 
among Nemophora species ; conspecific males and females, even if 
collected simultaneously, were described as different species. This paper 
aims to clarify the taxonomy and distribution of these two closely 
related species. 


Materials and methods 


In the course of this study all primary types were examined. Following 
ICZN (recommendations 72E, 72F), all original labels of the specimens 
designated as lectotypes / paralectotypes, as well as old holotypes, are 
cited in full. Label size is given in millimeters ; the original text of 
the labels of type specimens is provided within quotation marks ; recent 
locality names are added in square brackets ; an oblique line (/) is 
used to separate parts of labels written on different lines. The spelling 
of locality names, except for the primary types, follows The Times 
Atlas of the World (Comprehensive Edn.), 1968. Localities in Japan 
were attributed to prefectures following Utech (1980). Altitudes are 
given in meters above sea level to an accuracy of 50 m. 


Male genitalia were investigated as described by Kozlov (1993) ; female 
genitalia were not considered here. The interocular index was measured 
following the formula of Davis (1975), being the ratio between the 
vertical diameter of the compound eye and the interocular distance 
measured at a point on the frons midway between the base of the 
antennal sockets and the anterior tentorial pits. Other measurements 
were made as shown in Fig. 1. 


40 


Abbreviations of museums and institutions : 


BMNH The Natural History Museum, London, U.K. 

EIHU Entomological Institute, Faculty of Agriculture, Hokkaido 
University, Sapporo, Japan. 

ELUO Entomological Laboratory, University of Osaka Prefecture, 


Osaka, Japan 
MINGA Muzeul de Istoria Naturala ‘Grigore Antipa’, Bucharest, Ru- 


mania 
MZH Zoological Museum in Helsinki, Finland 
NML National Museum of Natural History, Leiden, The Netherlands 
NHMW Natural History Museum, Wien, Austria 
USNM U.S. National Museum of Natural History, Smithsonian Ins- 
titution, Washington, D.C., U.S.A. 
ZIN Zoological Institute, St. Petersburg, Russia 
ZMB Zoological Museum, Humboldt University, Berlin, Germany 
ZMUC Zoological Museum, University of Copenhagen, Denmark. 


Fig. 1. Measurements of head and forewing : id — interocular distance ; | — forewing 
length ; md — minimum distance between eyes ; pf — position of outer border of 
medial yellow band ; vd — vertical eye diameter ; wf — width of medial yellow band. 


4] 


Phylogenetic relationships 


There is no subgeneric classification of Nemophora, but four species 
groups, “A”-“D”, were recognized by Meyrick (1912). However, even 
this classification appears doubtful, and different synonyms of N. 
decisella were included by Meyrick in different species groups. 


Our suggestion of a sister-group relationship between the two species 
discussed here is based on the only synapomorphy (sexual dimorphism 
in the coloration of the forewing base) that we can recognize. No other 
Nemophora species except N. heteroxantha Diakonoff, as we can 
conclude from examination of primary descriptions and the BMNH 
collection, share this apomorphy with N. decisella and N. cantharites. 
However, validity of N. heteroxantha described from E. Java is proble- 
matic (see below), and we do not consider this species here. The position 
of N. decisella and N. cantharites within the genus Nemophora cannot 
be refined further at present as some two-third of the included species 
are poorly known. 


Nemophora decisella (Walker, 1863) (Figs 2-5, 8, 9, 12-17) 


Nemotois decisella Walker, 1863, List Lepid. Het. Brit. Mus. 28 : 505. Holotype 
@: Indonesia, Sumatra [given erroneously in the original description 
as “China” — by a curious error the English descriptions and localities 
of this species and N. sinicella were switched in Walker’s publication ; 
the Latin disgnoses are correct (Meyrick, 1912)] ; labelled : circle diam 
8 with red border, print “Type”; blue-grey circle diam 6, black ink 
“Sumat/ra”, reverse side “54/76” ; 14 X 24, wide black border, black 
ink + print “Nemotois / deciscella. [Sic!] Wkr. / Cat. Lep. BM. XXVIII 
p. 505. 1863 / TYPE 3 descr.” ; 5 X 35, print “22. Nemotois decisella.” 
(BMNH) [examined]. 

Nemotois decisella Walker : Meyrick, 1912 : 7 (Sumatra). 

Nemophora decisella (Walker) : Diakonoff, 1951 : 173-174 (W Java) ; Robinson 
et al., 1994: 23 (Burma, Thailand, W Malaysia, Sumatra, Java, 
Anambas Is., Brunei, Kalimantan), fig. 33 (male head), plate 3 fig. 2 
(male, colour illustration). 

Ucetia bifasciella Walker, 1866, List Lepid. Het. Brit. Mus. 35 : 1821. Holotype 
© [erroneously cited as male in original description]: Indonesia, Java 
[type locality confirmed from BMNH accession register] ; labelled : circle 
diam 8 with red border, print “Type”; 5 X 7, print “60.15 / E[ast]. 
I[ndian]. C[ompany].” ; 6 X 8, black ink “772” ; 14 X 24, wide black 
border, black ink +print “Ucetia / bifasciella / Wkr. / Cat. Lep. Het. 
BM. 35. p. 1815. [Sie!]; 1866 / TYPE & .descer. 1) 0s) 5, 20028 some 
“Ucetia bifasciella” ; 4.5 X 14, print “a. Java” (BMNH) [examined]. 

Nemotois bifasciella (Walker) : Meyrick, 1912 : 7 (Java). 


42 


Nemophora bifasciella (Walker) : Diakonoff, 1951 : 179 (Java). 

Nemophora bifasciella irrorata Diakonoff, 1951, Treubia 21 : 179-181. Holotype 
© : Indonesia, E Java, Tengger Mts ; labelled : 4 X 7, print +black ink 
Gen Noe 28772 5.53% Sprint black ink “East Java, 2000 m 
/ Kletak / at light [deleted by ink] 19.V.1941 / A.M.R. Wegner leg.” ; 
15 X 16, thin black frame, print + black ink “TYPE: © / Nemophora 
| bifasciella / WALK / subsp. / irrorata / A.Diakonoff 1950”; red 
13 X 17, print + black ink “Museum Leiden / Holotype © / Nemophora 
| bifasciella / irrorata / Diakonoff, 1951 : 179” (NML) [examined]. 
Paratypes : 1 ©, labelled : 5.5 X 15, print + black ink “East Java, 2000 
m / Kletak / at light [deleted by ink] 6.V.1940 / A.M.R.Wegner leg.” ; 
15 X 16, thin black frame, print + black ink “PARATYPE: © / 
Nemophora / bifasciella / irrorata / A.Diakonoff 1950” (NML) 
[examined]. | © : same as previous, except the collecting date (10. VI.1941) 
(NML) [not examined]. Syn. n. 

Nemotois paradisea Butler, 1881, Trans. ent. Soc. London: 592. Holotype 
@: Japan, Honshu; labelled: circle diam 8 with red border, print 

~ “Type” ; 5.5 X 10, black ink “Tokei 80-97” [according to the BMNH 
accession register, this material was collected by Fenton], reverse side 
“Nemotois / paradiseus / Butler. Type” ; 14 X 24, wide black border, 
black ink + print “Nemotois / paradisea / Btlr. / Tr. Ent. Soc. Lond. 
592 (1881) / TYPE £ descr.” (BMNH) [examined]. Syn. n. 

Nemotois paradisea Butler : Meyrick, 1912 : 7 (Japan). 

Adela imperialis Rebel, 1900, Iris 13 : 187-188. Lectotype & (here designated) : 
Russia, “Amur” ; labelled : 5 X 7, print “630” ; 11 X 23, black ink “Adela 
Type / imperialis Rbl” ; 9 X 15, print + black ink “Stgr. 900 / Amur” ; 
13 X 18, print black ink “Lectotype & / Adela / imperialis / Rebel, 
1900 / teste M.Kozlov, 1994” (NHMW). Paralectotype @ (here desi- 
gnated) : same locality ; labelled : 3 X 7, print “Amur”; 5 X 21, black 
ink “Imperialis m[ihi].” ; 9 X 18, black ink “Nemotois / Paradisea Btl./ 
Imperialis Stg. 1.1./ Sn.” ; 5 X 14, print “Zool. Mus. / Berlin”; 13 X 
18, print + black ink “Paralectotype & / Adela / imperialis / Rebel, 
1900 / teste M.Kozlov, 1994” (ZMB). Syn. n. 

Nemotois imperialis (Rebel) : Meyrick, 1912 : 7 (E. Siberia). 

Adela imperialis Rebel : Szent-Ivany, 1945, Fragmenta Faunistica Hungarica 
8 (1) : 9 (Hungary) (err. det.). 

Nemaphora [sic!] imperialis (Rebel) : Gozmany, 1965 : 40 (Budapest) (err. 
det.). 

Adela sythoffi Snellen, 1901, Tydschr. Ent. 44: 77, pl. 5 fig. 4. Lectotype 
© (here designated) : Indonesia, W Java, Preanger ; labelled: 7 X 9, 
black ink “W. Java / Preanger / 5000 vt / Sythoff ©” ; 13 X 18, print 
+ black ink “Lectotype © / Adela / sythoffi / Snellen, 1901 / teste 
M. Kozlov, 1994”; red 13 X 17, print + black ink “Museum Leiden 
/ LECTOTYPE @ / Adela / sythoffi / Snellen, 1901 : 77” (NML) 
[examined]. Paralectotype © (here designated) : same locality ; labelled : 
7 X 9, black ink “W. Java / Preanger / 5000 vt / Sythoff 9”; blue 


43 


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13 X 17, print + black ink “Museum Leiden / PARALECTOTYPE 
Q / Adela / sythoffi / Snellen, 1901” (NML) [not examined]. 

Nemotois sythoffi (Snellen): Meyrick, 1912: 7 (as junior synonym of N. 
bifasciella). 

Nemotois baibarana Matsumura, 1927, J. Coll. Agric. Hokkaido Imp. Univ., 
19: 7. Holotype © : Taiwan; labelled: 8 X 12, print “BAIBARA / 
YeSaltor7 Kikuchi2 reverse side, pencil 21 Feb; / 192672 9x 19, 
black ink “Adela Mats. / baibarana”; red 7 X 16, print “Type / 
Matsumura” ; red 10 X 21, black ink + print “Holo-type / Nemotois / 
baibarana / MATSUMURA” (EIHU) [examined]. Syn. n. 

Nemotois honei Meyrick, 1935, Mat. Microlepid. Chin. Prov. : 94. Lectotype 
Q (here designated) : China, Nanking ; labelled : circle diam 8 with violet 
border, print “Lecto-/type” ; 4.5 X 9.5, black ink “Nanking / China / 
Hföne]. 15.8.[19]33” ; 13 X 16, black ink + print “Nemotois / hoeneella / 
Car. & Meyr. / E. Meyrick det. / in Meyrick Coll.” ; 6 X 13, print 
“Meyrick Coll. / B. M. 1938-290” ; 3 X 18, black ink “hoeneella Car.” ; 
12 X 20, print “Erroneously labelled / by Meyrick — specimens / of 
honei and honeella | were switched in the / Meyrick collection. / Teste 
M. Kozlov, 1994” ; 13 X 18, print + black ink “Lectotype 9 / Nemotois / 
honei / Meyrick, 1935 / teste M. Kozlov, 1994” (BMNH) [examined]. 
Paralectotypes (designated by A. Popescu-Gorj, see Comments) : same 
locality ; 14, labelled : blue 5 X 20, print “Nanking (China) / 15.6.1933 
H. Hone”; 13 X 22, black ink “Nemotois / hoenei [sic!] Car / sp. 
nov.” ; 9 X 13, black ink “Type” ; 14 X 20, red margins, print + black 
ink “Romania [along the left margin] / LECTOTYPE / Nemotois @ / 
honei / Car. & Meyr. DES. / Dr. A. POPESCU-GORJ” ; 11 X 18, 
black ink “Lectotype © in / BMNH design. by / Kozlov & Robinson / 
1995 ; this speci- / men is paralectotype”, reverse side “M. Kozlov, 1995”. 
1 Q, labelled : blue 5 X 20, print “Nanking (China) / 15.6.1933 H. Hone” ; 
9 X 7, black ink “28.”; 14 X 20, violet margins, print +black ink 
“Romania [along the left margin] / ALLOLECTOTYPE / Nemotois 
© / honei / Car. & Meyr. DES. / Dr. A. POPESCU-GORJ”. 2 99, 
labelled : blue 5 X 20, print “Nanking (China) / 15.6.1933 H. Hone” ; 
14 X 20, green margins, print + black ink “Romämia [along the left 
margin] / PARALECTOTYPE / Nemotois 2 / honei / Car. & Meyr. 
DES. / Dr. A. POPESCU-GORJ”. 1 3, labelled: 7 X 17, black ink 
“Nanking, China / 15.V1.1933. H.Höne” [recent label added by 
A.Popescu-Gorj to the unlabelled specimen]; 14 X 20, green margins, 
print + black ink “Romänia [along the left margin] / PARALECTO- 
TYPE / Nemotois 9 / honei / Car. & Meyr. DES. / Dr. A. POPESCU- 
GORJ” ; 9 X 17, black ink “Incorrect recent / labelling - À type / from 
Mokanshan. / M. Kozlov, 1995” (MINGA) [examined]. Syn. n. 

Adela aurantibasella Caradja, 1938, Stettiner Ent. Ztg. 99 : 252-253. Holotype 
@: China, Shaowu [erroneously or alternatively cited as Lackbaumpass 
in the original description] ; labelled: 9 X 17, black frame, black ink 
“Shaowu / Fukien / 14.6”; 9 X 17, blue ink “Nu este tip [Not a type, 


45 


Romanian] | DP Gory’; 15 X 25, black frame, black ink “Adela / 
aurantibasella / Type ® Car” ; 13 X 18, black ink “This specimen / 
was considered / as HOLOTYPE of / Adela auranti- / basella Caradja / 
by M. Kozlov, 1995” (MINGA) [examined]. Syn. n. 


Redescription 


MALE : Forewing length 7.0-10.6 mm ; wing expanse 15-23 mm. Vertex 
yellowish brown, sometimes with ochreous tint ; frons yellow. Proboscis 
and palpi light yellow to light brown. Labial palpus short (about 0.3 
X vertical eye diameter) ; maxillary palpus very short, about half length 
of labial palpus. Compound eye enlarged, interocular index 0.9 to 1.8 ; 
occipital distance less than 0.45 X vertical eye diameter. Antenna 2.1- 
2.5 X length of forewing, with simple inwardly directed pegs. Scape 
and proximal region of flagellum (about 0.6 X forewing length) with 
cupreous brown or blackish slightly raised scales; distal region of 
flagellum silver white to grey. Tegula and thorax brown, greyish brown 
or bronze. Forewing with yellow transverse band bordered on both 
sides by a single row of dark brown scales followed by 3 to 5 rows 
of silver scales ; ground colour surrounding silver bands darker brown 
than at base and apex of forewing, but this difference obvious only 
in fresh specimens ; basal field with dark brown coloration spreading 
from medial fascia along costal margin, sometimes creating the 
impression of a separate dark, iridescent costal spot near wing base. 
Width of yellow band variable, from very narrow, as wide as silver- 
grey bands, to 2.0-2.5 X width of these bands. Most specimens with 
margins of yellow medial band almost parallel, but in some examples 
the yellow band is slightly narrower at costa than at termen; 
exceptionally it can be 1.5 times wider at costa than at termen ; forewing 
ground colour proximal to fascia similar to that of thorax and tegula ; 
distal area of forewing suffused with yellow scales and slightly paler. 
Cilia brown, bronze at apex. Hindwing light brown ; costal area grey ; 
cilia brownish-grey. Legs brown with purple iridescence, bases of 
tarsomeres yellow. Fore tibia with apical tuft of blackish scales ; 
epiphysis at two-fifths length of tibia, not reaching its tip. Abdomen 
brownish grey to yellowish. 


FEMALE : Forewing length 6.5-11.6 mm, wing expanse 14-24 mm. Eye 
enlarged ; interocular index 0.7-0.8. Antenna 0.9-1.2 X length of 
forewing, with basal region of flagellum thickened by raised scales 
which are ochreous-yellow in the basal third to purplish-brown in the 
distal third ; extent of raised scales along antenna varying from 0.5 
to 0.7 length of forewing ; tip of flagellum yellow, with dark brown 


46 


ring at apex of each flagellomere. Thorax and tegulae yellow to light 
brown. Basal one-fifth of forewing yellow, except for very narrow dark 
brown costal area. Otherwise similar to male. 


MALE GENITALIA: Tegumen dome-shaped, with prominent medial 
ridge. Socii small, slightly less than diameter of aedeagus. Vinculum 
often with long cylindrical posterior region (up to 1/4 of total length), 
almost triangular anteriorly; posterior margin usually prominent 
medially. Valva significantly longer than tegumen, with prominent lobe 
on ventral margin; tip very narrow, pointed ; dorsal margin slightly 
emarginate ; viewed laterally, the valva appears almost rhomboidal. 
Valvae fused medially. Aedeagus straight to bowed, with pair of lateral 
Carinae near apex; base slightly swollen. Juxta short (about 0.6 X 
aedeagus length), the arrow-head very narrow, with pointed apex. 


en 


Figs 8-11. Forewing pattern : 8, 9 — Nemophora decisella ; 10, 11 — N. cantharites. 
8, 10 — males ; 9, 11 — females. Scale : 1 mm. 


47 


Figs 12-22. Male genitalia: 12-16 — Nemophora decisella, Amur reg. ; 17 — N. 
decisella, Sumatra ; 18-22 — N. cantharites, Mindanao. 12, 18 — genital complex, 
ventral view ; 13, 19 — same, lateral view ; 14, 20 — aedeagus, ventral view; 15, 17, 
21 — same, lateral view ; 16, 22 — juxta. Scale : 0.25 mm. 


48 


BroLocy : In the northern part of its range this appears to be a late 
summer species, adults having been collected from August to October. 
In the south, specimens were collected in all months except November. 
It is likely that N. decisella in SE Asia has more than one generation. 
In Japan it inhabits mountain regions (Okano, 1959) ; in SE Asia it 
has been collected in lowland and montane primary forest to 2030 
m (Robinson et al., 1994). The difference in temperature between these 
habitats is extreme and suggests that this species has remarkable 
plasticity. 


This is normally a diurnal species, but a very few specimens have also 
been collected at mercury-vapour light in Borneo in the early evening. 


DISTRIBUTION (Fig. 29): Russia (Khabarovsk reg., Primorye), Japan 
(Honshu, Shikoku, Kyushu, Tsushima), Korea, China (incl. Taiwan), 
Burma, Thailand, West Malaysia (incl. Langkawi), Indonesia (Sumatra, 
Java, Anambas Is, Kalimantan), Brunei, East Malaysia (Sarawak, 
Sabah). 


GEOGRAPHICAL VARIATION : Although specimens of decisella from its 
two most widely separated known localities, the Amur region and Java, 
look very different, all the characteristics investigated (see Fig. | for 
explanations) show clinal latitudinal variation. Forewing length in both 
males and females (Fig. 23), the relative size of the compound eyes 
(Fig. 26) and the interocular index in males (Fig. 27) are greater the 
further south the origin of the specimen. Concomitant with an increase 
in male eye size, the occipital distance between the eyes decreases from 
0.25-0.45 X vertical eye diameter in specimens from the Amur region 
and Japan to zero or almost zero in specimens from near the equator 
(Fig. 28). The relative width of the forewing yellow band decreases 
southwards (Fig. 24) while the position of the fascia, basal in northern 
specimens, becomes progressively closer to the middle of the wing 
(Fig. 25). No variation in these traits correlate with the longitude of 
collecting-localities. 


Variation in male genital structure is expressed mainly in the form 
and extent of the medial lobe of the valva, the shape of the aedeagus, 
and the relative size of the lateral carinae on the aedeagus. Since only 
a few specimens were dissected, any geographical trend in variation 
is difficult to quantify ; however, the southern populations seem to 
have larger and better sclerotized lateral carinae (cf. Figs 9 and 11). 
Observed variation concerns only the relative size of some structures, 
and is not unusual in comparison with the intraspecific variation 
observed in other Nemophora species. 


49 


CoMMENTS : Due to its wide range, geographic variation and sexual 
dimorphism, this species has been described under at least nine different 
names, the oldest being N. decisella (Walker, 1863) based on a single 
male from Sumatra. The next available name, N. bifasciella (Walker, 
1866), is based on a single female from Java. It should be noted that 
Walker misidentified the sex of the type specimen of bifasciella. 
However, the specimen labelled as the type in the BMNH collection 
corresponds completely to Walker’s description. N. sythoffi, described 
from Java by Snellen (1901), was synonymized with N. bifasciella by 
Meyrick (1912). Other descriptions of this remarkable species were 
based on specimens from the Russian Far East, Japan and China. 
In Japanese publications (Okano, 1959 ; Moriuti, 1982) the species is 
known under the name N. paradisea (Butler). 


ie nes 0.1 4 


© 
| 
© females) 0 1 3 e N} (2 


= 05110 : ‘ 
0.09 SES 

20 

O 

TD 

= 

Q 

=) 

O 

2 

O 

= 


Occipital distance 


0 20 40 
Latitude Latitude 


Figs 23 — 28. Variation of external characters of Nemophora decisella in relation to 
the latitude of localities : 23, forewing length, mm (I, see Fig. 1) ; 24, relative width 
of medial yellow band (wf/l) ; 25, position of outer border of medial yellow band 
in forewing (pf/1) ; 26, relative size of male compound eye (vd/1) ; 27, male interocular 
index (vd/id) ; 28, male occipital distance (md/vd). Each dot corresponds to one 
specimen. 


50 


N. heteroxantha Diakonoff, 1951, described from Kletak Pass, Tenger 
Mts in E Java (the type locality of N. bifasciella irrorata), 1s probably 
also a synonym of N. decisella. No males of N. heteroxantha are known, 
and females differ in some traits from N. decisella. Since we did not 
consider variation in female genitalia of N. decisella, the taxonomic 
value of differences pointed out by Diakonoff (1951) remains unclear 
and at the recent level of knowledge we consider N. heteroxantha as 
a distinct species. 


Rebel (1900) in his original description of Adela imperialis (based on 
two males from the Amur region of Russia) cited Staudinger (in litt.) 
as the author of the species, and mentioned the name ‘paradisea B.- 
Haas in litt.’ as a junior synonym. Thus, most probably, Rebel based 
his description of imperialis on specimens that had already been 
correctly determined as N. paradisea Butler by Bang-Haas or Stau- 
dinger. N. imperialis (Rebel) was listed by Meyrick (1912) as a valid 
species and then completely forgotten until the name was used again 
by Szent-Ivany (1945) and then by Gozmany (1965) for an erroneously 
determined melanic specimen of N. degeerella (L.) from Hungary 
(Küppers, 1980 : 263). 


The two specimens identified here as syntypes of Adela imperialis were 
discovered in the collections of NHMW and ZMB. We designate the 
specimen from NHMW as lectotype because it was already marked 
as “Type” and many of Rebel’s types are already located in Wien. 


N. honei (Meyrick) was described from seven specimens, two males 
and five females. We designate the female specimen in BMNH as 
lectotype. Although most of Caradja’s material is housed in MINGA, 
Bucharest, Meyrick states explicitly in the introduction to his paper 
(Meyrick, 1935) that he retained the primary type of each new species 
unless stated otherwise. | 


Although a specimen of honei was labelled as lectotype in the MINGA 
collection by A. Popescu-Gorj, this species was omitted from his 
catalogue (Popescu-Gorj, 1992) and the designation is unpublished ; 
the five MINGA syntypes are therefore paralectotypes. One of these 
Specimens (male) had no original data label, and a recent label 
“Nanking, China / 15.V1.1933 H.Höne” has been added by A. Popescu- 
Gorj. However, according to the original description, the second male 
syntype originated from Mokanshan, and therefore this recent label 
should be considered incorrect. 


Popescu-Gorj (1992) was unable to find the holotype of N. aurantibasella 
(Caradja) in the MINGA collection, and we can confirm that no female 


SI 


specimen of decisella is labelled “Lackbaumpass, 12.5” as in the original 
description. However, one female specimen fits perfectly the original 
description, particularly in its exceptionally small (14 mm) wing 
expanse. This specimen bears a type label of aurantibasella hand-written 
by Caradja, but is labelled “Shaowu, 14.6”. The loss of a unique type 
from the Caradja collection would be unprecedented, and we suggest 
the alternative explanation , that the label data was wrongly cited (or 
a geographic alternative given) in the original description. We consider 
this female to be the holotype of aurantibasella. 


MATERIAL EXAMINED: Russia: | 4, Khabarovsk (ZMUC); 3 @@, 2 99, 
“Amur” (Staudinger) (ZIN); 1 &, same label (NML); 2 44, “Amur” 
(BMNH); 2 29, Khabarovsk reg., Khingan mt., 28.V11.1877 (ZIN) ; 3 38, 
3 99, S Primorye, Vladivostok, Okeanskaya, 26. VII-4.VIII.1950 (Zagulajev) 
(ZIN); 1 4, 1 9, S Primorye, 20 km SE Ussurisk, Gornotaezhnoye, 1- 
3.V111.1966 (Zabello) (ZIN); 1 &, S Primorye, Khasan distr., Sidemi 
(Jankowski) (BMNH); 1 &, 2 99, S Primorye, Khasan distr., 3 km SE 
Andreevka, 1-11. VIII.1985 (Sinev) (ZIN). Japan: 3 64, Honshu, Fukusima 
Pref., Bantai Mt., 5.V111.1927 (Issiki) (USNM); 1 4, Honshu, Fukui Pref., 
Tsuruga, 4.1X.1917 (Roshkovskij) (ZIN) ; 1 9, Honshu, Gifu Pref., Katayama, 
8.V111.1920 (Takeuchi) (USNM); 1 @, Honshu, Kyoto Pref., Hanase, 
28.V111.1953 (Issiki) (USNM) ; 1 ©, Honshu, Shiga Pref., Yogo-cho, Kanzanji, 
28.V111.1993 (Hirowatari) (MZH) ; 1 À, 2 99, Honshu, Osaka Pref., Iwawaki 
Mt., 19.V111.1920 (Issiki) (USNM) ; 1 4, 1 9, Honshu, incomplete data, 1886 
(Pryer) (BMNH); 1 ©, Kyushu, Kagoshima Pref., Kanoya, 11.1X.1895 
(Wileman) (BMNH); 1 ©, Kyushu, Kagoshima Pref., Takakumayama, 
13.1X.1895 (Wileman) (BMNH). South Korea: | À, incomplete data (ZIN) ; 
1 4, V1.1926 (Issiki) (USNM). China: 3 66, 11 Q9, Ichang, Chang-Yang, 
1200-1800 m, VIII.1886 (Leech) (BMNH); 2 @@, Kia-Ting-Fu, V-VII.1891 
(Leech) (BMNH); 2 44, 1 9, Sztschwan, Tai-tou-ho (Déjean) (BMNH) ; 1 
@, Taiwan, Musya, 19.V11.1925 (Issiki) (USNM); | 4, Taiwan, V11.1947 (Li) 
(USNM); 1 4, Taiwan, Koshun, 25.1V-25.V.1918 (Sonan, Miyake & Yoshina) 
(USNM); 1 4, 1 9, Taiwan, Parasan, 27.VII.1943 (Issiki) (USNM). Burma : 
2 66, King Island [Kadan Kyun], Mergui, 7.11.1924 (Archibald) (BMNH) ; 
11 44, Mergui, Tenasserim, 1888 (Doherty) (BMNH) ; 1 44, incomplete data, 
2.X.1895 (Hampson) (BMNH). THAILAND: 2 @@, Uthai Thani Distr., 
Khao Nang Rum, 400 m, 1.111.1986 (Allen) (BMNH). West Malaysia: 1 &, 
Pahang, Fraser’s Hill, 400 m, 1.V11.1933 (Pendlebury) (BMNH); I 4, Kedah, 
Langkawi Is., 15.1V.1928 (Pendlebury) (BMNH); | 4, Pahang, Cameron 
Highlands, Gunung Brinchang, 1980 m, 14.V.1939 (Pendlebury) (BMNH) ; 
1 À, 1 9, Perak, Padang Rengas, low country, 1891 (Doherty) (BMNH) ; 
1 6, 1 9, Perak (Doherty) (BMNH) ; 1 4, Trong, 11.1899 (Abbott) (USNM) ; 
1 4, KhowSaiDow Mt. 350 m, 11.1899 (Abbott) (USNM). East Malaysia : 
2 64, Sarawak, Kuching, 1.1907 (J. H[ewitt]) (BMNH); 1 4, same locality, 
16.1V.1900 (J. H[ewitt]) (BMNH); 1 &, Sabah, Gunong Monkobo, 945 m, 
dipterocarp forest, 14-23. VIII. 1987, collected at light (Tuck) (BMNH). Brunei : 


yy) 


1 9, Rampayoh R. (north), LP 195, lowland forest, 1-3.111.1982 (Robinson) 
(BMNH). Indonesia: 11 35, Kalimantan, Riam Kiwa, 150-350 m, 1891 
(Doherty) (BMNH); 1 4, 1 ©, E Kalimantan, Tabang, Bengen River, 125 
m, 20.1X.1956 & 26.X.1956 (Wegner) (NML); | &, Anambas Is., P[ulau] 
Jemaja, Letung (Chasen / Raffles Mus.) (BMNH); | 6, W Sumatra, Lebong 
Tandai 1920-1923 (Brooks) (BMNH); 1 4, 2 29, SW Sumatra, Marang, 
sea level, VIII-IX.1890 (Doherty) (BMNH); 1 9, SW Sumatra, Liwa, 1000- 
1350 m, VII-V111.1890 (Doherty) (BMNH); 1 ©, Java, Telawa, 6.X11.1937 
(Kalshowen) (NLM); | &, W Java, mt. Megamendung, 700 m, 14.1X.1952 
(Wegner) (NLM); 1 46, W Java, Udjung Kulon, Njewaän, 18.V11.1955 
(Wegner) (NLM); 1 à, E Java, Nongkodjadjar, 1350 m, V.1934 (Kalis) 
(BMNH); 1 4, S Java, Kalipari, 350 m, 1891 (Doherty) (BMNH); | 6, 
Java, incomplete data (BMNH). Unidentified localities: 1 4, Lambiden, 
V.1892 (D[oherty]) (BMNH); 1 ©, Japan, Kurigehara, 6.VIII.1881 (Lewis) 
(BMNH). 


Nemophora cantharites (Meyrick, 1928) (Figs 6, 7, 10, 11, 18-22) 


Nemotois cantharites Meyrick, 1928, Exotic Microlepidoptera 3 : 464. Holotype 
© : Philippines, Mindanao, Kolambugan ; labelled : circle diam 8 with 
red border, print “Holo-/ type” ; 5 X 10, black ink “Mindanao / Philip- 
pines / Blanks]. .6.[19]17”; 3 X 20, black ink “cantharites Meyr.” ; 
13 X 16, black ink + print “Nemotois / cantharites / Meyr. / E. Meyrick 
det. / in Meyrick Coll.” ; 6 X 13, print “Meyrick Coll. / B. M. 1938- 
290” (BMNH) [examined]. 

Nemophora cantharites (Meyrick) : Diakonoff, [1968]: 294, fig. 797 (female 
wing pattern) (Mindanao). 

Nemotois heliochalca Meyrick, 1928, Exotic Microlepidoptera 3: 464-465. 
Holotype & : Philippines, Mindanao, Kolambugan ; labelled: circle 
diam 8 with red border, print “Holo-/type”; 5 X 10, black ink 
“Mindanao / Philippines / Blanks]. .6.[19]17”; 13 X 16, black ink + 
print “Nemotois / heliochalca / Meyr. / E. Meyrick det. / in Meyrick 
Coll.” ; 6 X 13, print “Meyrick Coll. / B. M. 1938-290”; blue 4 X 
13, print “Abdomen / missing” ; 3 X 21, black ink “heliochalca Meyr.” 
(BMNH) [examined]. Syn. n. 

Nemophora heliochalca (Meyrick) : Diakonoff, [1968] : 294, figs 468-469 (male 
genitalia), 799 (male wing pattern) (Mindanao) ; Robinson et al., 1994: 
23 (Philippines). 


Redescription 


MALE : Forewing length 9.4-10.3 mm ; wing expanse 19-23 mm. Vertex 
and frons ochreous to ferruginous. Proboscis and palpi light brown. 
Labial palpus short (about 0.3 X vertical eye diameter) ; maxillary 
palpus of same length as labial palpus, with terminal segment elongated, 
slender apically. Compound eye enlarged, interocular index about 1.5 ; 


53 


occipital distance 0.05-0.08 X vertical eye diameter. Antenna 2.4-2.5 X 
length of forewing, with simple inwardly directed pegs. Scape and 
flagellum uniformly brown ; basal region of flagellum (about 0.3-0.4 X 
forewing length) with cupreous brown or blackish slightly raised scales. 
Thorax bronze ; tegulae cupreous brown. Forewing with narrow bright 
ochreous transverse band at two-fifths, bordered on both sides by a 
single row of dark brown scales followed by 4 to 7 rows of light golden 
scales ; ground colour surrounding golden bands darker brown than 
at base and apex of forewing, but this difference obvious only in fresh 
specimens ; basal field with dark coloration spreading along radial stem, 
costal area bronze, terminal zone light brown ; ground colour of distal 
region of forewing light brown, suffused with golden scales. Cilia 
bronze. Hindwing brown ; costal area grey ; cilia brownish-grey. Legs 
brown, tibiae cupreous brown. Fore tibia with apical tuft of blackish 
scales ; epiphysis at one-half of the tibia, not reaching its apex. 
Abdomen brownish grey to yellowish. 


FEMALE : Forewing length 8.6-9.5 mm, wing expanse 18-21 mm. Eye 
enlarged ; interocular index about 0.9. Antenna about 1.2 X length 
of forewing, with basal third thickened by raised blue-purple to black 
scales ; tip of flagellum grey, with dark grey rings at apex of each 
flagellomere. Thorax and tegulae dark indigo-green. Ground colour 
of forewing dark brown, with indigo-green iridescence ; medial fascia 
dark brown, bordered by light golden bands as in male. Cilia dark 
brown. Hindwing dark brown, cilia brown. Legs dark brown, with 
indigo-green iridescence. Otherwise similar to male. 


MALE GENITALIA: Tegumen dome-shaped, with prominent medial 
ridge. Soci small, slightly less than diameter of aedeagus. Vinculum 
almost triangular. Valva of same length as tegumen, with prominent 
lobe on ventral margin ; tip widely rounded ; dorsal margin straight. 
Valvae not fused medially. Aedeagus with apical sclerotized band almost 
perpendicular to longitudinal axis (in lateral view) ; base of aedeagus 
wide, almost triangular (in dorsal view). Juxta short (about 0.5 X 
aedeagus length), the arrow-head very narrow, with elongate pointed 
apex. 


DISTRIBUTION (Fig. 29) : Philippines (Mindanao). 


COMMENTS : Although the description of N. cantharites is based on 
a single specimen which must be considered the holotype, a female 
in the USNM collection is labelled as a paratype (Diakonoff, [ 19681) ; 
it has no type status. 


54 


Fig. 29. Distribution of Nemophora decisella (dots) and N. cantharites (triangle). 


MATERIAL EXAMINED : Philippines : 23 6&, 6 2%, Mindanao, Kolambugan, 
25.V-20.V1.1914 (Wileman) (BMNH); 4 44, 2 29, same locality, VI.1914 
(Banks) (USNM). 


Acknowledgements 


We are extremely grateful to T. Hirowatari, R. de Jong, N. P. Kristensen, 
B. Krutzsch, T. Kumata, M. Lödl, E. Nieukerken, A. Popescu-Gorj, D. Rusti 
and S. Sinev for providing us with much useful information and for the loan 
or donation of specimens and to K. Tuck for commenting an earlier draft 
of the paper. M. Kozlov acknowledges with thanks the financial support of 
the British Council for a two week study trip to the BMNH, and thanks 
Michael Shaffer for his assistance during that visit. The research visit of M. 
Kozlov to MINGA was jointly funded by the Finnish and Rumanian 
Academies. Photographs were taken by the BMNH Photographic Unit. 


References 


Davis, D. R., 1975. West Indian moths of the family Psychidae with descrip- 
tions of new taxa and immature stages. Smithson. Contr. Zool. 188 : 
1-66. 


38 


DiAkONOFF, A., 1951. Records and descriptions of Microlepidoptera (4). 
Treubia 21 : 133-182. 

DIAKONOFF, A., 1955. Microlepidoptera of New Guinea: results of the third 
Archbold Expedition. Pt. V. Verh. K. ned. Akad. Wet., Ser. 2, 50 (3): 
1-211. 

DIAKONOFF, A., 1967 [1968]. Microlepidoptera of the Philippine Islands. Bull. 
U.S. natn. Mus. 257 : 1-484. 

GozMANY, L., 1965. Incurvariidae — hosszucsapu molyok. In Gozmany, L. 
& Szöcs, J.: Microlepidoptera I. Fauna Hungariae. XVI. Kötet - 
Lepidoptera. 2. Füzet (no. 76). pp. 17-41. Akadémiai Kiadö, Budapest. 

ICZN, International Commission on Zoological Nomenclature, 1985. Inter- 
national Code of Zoological Nomenclature, 3rd edition. 338 pp. Inter- 
national Trust for Zoological Nomenclature, London and University of 
California Press, Berkeley & Los Angeles. 

Koz tov, M. V., 1993. New species of Cauchas Zeller (Lepidoptera : Adelidae) 
from the Altai and Tianshan Mountains. Nota lepid. 16 : 113-123. 
Kuppers, P. V., 1980. Untersuchungen zur Taxonomie und Phylogenie der 
Westpaläarktischen Adelinae (Lepidoptera : Adelidae). 497 pp. Verlag 

M. Wahl, Karlsruhe. 

MEYRICK, E., 1912. Lepidoptera Heterocera (Tineae). Fam. Adelidae. Genera 
Insect. 133 : 1-12. 

Meyrick, E., 1935. List of the Microlepidoptera of Chekiang, Kiangsu and 
Hunan. /n Caradja, A. & Meyrick, E.: Materialen zu einer Microle- 
pidopteren Fauna der chinesischen Provinzen Kiangsu, Chekiang und 
Hunan. 96 pp. R. Friedlander & Sohn, Berlin. 

MoriUTi, S., 1982. Incurvariidae. /n Inoue, H. et al. : Moths of Japan, Vol. 2, 
pp. 155-156. Kodansha, Tokyo. 

Nye, I. W. B., & FLETCHER, D. S., 1991. The generic names of moths of 
the World. Vol. 6 : Microlepidoptera. xxıx + 368 pp. London. 

Oxano, M., 1959. Adelidae. Jn Inoue, H. et al. (Eds.): Iconographia 
Insectorum Japonicorum : Colore naturali edita. 2nd edn., pp. 277-278. 
Tokyo. 

Poprescu-GorJ, A., 1992. Le catalogue des types de Lépidoptères gardés dans 
les collections du Muséum d’Historie Naturelle «Grigore Antipa» 
(Bucarest) (fam. Micropterigidae -Pterophoridae). Trav. Mus. Hist. nat. 
«Gr. Antipa» 32 : 131-184. 

REBEL, H., 1900. Neue paläarktische Tineen. Dr. ent. Z. Iris 13: 161-188. 

Rosinson, G. S., Tuck, K. R., & SHAFFER, M., 1994. A field guide to the 
smaller moths of South-East Asia. 309 pp., 51 figs, 32 col. pls., Malaysian 
Nature Society, Kuala Lumpur. 

SNELLEN, P. C. T., 1901. Beschrijvingen van nieuwe exotische Tortricinen, 
Tineinen en Pterophorinen. Tijdschr. Ent. 44 : 67-98. 

SZENT-IVANY, J., 1945. Faunistische und ökologische Angaben über die 
Adelinen (Lepidopt.) des Karpatenbeckens mit Beschreibung zwei neuer 
Arten. Fragm. Faun. hung. 8 (1) : 7-10. 

Urecu, F. H., 1980. Provinces and prefectures of Japan : a biogeographical 
aid. Ann. Carneg. Mus. 49 : 161-175. 


56 


Nota lepid. 18 (1) : 57-76 ; 31.1.1996 ISSN 0342-7536 


Behaviour and morphology of an 
aphytophagous lycaenid caterpillar : 
Cigaritis (Apharitis) acamas Klug, 1834 
(Lepidoptera : Lycaenidae) 


Matthias SANETRA* & Konrad FIEDLER** 


* Institut fiir Zoologie, Technische Hochschule Darmstadt, Schnittspahnstr. 3, D-64287 Darmstadt, 
Germany 

** Theodor-Boveri-Biozentrum der Universitat, LS Verhaltensphysiologie und Soziobiologie, Am 
Hubland, D-97074 Wiirzburg, Germany 


Summary 


Interactions between caterpillars of Cigaritis (Apharitis) acamas Klug, 1834 
from Cyprus and their host ants, Crematogaster cf. jehovae Forel, 1907, were 
observed in captivity. At least final instar caterpillars permanently live inside 
ant nests where they mainly prey on ant brood. To a lesser extent, ant regur- 
gitations also contribute to the caterpillars’ nutrition. Caterpillars are perma- 
nently antennated and cleaned by their attendant ants. Details on the mor- 
phology of the myrmecophilous organs in Cigaritis larvae are provided for 
the first time. The relations of Cypriot Cigaritis acamas to other members 
of the genus from neighbouring arid zones and to related lycaenid species 
are discussed, especially with respect to myrmecophily and ecological requi- 
rements. 


Zusammenfassung 


Zwei Raupen von Cigaritis (Apharitis ) acamas Klug, 1834 wurden auf Zypern 
einzeln in den Nestern der bodennistenden Ameisenart, Crematogaster cf. 
Jehovae Forel, 1907 gefunden. In Labornestern wurden die Interaktionen 
zwischen Raupen und Ameisen beobachtet. Die Raupen wurden ständig von 
ihren Wirtsameisen betrillert. Die Ameisen leckten die gesamte Oberflache 
der Raupen ab und zogen deren lange laterale Borsten durch ihre Mandibeln. 
Sekrete aus dem dorsalen Nektarorgan wurden sofort von den Ameisen 
aufgenommen, die auch den fliissigen Kot der Raupen abnahmen. Reaktionen 
der Ameisen auf Ausstiilpungen der Tentakelorgane wurden nicht beobachtet. 
Hauptnahrung der Altraupe ist die Brut der Wirtsameisen, daneben werden 
auch durch taktile Stimulation mit den Vorderbeinen trophallaktische Regur- 
gitationen erbettelt. Die Beobachtungen zeigen, daß zumindest die ausgewach- 
sene Raupe von C. acamas (vorletztes und letztes Stadium) als permanenter, 
voll integrierter Myrmekophiler in den Nestern von Crematogaster-Arten lebt, 
wo sie den Wirten durch Brutfraß beträchtlichen Schaden zufügt. Die Epi- 


57 


dermalorgane der Raupe zeigen starke Anpassungen an ihre obligate Myr- 
mekophilie. Neben einem dorsalen Nektarorgan und den fiir Raupen der 
Tribus Aphnaeini typischen großen Tentakelorgane besitzt die Larve sehr 
zahlreiche Porenkuppelorgane und dendritische Haare, außerdem laterale 
Gruppen langer Borsten und — caudal vom Nektarorgan gelegen — eine 
Gruppe pilzförmiger Haare mit stark poriger Oberfläche. Struktur und 
Verhaltensbeobachtungen legen nahe, daß alle Organe an der Kommunikation 
mit Ameisen teilhaben. Die Beobachtungen an zypriotischen C. acamas werden 
mit Angaben zur Larvalbiologie verwandter Bläulinge verglichen und im 
Kontext mit der Zoogeographie und Evolution der Aphnaeini diskutiert. 


Resume 


Les auteurs ont trouvé sur l’île de Chypre deux chenilles de Cigaritis (Apha- 
ritis) acamas Klug, 1934, isolées, dans les nids de la fourmi terricole Cremo- 
gaster cf. jehovae Forel, 1907. Ils ont observé en laboratoire les relations entre 
ces chenilles et les fourmis dans des nids artificiels. Les chenilles étaient 
constamment titillées par les fourmis-hôtes, qui les léchaient sur tout le corps 
et tiraient à travers leurs mandibules les longs poils latéraux ; les sécrétions 
des organes nectarigènes dorsaux étaient immédiatement absorbées, de même 
que les excréments liquides. On n’a pas constaté de réaction des fourmis aux 
dégagements des organes-tentacules. La nourriture principale de la chenille 
adulte est constituée par les œufs et larves des fourmis-hôtes ; par stimulation 
tactile avec les pattes antérieures, celles-ci mendient aussi des régurgitations 
trophallactiques. Ces observations montrent que la chenille adulte de C. 
acamas (au moins aux dernier et avant-dernier stades) vit dans les nids des 
forumis Crematogaster comme hôte myrmécophile permanent, complètement 
intégré. En se nourrissant des larves de ses hôtes, elle leur cause de sérieux 
dégâts. Les organes épidermiques de cette chenille montrent une adaptation 
très nette à leur vie en myrmécophilie obligatoire. Outre un organe nectarigène 
dorsal, et les grands organes-tentacules typiques des chenilles de la tribu des 
Aphnaeini, cette chenille présente de très nombreux organes pore-coupole et 
des poils dentritiques, ainsi que des groupes latéraux de longues soies avec, 
du côté caudal de l'organe nectarigène, un groupe de poils en forme de 
champignon dont la surface est très poreuse. Cette structure et les compor- 
tements observés suggèrent que tous les organes participent à la symbiose 
avec les fourmis. Les auteurs comparent leurs observations sur C. acamas 
de Chypre avec ce qu’on sait de la biologie larvaire des Lycènes apparentées 
et, dans ce contexte, discutent de la zoogéographie et de l’évolution des 
Aphnaeini. 


Introduction 


Myrmecophily within the Lycaenidae is extremely widespread, and a 
variety of relationships, especially between caterpillars and ants, have 
been described. These interactions range from broadly unspecific 


58 


peaceful coexistence or facultative and unspecific mutualism to highly 
advanced mutualistic and sometimes parasitic associations with specific 
host ants (for review see Fiedler, 1991). Caterpillars of the predominantly 
African tribe Aphnaeini exhibit an extraordinarily high degree of 
myrmecophily. As far as their life-cycles are yet known, most Aphnaeini 
species are obligatorily associated with specific ants (Clark & Dickson, 
1971 ; Fiedler, 1991). In particular, various reports on these fairly close 
and specific relationships between Aphnaeini caterpillars and ants (e.g. 
Larsen & Pittaway, 1982) hint towards commensalic or parasitic 
exploitation of host ant colonies (mostly in the myrmicine genus 
Crematogaster ) rather than suggest mutualistic symbiotic associations. 
Thus, the peculiar ecology of Aphnaeini butterflies and the highly 
specialized morphology of their immature stages deserve further 
investigation. 


The genus Cigaritis Donzel, 1847, consisting of two so-called subgenera 
(sometimes treated as separate genera), is the only clade of the 
Aphnaeini to reach the southern margin of the western Palearctic realm. 
Cigaritis s. str. contains three species endemic to northwestern Africa 
(zohra, allardi, siphax), which have adapted to a more temperate 
Mediterranean climate, while the six to nine species (depending on 
the taxonomic status attributed to some local forms or subspecies) of 
the subgenus Apharitis are centred in the arid zones of North Africa, 
Arabia and Asia (Larsen & Pittaway, 1982). Cigaritis (Apharitis) 
acamas Klug, 1834 is by far the most widely distributed member of 
the genus, recorded from the fringes of the Sahara across the Middle 
East to northwestern India. It is therefore not surprising that it has 
also managed to reach the island of Cyprus (Riley, 1925). 


Cigaritis 1s closely related to the paleotropical genus Spindasis, which 
occurs with some 50 species throughout all of Africa south of the 
Sahara desert and extends across India eastwards to Sundaland, the 
Philippines, southern China and the Japanese island of Okinawa. While 
Cigaritis s. str. appears to be a monophyletic unit, the separation 
between Apharitis and Spindasis is all but clear-cut (e.g. Riley, 1925 ; 
Wynter-Blyth, 1982), and a thorough cladistic revision may well reveal 
that all taxa in question are best grouped into a more inclusive genus 
Spindasis to avoid paraphyletic rest groups. 


Very little information is available on the biology of the genus Cigaritis, 
and the life-histories of all its species are far from being thoroughly 
investigated. Adults of Cigaritis are known to occur only very locally 
at low population densities, with the exception of the North African 
endemics C. zohra (Donzel, 1847) and C. allardi (Oberthiir, 1909) which 


59 


can be very abundant in suitable habitats, but also very local (Thomas 
& Mallorie, 1985 ; Sanetra, personal observations). Caterpillars of 
Cigaritis are more or less closely associated with ants of the genus 
Crematogaster, and Dumont (1922) provided the earliest report on 
these myrmecophilous life habits. He observed that caterpillars of C. 
myrmecophila in Tunisia spent the day in subterranean tunnels in the 
outskirts of Crematogaster nests, but came out to feed at ae on 
Calligonum comosum (Polygonaceae). 


In Oman, Larsen & Pittaway (1982) discovered three-quarters grown 
caterpillars as well as a pupal exuvia of C. acamas inside the galleries 
of a Crematogaster ant inhabiting palm trunks, and these authors also 
observed adoption behaviour of newly hatched larvae, which were 
carried off into the nest by foraging ants. Larvae of herbivorous C. 
zohra from Morocco rest and diapause specifically in nests of Cre- 
matogaster laestrygon (Rojo de la Paz, 1992). Facultative myrmecophily 
occurs in C. allardi whose caterpillars feed on green plants and are 
attended by various Crematogaster species, but which never penetrate 
into ant nests (Rojo de la Paz, 1992). Similarly, caterpillars of the 
Asian C. epargyros (Eversmann 1854) have been reported to feed on 
plants in company with unspecified ants (Lukhtanov & Lukhtanov, 
1994). 


The findings of Larsen & Pittaway (1982) suggest that at least in Arabia 
C. acamas caterpillars spend their whole life cycle inside ant nests, 
and raises the important question as to whether the larvae of C. acamas 
are aphytophagous commensales or even parasites of their ant hosts. 
We here present observations on C. acamas caterpillars from Cyprus 
inside artificial ant nests, which shed new light upon the feeding strategy 
of this species. In particular, we demonstrate for the first time that 
final instar caterpillars essentially prey on the brood of Crematogaster 
ants. Furthermore, we present the first investigation of the epidermal 
organs of Aphnaeini caterpillars using SEM techniques. 


Material and methods 


Two caterpillars of C. acamas from Cyprus (for detailed information 
see next section) were reared together with their host ants, Crematogaster 
cf. jehovae Forel, 1907, in the laboratory for about five weeks. When 
collected, one of the caterpillars had already reached the final instar, 
while the other one was in the penultimate (probably fourth) instar. 
Crematogaster colony fragments containing caterpillars were maintained 
in artificial nests at a temperature cycle of 15 : 25°C (night : day). Both 
fragments did not comprise a fertile queen, hence egg production was 


60 


interrupted. The behaviour of caterpillars and their interactions with 
ants Were observed every second day with variable duration using a 
stereomicroscope (magnification 12-25fold). 


When the caterpillars died (see below), they were stored in a freezer, 
later fixed in ethanol and finally post-fixed in Bouin’s fixative. One 
caterpillar was then critical-point dried, sputtered with gold and 
examined in a scanning electron microscope (Zeiss DSM 962). The 
other caterpillar was externally surveyed using a Wild Macroscope M 
400 (magnification 10-64fold). 


Results 


Habitat in Cyprus 


As within its whole range, C. acamas has a very patchy distribution 
on Cyprus, from which island only five localities have as yet been 
recorded (Parker, 1983 ; Manil, 1990). It was early spring (2.1v.1994), 
when the first author was fortunate enough to discover a new site 
in the Greek part of Cyprus, Distr. Paphos. The locality was situated 
on a northeastern slope in the vicinity of the small village Kritou 
Marottou (elevation about 500 m) at the rise of the Troodos mountains. 
The vegetation of the habitat can be characterized as a typical 
Mediterranean garrigue, in the spring aspect dominated by flowers like 
Asphodelus albus (Liliaceae). 


Within a small spot (estimated to be about 20 m in diameter), at which 
nests of the subterranean Crematogaster cf. jehovae were suddenly quite 
abundant, a single caterpillar of C. acamas was detected within the 
nest chambers. Thorough inspection of a further dozen of Crematogaster 
nests resulted in the discovery of a second caterpillar. The ant nests 
were built in the soil under stones and compared with arboreal 
Crematogaster species colonies appeared to be relatively small, contain- 
ing a few hundred workers. In the bordering area of the spot, other 
ant species out of the myrmicine genera Tetramorium, Pheidole, Messor 
and Aphaenogaster were the dominant subterranean ants. 


Interactions between caterpillars and ants 


After the Cigaritis caterpillars had been transferred into artificial nests 
together with their host ants, they spent most of the time motionless 
inside the nest chambers. No signs of aggressiveness were observed 
between the Crematogaster ants and the caterpillars. On the contrary, 
caterpillars were highly attractive to their hosts (Fig. 1). While resting 


61 


Figs 1-2. Mature caterpillar of Cigaritis acamas. 1 — Attended by a group of 
Crematogaster worker ants in a laboratory nest. Note the conspicuous black sheaths 
of the tentacle organs on the eighth abdominal segment ; 2 — Feeding on Crematogaster 
brood. The predacious caterpillar is groomed and antennated while preying on the 


ants’ young. (Photographs : M. Sanetra). 


62 


within the nest they were permanently surrounded by a large group 
of ant workers, estimated to number about 20-30 individuals. The 
attendants intensively groomed the caterpillars, thereby licking them 
all over their outer body parts as they usually did with their own 
nestmates and brood. In addition, they frequently pulled the long lateral 
setae of the caterpillars through their mandibles one by one very 
carefully. In order to take up the caterpillars’ liquid excrements, the 
ants sometimes licked the anal region of caterpillars. We never observed 
a caterpillar to discharge solid faeces. After moulting of the prefinal 
instar caterpillar into the ultimate instar, the ants tore the remaining 
exuvia apart and converted it into food. 


The two myrmecophilous organs commonly occurring within the family 
Lycaenidae, the dorsal nectar organ (DNO) and the tentacle organs 
(TOs), were well developed and functional in the caterpillars investigated 
from Cyprus. The vicinity of the DNO attracted the ants and was 
intensively antennated. When secretions were released from the DNO, 
these were eagerly imbibed by the attendant ants. Most of the time 
the TOs remained retracted. Eversions of tentacles could be observed 
either when the caterpillars wandered about or if they were disturbed. 
However, there was no visible reaction of the ants towards the 
periodically everted tentacles. About two weeks after collection 
(15.1v.1994) one of the caterpillars had freshly moulted into the final 
instar. Its colours were brighter than before and interestingly, though 
the caterpillar was not crawling around, its TOs were highly active. 
Again no specific reaction of the ants could be detected. 


Caterpillars were fed via trophallaxis by their host ants in a similar 
way to ants feeding each other. With its forelegs the caterpillar 
stimulated an ant to regurgitate a droplet of liquid from its crop. This 
kind of feeding behaviour was observed occasionally (3-4 times during 
approx. 5 h of irregular observation periods), but not as frequently 
as would be expected if caterpillars totally depended on trophallactic 
feeding. Eventually, it became evident what the main food source is 
of caterpillars inside ant nests enabling larval growth until pupation. 
A hungry caterpillar approached the ants’ brood chamber, grasped a 
piece of brood with its forelegs and pulled it down beneath the head 
(Fig. 2). Then immediately the caterpillar retracted its head under the 
sclerotized prothoracic shield, as if to protect itself against possible 
ant-attacks. Within a few minutes a caterpillar devoured several brood 
items (usually 3-5) in the described manner. The caterpillars seemingly 
preferred the larger brood pieces, such as prepupae and pupae, and 
later switched over to the younger larvae. 


63 


Since the predatory behaviour described above was observed in the 
final instar caterpillars during most observation periods, one can 
imagine that predation caused considerable harm to the ant nests. 
Unfortunately, neither of the Crematogaster colony fragments had a 
fertile queen, and due to the lack of new egg production the brood 
resources of both ant nests were depleted within a few days (by the 
18th and 25th April, respectively) because of the caterpillars’ great 
appetite. The total number of brood items which had been eaten by 
each caterpillar was estimated to be about 500 per nest, mainly small 
larvae. In the meantime, we tried to adapt the caterpillars to brood 
of the myrmicine genus Leptothorax, but without success. At the 
beginning of May both caterpillars, though almost fully grown, died 
of disease. 


Morphology of the mature caterpillar 


The mature caterpillars, 13 mm long and 3.5 mm in diameter, are 
almost round in cross-section (not flattened and woodlous-shaped as 
many Lycaenidae larvae). Their ground colour is whitish, with complex 
red markings all over the dorsal surface (dark elements in Figs. 1- 
2). Inspection with the stereomicroscope and the SEM revealed the 
following characteristic epidermal organs : 


a) The prothoracic shield (Fig. 3) is sclerotized and appears blackish. 
Its central part is almost free of setae, but towards the margins there 
occur dendritic setae in numbers. The anal plate is flattened and less 
strongly sclerotized than the prothoracic shield. 


b) A large dorsal nectar organ (DNO) on the seventh abdominal 
segment (Fig. 7). 


c) Associated with the DNO is a group of large, mushroom-shaped 
setae (diameter of distal flattened end 30-40 um). About 10-12 of these 
setae form a transverse row just posterior to the DNO (Figs. 7-8). 


— 


Figs 3-6. SEM photographs of epidermal structures observed in a mature C. acamas 
caterpillar. 3 — Smooth, strongly sclerotized prothoracic shield. The shield surface 
carries only small numbers of dendritic setae. Laterally and anteriorly the prothoracic 
shield is surrounded by long setae with a dentate surface. Scale : 500 um ; 4 — Long 
seta with dentate surface and group of much shorter dendritic setae. The long setae 
are regularly pulled by the ants through their mandibles. Scale : 100 um ; 5 — Dendritic 
hair on the sheath of a tentacle organ. Note the regular nests of microtrichia on the 
membraneous surface of the eversible part of the tentacle. Scale : 20 um ; 6 — Spiracle 
of an abdominal segment surrounded by dendritic setae and pore cupola organs (arrow). 
Scale : 50 ym. (Photographs A. Gerber/K. Fiedler). 


64 


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65 


The surface of the mushroom-setae is highly perforated (pore diameter 
0.1-0.15 um; Figs. 9-10) which, together with their large size and 
position, suggests that they are secretory and somehow involved in 
myrmecophily. Clark & Dickson (1971) figured apparently similar setae 
for a number of African Aphnaeini, but these are mostly situated at 
other locations (e.g. spiracles). Therefore, the homology of the mu- 
shroom-like setae in C. acamas awaits to be elucidated based on a 
broader survey of Aphnaeini immatures. 


d) The tentacle organs (TOs), located on the eighth abdominal segment, 
are of the typical Aphnaeini type. They are inserted in black, strongly 
sclerotized, elevated sheaths which are easily seen without any optical 
equipment. The sheaths of the TOs are equipped with dendritic setae 
(Fig. 11; see below). At the top of each sheath there is a “crown” 
of long setae with a granulate surface. The eversible part of the TOs 
is membraneous, covered with nests of microtrichia (Fig. 5), and carries 
long (0.5-1 mm), spiny setae on its distal end (Figs. 11-12), somehow 
resembling the spectacular TOs of Oriental Curetis larvae (Fiedler et 
al., 1995). 


e) Pore cupola organs occur, scattered all over the skin. There are no 
conspicuous clusters of these glands around the DNO or the spiracles. 


f) Tufts of long (0.6-1 mm) erect setae with a spiny surface (Fig. 4) 
occur in characteristic fashion. Above and below each abdominal 
spiracle there are two such tufts per segment, each comprising 4-8 hairs. 
Smaller groups (with ca. 4 hairs) occur at comparable locations on 
the thoracic segments. 


g) Much shorter (50-80 um) dendritic setae (Figs. 5-6) are present in 
very large numbers almost all over the larva, including the prothoracic 
shield and the sheaths of the TOs. Dendritic setae are likely to play 
a role in myrmecophily (Ballmer & Pratt, 1992). However, they usually 
occur in smaller numbers and concentrated at particular locations. As 
far as we are aware, a similarly dense cover of dendritic setae has 
as yet been observed only in the Oriental obligate myrmecophile Rapala 
dieneces (Fiedler, unpublished data). 


h) Two groups of short (80-120 um), erect, smooth bristles (probably 
mechanoreceptors) occur near the opening of the DNO (Fig. 7) and 
amidst the two TOs. 


i) We found no evidence whatsoever of “dish organs” on the abdominal 
segments 5-7. Such organs have been described and figured from a 
couple of African Aphnaeini species (Spindasis, Crudaria: Clark & 
Dickson, 1971), but until more material becomes available for a study 


66 


Figs 7-10. SEM photographs of epidermal structures observed in a mature C. acamas 
caterpillar. 7 — Dorsal nectar organ on the seventh abdominal segment. The glandular 
opening (arrow) is accompanied by a group of short, erect setae. Posterior to the 
glandular opening there is a group of large mushroom-like setae. Scale: 250 um; 
8 — Group of mushroom-like setae behind the DNO, enlarged. Scale: 25 um; 
9 — Mushroom-like seta. Scale: 10 um; 10 — Top surface of mushroom-like seta 
with numerous large pores. Scale : 2 um. (Photographs A. Gerber/K. Fiedler). 


67 


of their fine structure, the nature and significance of these organs 
remains mysterious. 


Discussion 


The few available records of Cigaritis acamas caterpillars (Larsen & 
Pittaway 1982; this study) suggest that the species is obligatorily 
myrmecophilous over its whole range. Crematogaster cf. jehovae serves 
as host ant for C. acamas on Cyprus and Benyamini (1990) mentioned 
the same ant species with respect to Israeli populations. Crematogaster 
jehovae is morphologically very similar to C. auberti Emery, 1869 which 
has been reported to maintain close myrmecophilous relationships with 
caterpillars of Cigaritis myrmecophila in Tunisia (Dumont, 1922). On 
the other hand, the specific identity of the host ants from Oman remains 
to be elucidated. By comparison with other species of the genus 
(Table 1), ant-dependence can be regarded as a common feature in 
Cigaritis, since caterpillars of this genus have always been found more 
or less closely associated with ants. Neither in the genus Cigaritis nor 
in other Aphnaeini genera is there any evidence of a single species 
being myrmecoxenous (Fiedler, 1991). 


In all Cigaritis species where the attendant ants have as yet been 
identified, these belong to the myrmicine ant genus Crematogaster. This 
ant genus also serves as host for most Aphnaeini species (33 out of 
41 species, 80.5 %), for which the identity of associated ants has been 
reported (e.g. in the genera Aphnaeus, Spindasis, Axiocerses, Chlo- 
roselas, Phasis, Poecilmitis, Oxychaeta: Clark & Dickson, 1971; 
Fiedler, 1991 ; Schlosz & Brinkman, 1991 ; Callaghan, 1993). Exceptions 
only occur in two closely allied African genera (Aloeides and Erikssonia ; 
with the formicine genus Lepisiota = Acantholepis), in Axiocerses 
amanga and Poecilmitis pyroeis (with the formicine genus Camponotus ), 
and in a few Spindasis species and Axiocerses harpax (with the 
myrmicine genus Pheidole). This pattern of host use suggests that a 
tight connection with Crematogaster ants was the ancestral condition 
among Aphnaeini lycaenids. Shifts to novel ant hosts occurred only 
in a small number of lineages. 


The only host ant record for Cigaritis involving an ant genus other 
than Crematogaster is that of Cataglyphis “bicolor” (see Agosti, 1990 ; 
Keegans et al., 1992 for recent contributions to the still unresolved 
taxonomy in that ant genus) reported by Dumont (1922) for Cigaritis 
myrmecophila. Our current knowledge of the biology of Cataglyphis 
ants (Wehner, 1987) suggests that this record is based on a misinter- 
pretation of Dumont’s observations in the Tunisian desert. Dumont 


68 


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69 


supposed the small Crematogaster ants to be slaves of the much bigger 
Cataglyphis ants. However, the solitary Cataglyphis workers, which 
Dumont had observed near the nests of Crematogaster, were almost 
certainly individual foragers and had no sociobiological relationship 
to the Crematogaster ants. Workers of Cataglyphis are well known 
to forage dozens of metres away from their home nests (Wehner, 1987). 
Socially parasitic associations between Cataglyphis and Crematogaster 
ants have never been discovered as yet (Hölldobler & Wilson, 1990), 
and such a relationship would be extremely unlikely to occur among 
members of two different ant subfamilies (Buschinger, 1990). Finally, 
no associations between Cataglyphis ants and larvae or pupae of other 
lycaenids have ever been observed (Fiedler, 1991 and unpublished data). 
Hence, Cataglyphis should be eliminated from the host ant list of 
Cigaritis butterflies, unless new field data become available. Unfortu- 
nately, Larsen & Pittaway (1982), when citing Dumont (1922), have 
uncritically reinforced this erroneous assumption. 


Cigaritis caterpillars apparently use a broad range of feeding strategies. 
Our observations revealed that prefinal and final instars of Cypriot 
C. acamas mainly prey on ant-brood, but additionally receive trophal- 
lactic regurgitations. Such a feeding behaviour, though not definitively 
observed before, had been postulated for C. acamas from Oman where 
caterpillars spend their whole life inside ant nests (Larsen & Pittaway, 
1982). We assume that, once adopted into Crematogaster nests, larval 
behaviour and utilization of food sources are very similar between C. 
acamas populations from Cyprus and Oman. Besides this aphytopha- 
gous feeding strategy of C. acamas, caterpillars of other Cigaritis have 
been found to feed on green plants of the families Fabaceae, Poly- 
gonaceae, or Cistaceae, like C. allardi, C. zohra (Rojo de la Paz, 1992), 
C. myrmecophila (Dumont, 1922) and C. epargyros (Lukhtanov & 
Lukhtanov, 1994). However, at least in the latter two species, where 
available information is very scant, we can at present not rule out 
the possibility that caterpillars obtain at least part of their food as 
predators or commensales in ant nests. 


On the other hand, we can as yet not ascertain if Cypriot C. acamas 
caterpillars are entirely aphytophagous during all instars. Wynter-Blyth 
(1982) assumed Cassia trees to be food-plants in India, but it remains 
unclear whether or not he really had found C. acamas larvae feeding 
on plant tissue or deduced this from having observed females laying 
eggs on these trees. The observations of Larsen & Pittaway (1982) 
on Omani populations indicate that C. acamas females deposit their 
eggs on certain plants nearby Crematogaster ant nests (e.g. date trunks 
in Oman, Cassia trees in India probably for the same reason). Ant- 


70 


Figs 11-12. SEM photographs of epidermal structures observed in a mature C. acamas 
caterpillar. 11 — Partially everted tentacle organ. The eversible part is covered with 
nests of microtrichia. Long hairs with dentate surface structre are protruding from 
the (invaginated) top of the eversible part. The sheath carries a large number of dendritic 
hairs. Scale : 100 um ; 12 — Dorsal view of partly everted TO with central long hairs 
and cuticular fine structure (microtrichia). The long, stiff erect setae with dentate 
surface, which surround the distal end of the TO sheath, have been lost prior to 
fixation (arrow points to insertion of one such hair). Scale: 50 um. (Photographs 
A. Gerber/K. Fiedler). 


71 


dependent oviposition is common among obligatorily myrmecophilous 
lycaenid butterflies (Pierce & Elgar, 1985 ; Fiedler, 1991) and ensures 
the establishment of the “right” caterpillar-ant association from the 
egg stage onwards. Later, caterpillars are adopted by the ants either 
directly after hatching or during one of the following instars. A similar 
strategy occurs among Australian Acrodipsas species, whose entirely 
aphytophagous larvae are adopted immediately after hatching from 
the eggs laid nearby the host ants’ nest (Samson, 1989). 


We could trace more or less reliable information on oviposition sub- 
strates, larval food resources and/or myrmecophily for 86 Aphnaeini 
species (Fiedler, 1991 & unpublished data ; Schlosz & Brinkman, 1991 ; 
Callaghan, 1993). Although most authors agree on the extraordinarily 
close relationships between Aphnaeini immatures and ants, our ob- 
servations on C. acamas are the first proof that preying on ant brood 
does really occur in this tribe. Although preying on ant brood has 
been postulated for species like Oxychata dicksoni, Spindasis takanonis 
and others, conclusive evidence is still lacking. Otherwise among the 
Lycaenidae, there are very few taxonomic groups where this feeding 
strategy has independently evolved (review: Cottrell, 1984). Besides 
the well-known genera Maculinea (Thomas et al., 1993 ; Elmes et al., 
1994) and Lepidochrysops (Henning, 1983), where young caterpillars 
are herbivorous and later turn into carnivores, examples are known 
from a couple of tropical Miletinae species (Fiedler, 1991) or Australian 
Acrodipsas (Samson, 1989), and strongly suspected from Indian Zesius 
chrysomallus (Yates, 1932). 


Trophallactic feeding is more widespread among myrmecophilous 
insects in general (Hölldobler & Wilson, 1990), and among lycaenids 
in particular. Besides C. acamas, this type of interaction has been 
confirmed in Aphnaeus adamsi and Spindasis takanonis, and is likely 
to occur in various other Aphnaeini species (Spindasis nyassae, Chloro- 
selas umbrosa). Trophallaxis also occurs, at least occasionally, in some 
Miletinae species, in Shirozua jonasi, Anthene levis and Niphanda fusca 
(Fiedler, 1991). 


Morphologically, the caterpillars of C. acamas are largely similar to 
other Aphnaeini whose immatures have been figured (Clark & Dickson, 
1971). The lack of any other SEM studies on Aphnaeini species, 
however, presently hampers exact comparisons. The significance of the 
long lateral hairs for the interactions with ants is suggested by the 
characteristic behaviour of attendant ants, which pulled these hairs 
through their mandibles. This behaviour could assist in collecting some 
putative secretions. Likewise, the intensive licking behaviour all over 


72 


the surface could well be caused by the numerous dendritic setae. From 
other lycaenids it is known that antennation behaviour of tending ants 
concentrates at locations with dendritic setae (Ballmer & Pratt, 1992 ; 
Fiedler, unpublished data). The mushroom-like setae with their large 
pores (much larger than in most pore cupola organs of lycaenid 
immatures) and the morphological associations of these setae with the 
nectar organ also suggest a strong connection of these organs with 
myrmecophily. On the other hand, we failed to find any equivalents 
of so-called “dish organs” which have been reported from a couple 
of Aphnaeini species with extremely close associations with ants. Given 
their tight relationships towards ants, the structure and function of 
epidermal organs of Aphnaeini immatures is one of the most rewarding 
fields for further studies on myrmecophilous lycaenids. 


The Cypriot populations of C. acamas differ in some aspects from 
those investigated by Larsen & Pittaway (1982) in Oman. The habitat 
on Cyprus has a typical Mediterranean climate which contrasts with 
the extreme desert conditions in Oman. Furthermore, host ant species 
of both populations are quite different according to their nesting niche 
preferences (a subterranean Crematogaster species on Cyprus, an 
arboreal species in Oman). Thus, caterpillars of either population might 
have evolved local host preferences, which are probably correlated with 
the prevailing ecological conditions. Geographical “host races” with 
respect to the ant species used for larval development have recently 
been described from another obligate myrmecophile, Maculinea alcon 
(Elmes et al., 1994). 


The apparent differences in habitat and host ants may corroborate 
the idea that Cypriot and Arabian populations of C. acamas deserve 
the status of valid subspecies. Originally, all subspecies around C. 
acamas have been exclusively defined on the grounds of adult 
morphology, while ecological or life-history characters still remain 
largely unknown. 


Larsen (1983, 1990) and Larsen & Pittaway (1982) arranged the de- 
scribed taxa in a more reasonable zoogeographical manner. However, 
the subspecific taxonomy of C. acamas remains unsatisfactory. Ssp. 
cypriaca Riley, 1925 is probably best regarded as a synonym of 
nominotypical acamas Klug, 1834 (Type locality : “Syria”) which occurs 
in Asia minor and the eastern Mediterranean (from southern Turkey 
eastwards to Iran, southwards across Syria, Lebanon, Israel, East 
Jordan and the Sinai to Egypt and probably into northern Arabia : 
Larsen 1974, 1990: Larsen & Nakamura, 1983 ; Benyamini, 1990). In 
south-western Arabia there occurs C. acamas bellatrix Butler, 1886, 


72 


while C. acamas hypargyros Butler, 1886 ranges from Oman eastwards 
across Iran and Pakistan to Afghanistan and India. 


Nevertheless, given the extremely localized and fragmented distribution 
in relation to the arid environments and strong dependence on its host 
ants, the applicability of the traditional subspecies concept in a case 
like €. acamas appears doubtful. 


According to Larsen & Pittaway (1982), C. acamas and its congenors 
(like C. myrmecophila) are eremic species confined to arid environments, 
although nominotypical C. acamas also extends into the Mediterranean 
biome. At least these two species and probably many more members 
of the tribe Aphnaeini (e.g. the numerous local endemics of the genera 
Poecilmitis or Aloeides in southern Africa) are ant-dependent low 
density species which only occur at very restricted locations. Ant- 
dependence in obligate myrmecophiles like Cigaritis might have par- 
ticularly affected speciation processes, possibly leading to a great 
number of environmentally differentiated entities to which traditional 
taxonomic concepts can hardly be applied. Especially in xeric envi- 
ronments, where suitable habitats (“oases”) are patchily distributed, 
small in size, and often highly isolated, local specializations with regard 
to myrmecophily could be a driving force of evolution (Pierce, 1984 ; 
Smiley et al., 1988). Hence, some of the taxonomic difficulties within 
Cigaritis could be influenced by this highly specialized way of life. 
Furthermore, much of the variation in size and colour observed between 
usually small samples from widely distant localities may be due to 
the extremely harsh and variable environmental conditions (unpredic- 
table rainfall, resource competition inside ant nests). 


To approach a more complete understanding of the taxonomy and 
evolution of Cigaritis butterflies, but also to fill the still significant gaps 
in the mere documentation of the life-cycles of these interesting butterfly 
species with their spectacular biology, we hope that this report 
encourages a larger number of lepidopterists as well as myrmecologists 
to pay attention to these highly advanced offshoots from the Ethiopian 
fauna, when studying natural history at the southern fringe of the 
Palearctic realm. 


Acknowledgements 


We thank A. Gerber (Würzburg) for her assistance with the SEM and Dr. 
R. D. Schumann (Darmstadt) for his help attempting to rear caterpillars with 
Leptothorax ant brood. A. Schulz (Leverkusen) kindly provided the deter- 
mination of the Crematogaster species. 


74 


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76 


Nota lepid. 18 (1) : 77-92 ; 31.1.1996 ISSN 0342-7536 


Uniformity of wing spotting of Maniola jurtina (L.) 
in relation to environmental heterogeneity 
(Lepidoptera : Satyrinae) 


T. G. SHREEVE*, R. L. H. Dennıs** & W. R. WILLIAMs*** 


*School of Biological & Molecular Sciences, Oxford Brookes University, Headington, Oxford 
OX3 OBP, U.K. 

**Department of Biological Sciences, Keele University, Staffordshire STS 5BG, U.K. 

*** Computing Service, Science Laboratories, University of Durham, South Road, Durham DH! 
SEE UK. 


Summary 


Ventral hindwing spotting and wingspan of the meadow brown butterfly 
Maniola jurtina (L.) were measured on samples collected from 10 sites in 
North Wales which differ in climate, geology and habitat structure, known 
to produce significant diversity in another satyrine butterfly Hipparchia semele 
(L.). For both sexes of Maniola jurtina spotting was uniform over all sites, 
but size differed significantly. These results do not conform to models which 
relate hindwing spotting to weather or to habitat related activity differences 
and selection. Data on movements from another location, Brereton Heath, 
Cheshire, are used to suggest that the observed uniformity of spotting may 
partly result from inter-site mobility, preventing divergence in spot patterns 
between sites. In addition, colonisation history and continuously changing 
patterns of habitat availability in North Wales may limit temporal and spatial 
isolation of populations of M. jurtina. Differences in size are most likely related 
to environmental effects during larval development. 


Résumé 


Les auteurs ont mesuré les ocelles du dessous des ailes postérieures et 
l’envergure du Satyride «Le Myrtil», Maniola jurtina (L.), dans des séries 
récoltées dans le nord du Pays de galles, a 10 endroits dont on savait que 
leurs différences de climat, de géologie et d’habitat étaient à l’origine d’une 
diversité significative chez un autre Satyride, «L’Agreste», Hipparchia semele 
(L.). Chez les Maniola jurtina de tous ces endroits, les ocelles des deux sexes 
avaient la même forme, mais les envergures présentaient des différences 
significatives. Ces résultats ne concordent pas avec les règles qui établissent 
une relation entre le dessin des ocelles des ailes postérieures et le temps ou 
des différences d’activité et de sélection selon l’habitat. Les données sur les 
mouvements du Myrtil observés dans une autre station — Brereton Heath, 


Vi 


Cheshire — sont présentées pour suggérer que l’uniformité des ocelles constatée 
pourrait être en partie le résultat de la mobilité des Myrtils entre les localités : 
elle préviendrait la divergence dans l’aspect des ocelles selon l’habitat. Par 
ailleurs, la colonisation et les changements continus des types d’habitat 
disponibles dans le nord du Pays de Galles pourraient réduire l’isolation des 
populations de M. jurtina dans l’espace et dans le temps. Quant aux différences 
de taille, elles sont le plus vraisemblablement dues aux influences de 
l’environnement pendant le développement de la chenille. 


Zusammenfassung 


Untersucht wurden das Fleckenmuster der Hinterflügel-Unterseiten und die 
Flügelspannweite des Ochsenauges Maniola jurtina. Das Material stammte 
von 10 Fundorten in Nordwales, die sich in Klima, Geologie und Habitat- 
struktur unterscheiden. Letzteres führt bei einer anderen Satyrine, nämlich 
Hipparchia semele, zu einer signifikanten Diversität. Bei beiden Geschlechtern 
von Maniola jurtina war das Flügelmuster an allen Fundorten gleichförmig, 
die Größe dagegen signifikant unterschiedlich. Diese Ergebnisse stehen im 
Widerspruch zu Modellen, die das Hinterflügel-Muster mit dem Wetter oder 
mit habitatbedingten Aktivitätsunterschieden und Selektion korrelieren. Unter 
Berücksichtigung von Daten über Lokomotionsaktivitäten von einem anderen 
Fundort (Brereton Heath, Cheshire) wird vorgeschlagen, daß die beobachtete 
Gleichförmigkeit des Fleckenmusters teilweise daraus resultiert, daß zwischen 
den einzelnen Fundorten ein Individuenaustausch stattfindet. Dies verhindert 
Unterschiede im Fleckenmuster zwischen den Fundorten. Außerdem begrenzen 
die Besiedlungsgeschichte und das sich ständig verändernde Angebot an 
Habitaten in Nordwales die zeitliche und räumliche Trennung der Populationen 
von M. jurtina. Größenunterschiede sind höchstwahrscheinlich durch Umge- 
bungseinflüsse während der Larvalphase bedingt. 


Introduction 


Populations of the meadow brown Maniola jurtina (L.) (Lepidoptera, 
Satyrinae) may differ in marginal hindwing spotting, size, emergence 
time and emergence period (Brakefield, 1984 ; Brakefield & Shreeve, 
1992 ; Dennis, 1977). Such differences are not universal, with geographic 
patterns of uniformity being recognised throughout the geographic 
range of this butterfly (Dowdeswell & McWhirter 1967; Owen & 
Smith, 1993). Morphological differences which occur are described as 
the product of current selection associated with habitat structure and 
climate (Brakefield, 1984 ; 1987) or result from past events associated 
with changes in distribution and range during Pleistocene polyglaciation 
(Dennis, 1977 ; Shreeve, 1989). Differences of peak emergence times 
have been related to temperature experienced during development 
(Brakefield, 1987) whilst the duration of the emergence period has been 


78 


related to environmental heterogeneity and local adaptation of pop- 
ulations derived from a single ancestral population (Brakefield, 1987 ; 
Thomson, 1971; Dennis, 1971) or from more than one ancestral 
population (Dennis, 1977 ; Shreeve, 1989). 


Within any population of Maniola jurtina there may be seasonal 
changes of morphological characters. Typically there are decreases of 
mean spot number, of the proportion of individuals with costally 
positioned spots and of wing expanse during the flight season (Creed, 
et al., 1962; Brakefield, 1984; 1987). Such changes, which are not 
universal (e.g. Shreeve, 1989), are linked to differential development 
rates and pleiotropic effects. 


Inter-population variation of marginal hindwing spotting in Maniola 
jurtina has been related to temperature, activity and predator evasion 
(Brakefield, 1984). It is argued that hindwing spots are emphasised 
in warm habitats since they can fully function as deflective characters 
in secondary defence when individuals are capable of effective escape 
flights. In cool habitats emphasis is placed on reduced spotting and 
crypsis since hindwing spots cannot act deflectively if low temperatures 
preclude flight activity and therefore escape. The model of Brakefield 
also relates spotting to habitat heterogeneity and to differences between 
male and female activity. Hindwing spotting is reduced and more 
uniform in populations of homogeneous habitats than those of 
heterogeneous habitats because fewer spot patterns will be successful 
in the former. In any habitat males will be more prominently spotted 
than females since emphasis is on deflection in males as they engage 
in more flight activity and on crypsis in the less active female. 


Since hindwing spotting is heritable (Brakefield & van Noordwijk, 
1985), variation between populations within any geographic area will 
be related to differences in selection pressures between populations and 
gene flow between these populations. In this paper we examine 
hindwing spotting and size of Maniola jurtina from a variety of sites 
in North Wales. Observations on Hipparchia semele (L.) (Lepidoptera, 
Satyrinae) at these sites have disclosed marked significant variation 
between populations. Individuals of both sexes from the Great Orme’s 
Head are significantly dwarfed, have fewer and smaller wing spots and 
emerge earlier than butterflies from other sites. Individuals from sand 
dune habitats are significantly larger in wing expanse and have greater 
wing spotting than those from other lithologies, with the exception 
of those from dune populations adjacent to the Great Orme’s Head, 
which apparently have been influenced by gene flow from populations 
on the Great Orme’s Head (Dennis, 1972 ; 1977 ; unpubl. data). There 


79 


is therefore the expectation that Maniola jurtina from the same or 
similar sites will show interpopulation variation in response to similar 
environmental factors. 


Materials and methods 


To compare spotting and wing expanse of populations from different 
areas, samples of adults were collected during the peak flight period 
from 10 different sites in north Wales (Figure | and Table 1), many 
of which demonstrate heterogeneity in Hipparchia semele. For most 
sites samples of adults were collected over several days during the peak 
flight periods (mid-July) of the years 1969-1972. Individuals from 
Aberffraw were collected in mid-August 1972. Hindwing spots were 
recorded using xl0 or x30 magnification and wing expanse was 
recorded as mid-thorax to forewing-tip length to 0.5 mm. using a scale 
rule. Spot number rather than spot area was measured because the 
former provides a simpler measure of the total spot area and spot 
number and position have been used to characterise populations in 
previous studies (see Brakefield, 1984). 


Habitat heterogeneity was recorded by summing the number of 
different types of vegetation structures within the areas used by Maniola 
jurtina. Climatic data for the sample sites were not obtained directly 
but April-June mean temperature and July-September mean temper- 
ature and daily hours of sunshine are calculated for each site using 
the formulae given by White & Smith (1982). The first climatic variable 


Great Orme 

Little Orme 
© 
Deganwy 
© 
e Colwyn „Dulas Valley 
Conway//Pydew Bay 
Morfa 


Aberffraw 


G 


~ Dolgarrog 


Fig. 1. The location of sample sites for Maniola jurtina in North Wales. 


80 


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has a major influence on the timing of emergence (Brakefield, 1987), 
the others on the activity of adults during the flight season. 


Intra- and inter-habitat movement data were collected, using mark- 
release recapture (MRR) techniques, during the period June 30 to July 
29 1987, at Brereton Heath Country Park, Holmes Chapel, Cheshire. 
Intensive data on mobility could not be collected in North Wales 
because of the distances between the sites and also because development 
has eliminated some of the sites since the original samples were 
collected. At the Cheshire site individuals were sampled along a fixed 
transect route which covered the entire area, including 14 habitat zones 
and areas unsuitable for the butterfly. Habitats (Figure 2) could be 
clearly identified from demarcations in vegetation structure and the 
location of egglaying and mating individuals. Population estimates were 
made using the method of Jolly (Begon, 1979). 


Fig. 2. The location of fourteen population units of Maniola jurtina at Brereton Heath, 
Cheshire. 


82 


Geographic patterns of spotting and size 


For both sexes, spot pattern, spot combinations, mean spot number 
and costality indices are shown in Table 2a and 2b (see Brakefield 
& Shreeve, 1992 for spot pattern notation). There are no significant 
differences between samples from different sites in the total number 
of hindwing spots (males, x? (9) = 11.04, P>0.2; females, X? (13) = 
20.53, P > 0.3), nor for individual wing spots (spot 1: males, x? (9) 
RP PP 00 females, 25 609 2 > 0:3"; spot 2); females. 720 
aeties 0:5, spot 5 females; y7 4) — 12-45, Pe 0al ; spot 6: 
idles) — 10.85, 20.3; females, y* (4, = 12.45, PZ 01 ; other 
spots were either almost universally present or absent). For all sites, 
male spot number is unimodal at 2 spots and the commonest phenotype 
is splay 2; and in females, spot number is unimodal at 0 spots, the 
commonest phenotype being spot absence. 


Sixty-four spot combinations are theoretically possible from the six 
hindwing spot locations ; 15 were identified in North Wales and most 
were infrequent (Table 2a and 2b). Comparisons between sites for spot 
combinations (males, 010010 versus remainder, x? (9, = 14.25, P > 0.1 ; 
females, 000000 and 010000 versus remainder, x? (18 = 23.66, P > 0.1) 
also indicate homogeneity, as did a comparison between sites for costal- 
anal bias in spotting (individuals were scored as to sign, spots being 
registered as positive if costal, neutral if medial and negative if anal ; 
males — »-04) 2.018; females, y2, — 12:54, 201) Mult- 
variate analyses of sites for frequencies of spot combinations revealed 
no conformity for sex, geographic location, habitat type or climate, 
contrasting with observations of Hipparchia semele (see Dennis, 1972, 
S75). 


Mean wing expanse of males and females is shown in Table 3. For 
both sexes there are differences of mean wing expanse between sites 
(males, F 188, — 5.38 : females, Fo 190, = 4.65 ; P< 0.001 in both cases). 
The largest individuals are from Gloddeath and the smallest from 
Aberffraw. Mean size is not correlated with mean spot number in either 
sex (males, r, = 0.12, females r, = 0.261, N.S. in both cases). With the 
exception of female costality, mean spot number, costality and size 
of both sexes are not correlated with temperature (April-June) during 
the relevant developmental period, or temperature and sunshine hours 
during the flight period (July-September). Female costality is correlated 
with mean July-September daily hours of sunshine (r, = 0.79, P< 0.01). 


83 


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Table 3 


Mean wingspan of males and females of Maniola jurtina from ten sites in North 
Wales 


Mean Wingspan (mm) + s.e. 
l Female 


Great Orme 24.9 + 0.23 26.4 + 0.31 
Little Orme 24.5 + 0.20 WAGs3) ae (AU) 
Gloddaeth 27.5 + 0.28 
Pydew Oj a2 (Ds) 
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Mobility 


The distribution of habitats on Brereton Heath is shown in Figure 
2. The total population size rose to a peak of 163 males on July 12 
and 161 females on July 22. Mean residence time for males is 9.7d 
and for females 7.2d. Some 24.7 per cent (118 of 761) of movements 
were between the 14 habitat patches ; this together with the pattern 
of cross movements (Figure 3) indicates that individuals on the heath 
belong to a single population. The pairwise frequency of cross- 
movements can be closely matched by an orthodox gravity model 
(r, = 0.69, P< 0.001) : 
P; 
Mi D, Equation | 

where M is the expected interactions, P is population size, and D is 
the direct distance between the populations ; the number of unique 
captures per habitat unit is used as a surrogate for population size. 
Habitat areas differ substantially in size, isolation, relative population 
size and density (Table 4), factors which can influence the ratio of 
between to within habitat movements. Out-movements, as a fraction 
of within habitat movement, are much greater from the ten small 
populations than from the four largest habitats (r, = 0.67, P< 0.01). 
Out-movements also correlate negatively with population size (r, = - 
0.83, P < 0.001), population density (r, = -0.70, P< 0.01) and habitat 
area (r, = -0.65, P< 0.01). 


85 


Movements into habitats can be modelled as part of the gravity 
equation (r, = 0.62, P < 0.05) : 


N P e 
ED) =i Equation 2 
Seige D; 


where I is the index of input into an area. A better predictor of in- 
movements is population size (r, = 0.94, P < 0.01), but this relationship 
is influenced by the correlation of habitat size with the geographic 
parameters of neighbours in Equation 2. At Brereton Heath large 
habitats tend to be nearest to other large habitats (r, = 0.53, P< 0.05), 
but there may also be a tendency for immigrants into larger more 
densely populated habitats to stay longer than those in sparsely 
populated habitats, and thus have a higher probability of being recorded 
during transect MRR events. 


III 


>> 


Fig. 3. Combined recorded movements of males and females between population units 
at Brereton Heath, Cheshire. 


86 


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Movements of the butterfly on Brereton Heath indicate that it is capable 
of flying considerable distances in a short period of time and of crossing 
a range of obstacles including mown grass, open water and woodland. 
Only 2.8 per cent (39 of 1392) of all captures and only 3.6 per cent 
(27 of 761) of captures involving recorded movements were of ex- 
habitat individuals. When compared to the number engaged in cross- 
movements between habitats (24.7 per cent), this demonstrates that 
movements between habitats must be rapid. Recorded distances moved 
by the butterfly will depend on study area dimensions and for how 
long individuals can be traced or followed. The mean maximum range 
of individuals at Brereton Heath for those captured at least three times 
is for males 158.2 + SE 12.7 m (n= 130) and for females 118.3 = SE 
14.4 m (n = 66). Some 25.5 per cent of individuals (50 of 196) recorded 
linear ranges greater than the mean distance between habitats (204 m) 
and 4.6 per cent greater than the furthest neighbour distance (429 m). 
96.4 per cent achieved linear transfers greater than the mean nearest 
neighbour distance (19 m). 


Discussion 


Models of spotting in Maniola jurtina (Brakefield, 1984) and in other 
Satyrine butterflies, e.g. Coenonympha tullia (Dennis, Porter & Wil- 
liams, 1986), lead to expectations of spot pattern differences between 
populations from sites in North Wales which differ in climate, lithology 
and habitat structure. No differences were detected. The distinctive 
patterns of morphological variation in Hipparchia semele (Dennis, 
1972, 1977) were not repeated in Maniola jurtina. It is unlikely that 
this lack of variation is a simply a result of the sampling methodology 
. If the samples came from populations with true spotting differences 
then confounding effects may result from sampling times and unknown 
seasonal changes of spot number within populations. If mean spot 
number declines with season then there is an expected negative correla- 
tion of spot number with temperature experienced during development 
and if mean spot number increases with season there is an expected 
positive correlation. No such correlations were found. Non-significant 
correlations of spotting attributes with any climatic variables can be 
most easily explained by proposing that there are no differences in 
the overall spotting patterns of the sampled populations. Wing expanse 
differs between sites but there are no consistent trends that can be 
related to site characteristics, as in the case of Hipparchia semele, 
although the relatively late time of sampling may account for the small 
size of individuals from Aberffraw. 


88 


We think that there are interesting implications within these data. Wing 
spotting, but not size, is consistent over a wide range of habitat types. 
Three possible explanations can be given. The first is that selection 
for spotting is the same in all habitats. The second is that these spots 
are neutral in their effect. The third hypothesis is that individuals of 
Maniola jurtina are sufficiently mobile to prevent stabilisation of 
alternative spot patterns in different habitat patches. With the present 
data none of these three hypotheses can be rejected. However, the first 
is unlikely because the differences between the sites are sufficiently large 
to expect differences of flight activity and requirements for the 
expression of secondary defence mechanisms. The second hypothesis 
may also be unlikely because predator attacks are directed at hindwing 
spot positions in samples of Maniola jurtina from other areas (Bengtson, 
1981 ; but see Shreeve, 1989). Although data on mobility presented 
here is from a different area to North Wales the observed patterns 
of mobility demonstrate that the butterfly has a substantial capacity 
to move between habitat areas. Observations from Brereton Heath 
provide results intermediate between those of previous studies that have 
primarily considered intra-habitat (Brakefield, 1982) and inter-habitat 
movements (Baker, 1984). They contradict observations by Ford (1975) 
for movements between habitats on the Isles of Scilly and confirm 
those of Baker (1984) that the butterfly is a highly mobile species. 
Movements within the pattern of habitat patches at Brereton Heath 
suggest that the contraction and isolation of habitats, as has been 
occurring in North Wales, may encourage migration (sensu Baker, 1978) 
rather than lead to its reduction. Ground unsuitable for the reproduction 
and maintenance of the butterfly G.e. mown grass, dense woodland, 
open water, urban structures) are not obstacles to movement. Corre- 
lations between habitat area, population size and density, out-move- 
ments and in-movements indicate that the greatest determinant of when 
to stay as opposed to when to leave is the presence of other individuals. 
Larger denser populations encourage individuals to stay within habitats. 
Nevertheless, there is a continual leakage of individuals from the largest 
populations which may be expected to generate diversity for dispersal 
genes. 


Additional, circumstantial evidence for vagility in M. jurtina is provided 
by analysis of allozyme variation in south-east England (Goulson, 
1993a ; 1993b). Low mean Fsr values (0.015 - 0.023) point to little 
divergence between geographically separated locations and are consistent 
with values for other highly vagile insects whose populations are 
effectively panmictic. 


89 


Variation in size of individuals between the different sample sites cannot 
be explained by conventional models. Spot size and number are usually 
related to wing size, large individuals tend to have more and larger 
spots (Brakefield, 1984). Surprisingly there is no correlation between 
size and spot number for the North Wales samples. An explanation 
is that for the sampled populations immediate environmental effects 
(which may not be shown in averaged climatic data) contribute more 
to size variation than do any other effects. The absence of variation 
in size to match the disruptive geographic pattern in Hipparchia semele 
probably relates to the different habitats occupied by the two species 
and differences in their colonisation history within North Wales. 
Maniola jurtina does not frequent the scree slopes occupied by A. 
semele and would not have colonised the present habitats until forest 
clearance (<< 5ka B.P.). Unlike A. semele it would not have a history 
of isolation on the Great Orme extending back to c. 9ka B.P. (Dennis, 
1992). As habitats for A. semele are relatively stable compared to those 
of M. jurtina, which is dependent on changing patterns of cultivation 
and grazing, M. jurtina populations probably share a more recent 
colonisation history and probably have a history of repeated local 
extinction and recolonisation, the latter aided by their mobility. 


In conclusion, the results presented here demonstrate homogeneity of 
spot pattern between sites despite climatic and structural differences 
between sites. Spotting is similar to that over much of the range of 
the species. North Wales populations are similar to others and cor- 
respond to the general southern and central spot stabilisation of Britain 
(Dowdeswell & McWhirter, 1967 ; Dowdeswell, 1981). We suggest that 
in North Wales at least, inter-site mobility may largely preclude diffe- 
rentiation in climatically dissimilar sites and that differentiation of 
Maniola jurtina observed elsewhere will only occur when populations 
have been effectively isolated. Instability in habitats, shifting locations 
and recent colonisation history in North Wales have thwarted the 
evolution of size and hindwing spot pattern variation in Maniola 
jurtina. Differences between sites in adult size, are, we suggest, the 
result of the interaction of developmental rate and site temperature ; 
warm sites promote development and allocation of resources to body 
mass. 


Acknowledgements 


We thank Derek Whiteley for providing the figures and Andrea Turner for 
providing the French abstract. 


90 


References 


Baker, R. R., 1978. The Evolutionary Ecology of Animal Migration. Hodder 
and Stoughton, London. 

Baker, R. R., 1984. The dilemma: when and how to go or stay. In Vane- 
Wright, R. I. & Ackery, P. R. (Eds) The Biology of Butterflies. Symp. 
R. Ent. Soc. No. 11 : 279-276. Academic Press, London. 

Becon, M., 1979. Investigating Animal Abundance. Edward Arnold, London. 

BENGTSON, S. A., 1981. Does bird predation influence spot-number variation 
in Maniola jurtina ? Biol. J. Linn. Soc. 15 : 23-27. 

BRAKEFIELD, P. M., 1984. The ecological genetics of quantitative characters 
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P.R. (Eds) The Biology of Butterflies. Symp. R. Ent. Soc. No. 11 : 167- 
190. Academic Press, London. 

BRAKEFIELD, P. M., 1987. Geographic variability in, and temperature effects 
on, the phenology of Maniola jurtina and Pyronia tithonus (Lepidoptera, 
Satyrinae) in England and Wales. Ecol. Entomol. 12 : 139-148. 

BRAKEFIELD, P. M. & Noorpwisk, A. J. van, 1985. The genetics of spot 
pattern characters in the Meadow Brown butterfly Maniola jurtina 
(Lepidoptera : Satyridae). Heredity 54 : 275-284. 

BRAKEFIELD, P. M. & SHREEVE, T. G., 1992. Case studies in Evolution. Jn 
Dennis, R.L.H (Ed.) The Ecology of Butterflies in Britain. Oxford 
University Press, Oxford, pp. 197-216. 

CREED, E. R., DOWDESWELL, W. H., Forp, E. BB & McWhirter, K. G., 
1962. Evolutionary studies on Maniola jurtina : the English Mainland, 
1958-60. Heredity 17 : 237-265. 

Dennis, R. L. H., 1971. A model for temporal subspeciation. Ent. Rec. J. 
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Dennis, R. L. H., 1977. The British Butterflies. Their Origin and Establishment. 
E.W.Classey, Faringdon. 

Dennis, R. L. H., 1992. An evolutionary history of British butterflies. In 
Dennis, R. L. H. (Ed.) The Ecology of Butterflies in Britain. Oxford 
University Press, Oxford, pp. 217-245. 

Dennis, R. L. H., Porter, K. & WırLıams, W. R., 1986. Ocellation in 
Coenonympha tullia (Müller) (Lep. Satyridae) : II. Population differ- 
entiation and clinal variation in the context of climatically-induced anti- 
predator defence strategies. Entomologist’s Gaz. 37 : 133-172. 

DowDEswELL, W. H., 1981. The Life of the Meadow Brown. Heinemann, 
London. 

DowDEswELL, W. H. & McWhirter, K.,1967. Stability of spot distribution 
in Maniola jurtina throughout its range. Heredity, 22 : 187-210. 

Forp, E. B., 1975. Ecological Genetics. (4th ed.). Chapman and Hall, London. 

Goutson, D., 1993a. Allozyme variation in the butterfly, Maniola jurtina 
(Lepidoptera, Satyrinae) (L.) — evidence for selection. Heredity 71 : 386- 
398). 


91 


Goutson, D., 1993b. The evolutionary significance of bimodal emergence 
in the butterfly, Maniola jurtina (Lepidoptera : Satyrinae) (L.). Biol. J. 
Linn. Soc. 49 : 127-139. 

Owen, D. F. & Smiru, D. A. S., 1993. Spot variation in Maniola jurtina 
(L.) (Lepidoptera : Satyridae) in southern Portugal and a comparison 
with the Canary Islands. Biol. J. Linn. Soc. 49 : 355-365. 

SHREEVE, T. G., 1989. The extended flight period of Maniola jurtina 
(Lepidoptera, Satyridae) on chalk downland : seasonal changes of adult 
phenotype and evidence for a population of mixed origins. Entomologist 
108 : 202-215. 

THoMsoN, G., 1971. The possible existence of temporal subspeciation in 
Maniola jurtina (L.). Ent. Rec. J. Var. 83 : 87-90. 

WHITE, E. J. & Smirn, R. I., 1982. Climatological Maps of Great Britain. 
Institute of Terrestrial Ecology, Cambridge. 


92 


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Nota lepidopterologica 


Vol. 18 No. 2 Basel, 29.11.1996 ISSN 0342-7536 


Editor : Steven E. Whitebread, Maispracherstrasse 51, P.O. Box 27, CH-4312 Magden, 


Switzerland. FAX : +41-61-8412238. 


Assistant Editors : Emmanuel de Bros (Binningen, CH) 
Dr. Roger Dennis (Wilmslow, GB) 
PD Dr. Andreas Erhardt (Binningen, CH) 
Dr. Yves Gonseth (Neuchatel, CH) 
Dr. Alexander Pelzer (Wennigsen, D) 


Contents — Inhalt — Sommaire 


CuUPEDOo, F.: Die morphologische Gliederung des Erebia melampus- 
Komplexes, nebst Beschreibung zweier neuer Unterarten: Erebia 
melampus semisudetica ssp.n. und Erebia sudetica belledonnae ssp.n. 


(Sabyiidac) mm ee ee ne 95 
MAIER, C. & SHREEVE, T. G. : Endothermic heat production in three 

SPECIEKORNymphalidae 0 000 eee 127 
PASSERIN D’ENTREVES, P. : Révision des Scythrididae paléarctiques. VI. 

Les types de Scythrididae du Natural History Museum de Londres, 

du Muzeul de Historia Naturala «Grigore Antipa» de Bucarest et 

du Zoologisches Museum der Humboldt-Universität de Berlin (Pre- 

INES DES) Ne 139 
TENNENT, J. : The distribution of the genus Tarucus Moore, [1881], in 

the Maghreb States of Morocco, Algeria and Tunisia, with notes 

onsspeeiesidentiiication (Eyeaenidae) ..  . . .......... 161 
Short communication — Kurze Mitteilung — En bref 
L6BEL, H. & DRECHSEL, T. : Zur Lebensweise der Raupe von Euchalcia 

emichi (Rogenhofer, 1873) in Zentralanatolien (Türkei) (Noctuidae, 

Poste) we ER a a cea ats 179 
Book reviews — Buchbesprechungen — Analyses ............. 138, 160 
<i Eunopean Congress of Lepidopterology 3.0. 94 
DD itekaUAMIBORG iin NOS f= 1999) 5k 0. Le. sas kB goa deen ne 126 
Tage ee x 2... re 178 


C2) 


Nota lepid. 18 (2) : 94 ; 29.11.1996 ISSN 0342-7536 


Xth European Congress of Lepidopterology 


Miraflores (Madrid, Spain) 
3-7th May 1996 


The Council of the Societas Europaea Lepidopterologica (SEL) kindly 
invites all lepidopterists to attend the Xth European Congress of 
Lepidopterology to be held in Miraflores (Madrid, Spain) from 3rd 
to 7th May 1996. The site of the Congress is the University Hostel 
“La Cristalera”, at Miraflores de la Sierra, which is situated in the 
Sierra de Guadarrama, about 50 km north of the city of Madrid. 


Main topics will be : 


General and specific problems of taxonomy and evolution 
Ecology, endangered species and species protection 
Holarctic zoogeography and local faunistics 

New discoveries in Microlepidoptera 

Tropical Lepidoptera 

Applied lepidopterology 


For more details please contact : 


Xth European Congress of Lepidopterology, 
Department of Biology, 

Universidad Autonoma, 

E-28049 Madrid, 

Spain 


Tel. : +34/ 1-3978281 
Fax : +34/ 1-3978344 


94 


Nota lepid. 18 (2) : 95-125 ; 29.11.1996 ISSN0342-7536 


Die morphologische Gliederung des Erebia melampus- 
Komplexes, nebst Beschreibung zweier neuer Unterarten : 
Erebia melampus semisudetica ssp.n. und Erebia 
sudetica belledonnae ssp.n. (Lepidoptera, Satyridae) 


Frans CUPEDO 


Processieweg 2, NL-6243 BB Geulle, Niederlande 


Summary 


A morphological analysis of the Erebia melampus complex, with the descrip- 
tions of two new subspecies : Erebia melampus semisudetica ssp.n. and Erebia 
sudetica belledonnae ssp.n. (Lepidoptera, Satyridae) : A quantitative analysis 
of the differences between Erebia melampus (Fuesslin, 1775) and Erebia 
sudetica Staudinger, 1861 reveals the existence of three morphologically 
different groups within the E. melampus complex, referred to as the sudetica 
group, the melampus group and the momos group. Statistically, the differences 
are highly significant. The momos group is morphologically largely intermediate 
between that of sudetica and melampus, but in some characters it is extremely 
close to sudetica. A short series, however, is sufficient for a clear diagnosis. 
Typical momos Fruhstorfer, 1910 are dark, and poorly marked. There exist, 
however, brightly coloured and heavily marked populations, which are hardly 
distinguishable from sudetica without examination of the genitalia (ssp. 
semisudetica ssp.n.). E. sudetica has its main distribution outside the Alps. 
Alpine populations are found in the Bernese Alps (ssp. inalpina Warren) and 
in the French Alps (ssp. belledonnae ssp.n.). There is no evidence of any 
sympatric occurrence of sudetica and melampus, nor of sudetica and momos. 
E. melampus is confined to the western Alps, momos to the eastern Alps. 
Where they meet there is, at least in the Ortler Alps, a relatively narrow 
zone of intermixing. This situation is typical for taxa coming into secondary 
contact after a period of (glacial) isolation and differentiation. Hence it follows 
that the differentiation of the momos-group took place at the latest during 
the Würm glaciation. For lack of evidence on sympatric occurrence or hybri- 
disation it is impossible to conclude on the degree of speciation, and thus 
on the systematic rank of the taxa concerned. Therefore, nomenclature has 
been left as it was, the subspecies of the momos-group being handled as if 
they were subspecies of melampus, and sudetica as if it were a distinct species. 


Zusammenfassung 


Aus einer quantitativen Analyse der klassischen Unterscheidungsmerkmale von 
Erebia melampus (Fuesslin, 1775) und Erebia sudetica Staudinger, 1861 geht 


95 


hervor, daß der E. melampus-Komplex sich in drei morphologische Gruppen 
gliedern läßt, die als sudetica-Gruppe, melampus-Gruppe und momos-Gruppe 
bezeichnet werden. Die Unterschiede sind statistisch in hohem Maße signifikant. 
Die momos-Gruppe hält morphologisch die Mitte zwischen den beiden ande- 
ren. In einigen Merkmalen steht sie sudetica sehr nah. Anhand einer kleinen 
Stichprobe ist sie jedoch eindeutig zu erkennen. Typische momos Fruhstorfer, 
1910 sind düster und arm gezeichnet. Es gibt aber Populationen die so lebhaft 
und reich gezeichnet sind, daß sie nach ihren Flügelmerkmalen kaum von 
sudetica zu unterscheiden sind (semisudetica ssp.n.). Die Verbreitung von 
sudetica ist größtenteils extra-alpin, mit alpinen Enklaven in den Berner Alpen 
(ssp. inalpina Warren) und in den französischen Alpen (ssp. belledonnae 
ssp.n.). Gegenüber melampus und momos ist sie völlig allopatrisch. E. melam- 
pus ist auf die Westalpen, momos auf die Ostalpen beschränkt. Wenigstens 
an einer Stelle (Ortler-Alpen) hat sich eine schmale Mischzone gebildet. Die 
Situation ist typisch für einen sekundären, postglazialen Kontakt nach einer 
(glazialen) Isolation. Die Differenzierung der momos-Gruppe muß also 
spätestens während des Würm-Glazials stattgefunden haben. Da es keine 
Angaben über Sympatrie oder Hybridisation gibt, ist der spezifische Diffe- 
renzierungsgrad der drei Gruppen nicht festzustellen und ist eine taxonomische 
Einstufung grundsätzlich unmöglich. Deshalb wird nomenklatorisch jetzt 
nichts geändert. Die Unterarten der momos-Gruppe werden nomenklatorisch 
als Unterarten von melampus, sudetica dagegen als eine distinkte Spezies 
behandelt. 


Resume 


L’analyse statistique des caractères distinctifs entre Erebia melampus (Fuesslin, 
1775) et Erebia sudetica Staudinger, 1861 révèle qu’il existe en fait trois groupes 
morphologiques dans le complexe d’E. melampus s.l., nommés respectivement 
«groupe sudetica», «groupe melampus» et «groupe momos». Ce dernier est 
largement intermédiaire entre melampus et sudetica ; d’après quelques carac- 
tères alaires il est très proche de sudetica. En examinant un petit nombre 
d’individus, on arrive cependant a une détermination certaine. Les momos 
Fruhstorfer, 1910 typiques sont d’une couleur foncée, tandis que le dessin 
des ailes est relativement réduit. Mais il existe dans ce groupe des populations 
à couleur vive, aux ailes bien ornées et qui d’après leur dessin se distinguent 
à peine de sudetica (ssp. semisudetica ssp.n.). E. sudetica est répandu surtout 
en dehors des Alpes, les seules populations alpines se trouvant dans les Alpes 
Bernoises (ssp. inalpina Warren) et dans les Alpes de l’Isère et de Savoie 
(ssp. belledonnae ssp.n.). Il n’y a pas de preuve de cohabitation de sudetica 
et melampus, ni de sudetica et momos. E. melampus est répandu dans les 
Alpes occidentales, momos dans les Alpes orientales. Sur la ligne de contact, 
il existe, localement du moins, une zone relativement étroite de mélange ; 
situation typique qui se présente quand deux taxa, après une période de 
différentiation en isolation (glaciaire), se rencontrent secondairement. Faute 
de preuve de cohabitation ou d’hybridation, il est impossible de déterminer 
le degré de differentiation spécifique, et par conséquence d’établir le statut 


96 


taxinomique des groupes. Pour cette raison, la nomenclature est maintenue 
inchangée, c’est-à-dire que les sous-espèces du groupes momos sont traitées 
comme si elles étaient des sous-espèces de melampus, et sudetica comme une 
espèce distincte. 


Einleitung 


Schon 1861 erkannte Staudinger, daß im Erebia melampus-Komplex 
zumindest zwei morphologische Gruppen vertreten sind, die wir jetzt 
als Erebia melampus (Fuesslin, 1775) und Erebia sudetica Staudinger, 
1861 kennen. Letztere wurde als eine geographische Varietät beschrieben, 
eine Kategorie, die man jetzt als Subspezies bezeichnet (ICZN, 1985 : 
45g). Sie wurde von Warren (1949) aus typologischen Gründen zu einer 
selbstandigen Art erhoben. De Lesse, Vertreter einer biologischen Art- 
auffassung, wies schon bald danach darauf hin, daß Warrens Argu- 
mentation aus dieser Sicht unzulänglich sei (de Lesse, 1951a,b). Trotz- 
dem wurde dessen Auffassung von späteren Autoren fast ausnahmslos 
übernommen (Forster & Wohlfahrt, 1976; Gonseth, 1987 ; Higgins, 
1975 ; Higgins & Riley, 1980; Kudrna, 1986). Das ist verständlich, 
denn man ist seitdem allgemein davon ausgegangen, daß der E. 
melampus-Komplex immer eindeutig in zwei Gruppen aufzuteilen ist 
und daß beide Taxa in den Berner Alpen sympatrisch vorkommen 
(Forster & Wohlfahrt, 1976 ; Bretherton, 1966). Damit stünde die art- 
liche Selbständigkeit von sudetica außer Zweifel. Diese beiden Annah- 
men, obwohl generell akzeptiert, sind aber nie richtig überprüft worden. 


Mittlerweile hat sich herausgestellt, daß in den Berner Alpen kein 
sympatrisches Vorkommen vorliegt (siehe weiter im Text). Bezüglich 
der eindeutigen Trennbarkeit ist schon lange bekannt, daß die spezi- 
ischen Merkmale von E. melampus in den Ostalpen weniger ausgeprägt 
sind als in den Westalpen (Hoffmann & Klos, 1914). Mehr oder weniger 
sudetica-ähnliche Merkmale machen dort eine sichere Bestimmung oft 
unmöglich. Stangelmaier (1979) stellte die Population des Nockgebietes 
(Gurktaler Alpen) sogar zu E. sudetica. Die einfache Tatsache, daß 
dies seitdem weder bestätigt noch widerlegt werden konnte, zeigt, daß 
damit der Zweifel nicht beseitigt war. 


Das Ziel der vorliegenden Arbeit ist es, die Variationsbreite der Unter- 
scheidungsmerkmale von E. melampus und E. sudetica bei möglichst 
vielen Populationen zu bestimmen, um feststellen zu können, ob die 
übliche Zweiteilung des E. melampus-Komplexes überhaupt beibehalten 
werden kann. 


DY 


Material und Methoden 
Material 


Von 1658 männlichen Tieren wurden die Flügelmerkmale gemessen ; 
von 1032 die Genitalmerkmale. Für Herkunft und Größe der Stich- 
proben siehe Abb. | und Tabelle 1. Weil viele Tiere bezüglich der unter- 


Abb. 1. Herkunft der untersuchten alpinen Populationen. Für die Bezeichnung der 
Nummern, siehe Tabelle 1. Von den mit einem * versehenen Lokalitäten wurden nur 
kleine Serien berücksichtigt, die nicht statistisch ausgewertet wurden. 


suchten Merkmale asymmetrisch sind, wurden immer die rechten Flügel 
und Valven gemessen. Die Weibchen sind außer Betracht gelassen, weil 
einige Unterscheidungsmerkmale sich nur auf männliche Tiere beziehen 
und außerdem relativ wenig Weibchen zur Verfügung standen. 


Valvenmorphologie 


Die Valven bestehen bei dieser Gruppe aus einem breiten, hohlen, proxi- 
malen Teil und einem schmalen, massiven, distalen Teil (Abb. 2). Auf 


98 


Tabelle 1. 


Herkunft und Größe der Stichprobe, und einige Meßwerte. 

Nff = Zahl der untersuchten Tiere (Flügelmerkmale). Ngen = Idem (Genitalmerkmale). 
O, = Frequenz der Ozelle in Zelle 6 der Vfl.Os. F, = Frequenz des Flecks in Zelle 
lc der Hfl.Us. V = Valvenindex (Populationsmittelwert). 

* — In der Stichprobe nur einmal vertreten. 


Lokalität | Nfl. Ngen| 0, %) F(%) V 


. Altvater, CZ, Sudeten. 

. Retezat, Romania, S. Karpaten. 

. Puy Mary, F, Zentralmassiv. 

. Grindelwald, CH, Berner A. 

. La Morte, F, Taillefer. 

. Rivier d’Allemont, F, Belledonne. 
. La Toussuire, F, Arvan. 

. Terme di Valdieri, I, A. marit. 

. Risoul, F, Cottische A. 

. Col du Lautaret, F, Dauphiné. 

. Pralognan, F, Vanoise 

. Col de Roselend, F, Beaufortin. 

. Col de la Croix, CH, Berner A. 

. Baltschiedertal, CH, Berner A. 

. Laquintal, CH, Walliser A. 

. Klewenalp, CH, Vierwaldst. A. 

. St. Moritz, CH, Engadin. 

. Solda, I, Ortler A. 

. Daone, I, Adamello. 

. Vent, Sölden, A, Otztaler A. 

. Neustift, A, Stubaier A. 

. Schnalstal, I, Otztaler A. 

. P. Pordoi, P. Gardena, I, Dolomiten. 
. Kals am Gr. Gl., A, Hohe Tauern. 
. Heiligenblut, A, Hohe Tauern. 

. Mallnitz, A, Hohe Tauern. 

. Plöckenpass, Nassfeldpass, A, Karnische A. 
. Turracher Höhe, A, Gurktaler A. 
. Schladming, A, Niedere Tauern. 

. Knittelfeld, A, Niedere Tauern. 


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der dorsalen Seite befinden sich die Zähne, oft in mehreren Reihen. 
Besonders ausgeprägt sind die der äußeren Reihe, die ich als Late- 
ralzähne bezeichne. Sie sind lateral angesetzt, spitz-kegelförmig und 
stark chitinisiert (Abb. 3). Im allgemeinen sind sie größer als die dorsal 
angesetzten Dorsalzähne. Im proximalen Valvenabschnitt nähern sie 
sich dem dorsalen Valvenrand ; dort unterscheiden sie sich nur noch 
durch ihre Form von den Dorsalzähnen. Nach Warren (1961) unter- 
scheiden sich E. melampus und E. sudetica in der Zahl, der Anordnung 
und der Länge der Valvenzähne. 


99 


Ntot. 


N prox. Ndist. 


Abb. 2. Valve von Erebia melampus s.l., schematisch. 
Erklärung im Text (Gemessene Merkmale, a-c). 


O.1mm 


Abb. 3. Valve, vergrôsserter Ausschnitt. L = Lateralzahn. 


Flügelmerkmale 


Vorderflügel und Hinterflügel tragen oberseits und unterseits eine Reihe 
von Postdiscalflecken, die auf den Vorderflügeln zu einer Längsbinde 
verschmolzen sind, und die teilweise mit einer schwarzen, nie weiß- 
punktierte Ocelle versehen sind. 


100 


Nach Warren (1949) hat E. melampus auf der Hinterflügelunterseite 
weniger Flecke als E. sudetica, und niemals eine Ocelle in Zelle 6 der 
Vorderflügeloberseite ; der Fleck in Zelle 4 der Hinterflügel ist, im 
Gegensatz zu E. sudetica, wurzelwärts verlagert (siehe Abb. 4). 


Gemessene Merkmale 


a. Die Zahl der Lateralzähne (Nlat) und der Dorsalzähne (Ndors) 
sowohl im proximalen als auch im distalen Valvenabschnitt. Daraus 
ergeben sich die Zahl der proximalen Zähne (Nprox), der distalen 
Zähne (Ndist) und die Gesamtzahl (Ntot). Siehe Abb. 2. Es wurde 
die Ratio proximal/distal (P/D-Ratio) der Lateralzähne, der Dorsal- 
zähne und aller Zähne berechnet. 


b. Die Länge der ersten 10 Lateralzähne auf der proximalen Seite, 
und ihr Mittelwert (L). Weil der Zahnansatz nicht genau festzustellen 
ist, wurde nicht die wirkliche Länge gemessen, sondern der Teil über 
dem dorsalen Valvenrand. 


c. Die Länge des proximalen und des distalen Valvenabschnitts (Vp 
und Vd, Abb. 2), aus der der Valvenindex, V = Vp.100/(Vp + Vd) 
berechnet wurde. 


d. Die auf der Vorderflügeloberseite vorhandenen Ocellen. Daraus 
ergibt sich die Frequenz jeder Ocelle in der Population (O, bis O, 
— Frequenz der Ocelle in Zelle 2 bis 6). 


Abb. 4. Hinterflügeloberseite, mit dem gemessenen Winkel alpha. 


101 


e. Die auf der Hinterflügelunterseite vorhandenen Bindenflecke. Daraus 
ergibt sich F, (Frequenz des Flecks in Zelle Ic) bis Fe. 


f. Als Maß der Wurzelwärtsverschiebung des Postdiskalflecks in Zelle 
4 wurde auf der Hinterfliigeloberseite der Winkel (alpha) gemessen 
zwischen Ader 5 und der Senkrechten auf der Verbindungslinie von 
Ocelle 3 und 4. (Sıehe Abb. 4). 


Methode 


a, b und c wurden mit einem Monokularmikroskop gemessen ; a bei 
einer Vergrößerung von 400 X. Auch die kleinen, sich überlagernden 
Zähne in der Spitzenregion der Valven können so gut gezählt werden ; 
b unter Verwendung eines Okularmikrometers, Vergrößerung 400 X ; 
c : wie b, jedoch Vergrößerung 100 X ; d und e wurden mit dem bloßen 
Auge bestimmt, und f mit einem Stereomikroskop, unter Verwendung 
eines Okulargradbogens mit Segmenten von 5°, mit durchfallendem 
Licht. Vergrößerung 20 X. Hierzu wurden die Flügel mit 96% Ethanol 
befeuchtet. 


Ergebnisse 


In Tabelle 1 sind die O,, F, und V-Werte eingetragen ; in Tabelle 2 
bis 7 die Variation der gemessenen Valvenmerkmale, und in Tabelle 
8 die des Winkels alpha. Die alpinen melampus-Populationen (8 bis 
30) sind von Westen nach Osten angeordnet. 


Folgendes ist aus den Tabellen sofort ersichtlich : 


a. Die Merkmalverteilungen (und damit ihre Mittelwerte) lassen einen 
Sprung zwischen den Populationen 19 und 20 (Ortler-Alpen und 
Otztaler Alpen) erkennen. 


b. Die Populationen 20 bis 30 unterscheiden sich nicht nur von den 
westalpinen melampus-Populationen, sondern auch von E. sudetica. 
Ganz deutlich geht das aus Abb. 5 hervor, in der die Variation der 
Valvenmerkmale und des Winkels alpha graphisch dargestellt ist, und 
aus Tabelle 9, in die die Mittelwerte und Standardabweichungen der 
gemessenen und berechneten Werte pro Gruppe eingetragen sind. 


Offensichtlich läßt sich also der E. melampus-Komplex nicht in zwei, 
sondern in drei morphologische Gruppen aufgliedern : eine sudetica- 
Gruppe, eine Gruppe westalpiner melampus-Populationen und eine 
Gruppe ostalpiner Populationen. Weil nominotypische E. melampus 
der westlichen Gruppe angehören, nenne ich diese die melampus- 
Gruppe. Die östliche Gruppe werde ich weiterhin als die momos- 


102 


Tabelle 2 


Gesamtzahl der Valvenzähne (Ntot) 
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bis bis bis bis bis bis bis bis bis bis bis bis 
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Gruppe bezeichnen (!). Erst in der Dikussion werde ich auf ihren taxo- 
nomischen Status eingehen. Zunächst soll festgestellt werden, inwiefern 
die untersuchten Merkmale voneinander unabhängig sind, inwiefern 
die Unterschiede signifikant sind, und inwiefern es möglich ist, Popu- 
lationen und Einzeltiere der drei Gruppen voneinander zu unterscheiden. 


Unabhängigkeit der Merkmale 


Von vornherein war klar, daß nicht alle untersuchten Valvenmerkmale 
unabhängig voneinander variieren. Welche Merkmale sich gegenseitig 
beeinflussen, kann aber erst hinterher festgestellt werden. 


(1) momos Fruhstorfer, 1910 ist der älteste Name, der innerhalb dieser Gruppe 
verfügbar ist. Locus Typicus : Dolomiten. 


103 


Tabelle 3 
Zahl der Lateralzähne (Nlat) 


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— Zahl der Zähne : Nlat und Ndors sind zwei voneinander unabhängig 
meßbare Größen. (Was nicht heißt, daß ihre Werte nicht korreliert sein 
können.) Weil jedoch die Zahl der Lateralzähne bei den drei Gruppen 
fast gleich ist (Tabelle 9), variiert Ntot zwangsläufig mit Ndors. Ntot 
ist also kein unabhängiges Merkmal. 


— Anordnung der Zähne : Zahlenmäßig sind Nprox und Ndist von 
Ntot (also von Nlat und Ndors) abhängig. Die Ratio Nprox/Ndist 
(P/D-Ratio) ist das aber nicht. Diese gibt Information über die 
Anordnung der Zähne. Weil Zahl und P/D-Ratio der Lateralzähne 
nur wenig variieren, ist aber die gesamt-P/D-Ratio von der P/D-Ratio 
der Dorsalzähne abhängig. 


104 


Tabelle 4 
Zahl der Dorsalzähne (Ndors) 
IO IE CS IS CIRE CSI 6 
bis bis bis bis bis bis bis bis bis 
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— Länge der Zähne : Die mittlere Länge aller Zähne wird stark durch 
die Ratio Lateralzähne/ Dorsalzähne mitbestimmt, weil letztere kleiner 
sind. Die mittlere Länge der ersten zehn Lateralzähne ist davon aber 
unabhängig. 


Praktischer Wert der Unterscheidungsmerkmale 


— Zahl der Zähne: Der Unterschied der Zahl der Lateralzähne ist 
für praktische Zwecke zu gering und würde große Stichproben erfor- 
dern. Theoretisch wäre also Ndors als Unterscheidungskriterium zu 
wählen. Aus praktischen Gründen ist aber Ntot zu bevorzugen, weil 


105 


Tabelle 5 
Zahl der Proximalzähne (Nprox) 


TROT A TS AT MOI DIDI 
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diese ohne jegliche Erfahrung im Unterscheiden von Lateral- und 
Dorsalzähnen leicht zu bestimmen ist. 


— Anordnung der Zähne : Die Anordnung der Zähne auf der Valve 
wird vor allem von der P/D-Ratio der Dorsalzähne bestimmt. Es ist 
jedoch weitaus einfacher, die Gesamtzahl der Proximalzähne zu 
bestimmen. Diese wird zwar mitbestimmt von 1. Zahl und P/D-Ratio 
der Lateralzähne, und 2. Zahl der Dorsalzähne, aber effektiv hat nur 
diese letzte Zahl Einfluß auf das Ergebnis, und zwar in dem Sinne, 
dal3 dadurch der Unterschied zwischen den drei Gruppen hervorgehoben 
wird. (Siehe Tabelle 9.) Der Vorteil ist, daß Nprox einfach und schnell 
zu bestimmen ist. 


106 


Tabelle 6 
Zahl der Distalzähne (Ndist) 
Oeil 314436 a4l 746 


Lok. | bis bis is bis bis bis bis 
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— Länge der Zähne : Das Bestimmen der mittleren Länge der Late- 
ralzähne ist arbeitsintensiv, liefert aber ein brauchbares Kriterium zur 
Unterscheidung von sudetica und momos. 


Signifikanz der Unterschiede 


Für jedes der quantitativ variablen Merkmale (Genitalmerkmale und 
alpha) und für die daraus berechnete P/D-Ratios wurden die Gesamt- 
frequenzkurven der sudetica-Gruppe, der momos-Gruppe und der 
melampus-Gruppe paarweise miteinander verglichen und es wurde die 
Wahrscheinlichkeit (p), daß es sich um zwei Stichproben der gleichen 
Population handle, mit dem t-Test von Student berechnet. Die p-Werte 


107 


Tabelle 7 


Mittlere Länge der ersten zehn Lateralzähne (L), in mikrometer 


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Lok.| bis bis bis 1 is bis bis bis bis bis 
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sind in Tabelle 9 eingetragen. Bei einem Signifikanzniveau von 5% 
(1,67% nach Durchführung der Bonferroni-Korrektur) erweist sich 
melampus in allen studierten Merkmalen signifikant von sudetica und 
momos verschieden. Momos unterscheidet sich in Zahl und Lange der 
Valvenzähne signifikant von sudetica, nicht jedoch in deren Anordnung 
(P/ D-Ratio). Die Anordnung der Hinterflügelflecke (alpha) ist bei allen 
drei Gruppen signifikant verschieden. 


Vergleich der melampus-Gruppe und der sudetica-Gruppe 


Es sind diese Gruppen, die Warren vor Augen hatte als er sudetica 
in den Artrang erhob. Sie zu unterscheiden, war eigentlich nie ein 


108 


Tabelle 8 
Winkel alpha, in Graden. Erklärung : siehe Abb. 4 
-10 -5 0 
bis bis bis 
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Problem. In allen untersuchten Merkmalen unterscheiden sie sich 
signifikant (siehe Tabelle 9). Jetzt, da von einer dritten Gruppe die 
Rede ist, sollen die Unterschiede aber klar definiert werden. 


Unterschiede 


— Zahl der Zähne : Ntot ist bei melampus um 78%, Ndors sogar um 
146% größer als bei sudetica. 


— Anordnung der Zähne: Die P/D-Ratio der Dorsalzähne ist bei 
melampus um 225% größer als bei sudetica, Nprox um 266%. Aus 
Tabelle 1 (unter V) geht hervor, daß dies nicht durch eine größere 
Länge des proximalen Valvenabschnitts verursacht wird. 


109 


Tabelle 9 


Links : Mittelwert (M) und Ratio proximal/distal (P/D) der gemessenen Valvenmerk- 
male und des Winkels alpha, pro Gruppe. SD = Standardgbweichung 
Rechts: Aus Student’s t-Test hervorgehende p-Werte bei paarweisen Vergleich der 
drei Gruppen 


Sud Mom Mel Sud/Mel Sud/Mom Mom/ Mel 
348 330 354 I P R 
34.9 48.7 <0.01% <001% <0.01% 
5.8 8.3 
0.29 VS <0.01 % 100 % <0.01 % 
0.08 0.14 
152 16.0 <0.01% <001% <0.01% 
189 23 
0.46 0.53 <0.01 % 100 % <0.01 % 
0.12 0.16 


10,7 524] <0.01% <001% <001% 
Del Tes 


0.52 <0.01 % 3.8 % <0.01 % 
0.21 


SD 

Nprox M 16.3 <0.01 % <0.01 % <0.01 % 
SD 4.3 

Ndist M 
SD 

L M 


<0.01% <001% <001% 


— Zahnlänge : Die mittlere Lange der Lateralzähne (L) ist bei sudetica 
um 47% größer als bei melampus. 


— Die Kurven des Winkels alpha iiberschneiden sich im Bereich von 
10-20°. Die Hälfte aller melampus-Werte liegt darüber, die Hälfte aller 
sudetica-Werte darunter. 


— Die Frequenz von O, variiert bei sudetica je nach Population von 
24%-84%. Bei melampus wurde dieses Merkmal kein einziges Mal 
festgestellt. (Siehe Tabelle 1). 


110 


Länge der Valvenzähne Zahl der Valvenzähne 


% % 


% = 


SS 


7-8 9-10 11-12 13-14 15-16 17-18 19-20 21-22 23-24 25-26 1-5 11-15 21-25 31-35 41-45 51-55 


Zahl der Proximalzähne 


1-2 5-6 9-10 13-14 17-18 21-22 25-26 29-30 6-10 16-20 26-30 36-40 46-50 


50 
Alpha 


40 
30 
% 
20 
10 


0 
-10-5 -5-0 0-5 5-10 10-15 15-20 20-25 25-30 30-35 56-60 


Abb. 5. Variationskurven der gemessenen Merkmale, pro Gruppe. 
LO = E. sudetica ; + = E. melampus ; O = E. momos. 


111 


— Auf der Hinterflügelunterseite bildet nur F, ein zuverlässiges Un- 
terscheidungsmerkmal. Bei sudetica ıst dieser Fleck in allen Popula- 
tionen vorhanden (19%-90%), bei melampus wurde er nur in drei 
Populationen einmal gefunden. | 


— Die Variationsbreite der meisten Valvenmerkmale ist bei melampus 
größer als bei sudetica. Letztere ist also, in dieser Hinsicht, die homo- 
genere Gruppe, was bemerkenswert ist, weil ihr Areal am stärksten 
disjunktiert ist. 


Trennbarkeit 


Zusammenfassend kann man sagen, daß die von Warren (1936 ; 1949 ; 
1961) beschriebenen Unterscheidungsmerkmale reell sind (allerdings 
nur ın bezug auf diese beiden Gruppen), aber nicht absolut : für jedes 
Merkmal gibt es Tiere, die nicht zu bestimmen sind, die sog. Partial- 
übergänge sensu Lorkovic (Lorkovic, 1938-42 ; 1953 ; Sijaric, 1980). 
Unter allen untersuchten sudetica fand ich jedoch nur 11 Totalüber- 
gänge, ın der melampus-Gruppe keinen einzigen. Das heißt, daß auf 
Grund der untersuchten Merkmale nicht nur jede Population, sondern 
auch 98.4% der Einzeltiere mit Sicherheit entweder als sudetica oder 
als melampus bezeichnet werden können. 


Die momos-Gruppe 


Zuerst fällt auf, daß die momos-Gruppe in ihren quantitativ variablen 
Merkmalen genauso einheitlich ist wie die beiden anderen Gruppen. 
Die Merkmalverteilungen stimmen aber weder mit denen von sudetica 
noch mit denen von melampus ganz überein. 


Unterschiede 


— Zahl der Zähne. Sowohl Nlat als Ndors sind intermediär zwischen 
melampus und sudetica. 


— Anordnung der Zähne : In der P/D-Ratio der Lateralzähne unter- 
scheiden sich momos und sudetica nicht; in der P/D-Ratio der 
Dorsalzähne unterscheiden sie sich nicht signifikant. Der Unterschied 
gegenüber melampus ıst dagegen groß. 


— Die mittlere Länge der Lateralzähne hält die Mitte zwischen der 
von sudetica und der von melampus. 


— Die Anordnung der Flecke auf den Hinterflügeln ist fast wie bei 
sudetica. 


— F,. Bei sieben der untersuchten Populationen (Nr. 20, 22-27) fehlt 
dieser Fleck fast immer. Bei vier Populationen dagegen zeigt eine 


112 


beträchtliche Zahl der Falter (17%-25%) dieses Merkmal. Diese Fre- 
quenzen sind immerhin niedrig im Vergleich zu denen der sudetica- 
Gruppe. 


— Ok. In den vier obenerwähnten Populationen (21, 28, 29, 30) weist 
auch ein hoher Prozentsatz (>35%) der Tiere die Ocelle in Zelle 6 
der Vorderflügeloberseite auf. In den übrigen Populationen ist die Fre- 
quenz weitaus geringer (15%), oder fehlt die Ocelle ganz (Tabelle 1). 


Zusammenfassend : in einigen Merkmalen (P/D-Ratio, alpha) ist 
momos nicht oder nur schwer von sudetica zu unterscheiden ; in den 
anderen Merkmalen ist sie intermediär zwischen melampus und sudetica. 
Dabei sind die untersuchten Populationen nach ihren Valvenmerkmalen 
ziemlich homogen. Die Ausprägung sudetica-ähnlicher Flügelmerkmale 
ist aber recht unterschiedlich und erlaubt eine weitere Zweiteilung der 
Gruppe. 


Trennbarkeit 


Wenn man Einzeltiere betrachtet, findet man in jeder Population 
Exemplare, die man zu melampus oder zu sudetica stellen kann, zudem 
aber, für jedes Merkmal, eine beträchtliche Zahl von Übergängen. Das 
erweckt den Anschein einer kontinuierlichen morphologischen Varia- 
tionsreihe, die von melampus über momos zu sudetica führt. Auf 
Populationsniveau jedoch zeigt sich eine geographisch bedingte, va- 
riationsstatistische Diskontinuität, die es bis jetzt ermöglichte, jede 
Population anhand einer kleinen Stichprobe eindeutig entweder als 
melampus oder als momos oder als sudetica zu bezeichnen. Somit 
gliedert sich der E. melampus-Komplex zwangsläufig in drei morpho- 
logisch-geographische Einheiten. 


Kontaktzonen und eventuelle Sympatrie 
melampus-momos 


Aus dem Grenzgebiet melampus | momos liegt eine Reihe sehr kleiner, 
aber brauchbarer Serien vor. Tiere aus Ischgl, Sankt Anton und Serfaus 
(Tirol) gehören eindeutig zu melampus ; Tiere aus dem Kaunertal 
dagegen gehören zu momos. Offenbar bildet hier das Inntal die Grenze. 
Zwischen Ortleralpen und Ötztaler Alpen (Stichproben von Solda und 
Schnalstal) bildet das Etschtal eine natürliche Barriere. Es gibt aber 
Hinweise dafür, daß sich wenigstens auf dem rechten Etschufer Misch- 
populationen befinden. Es liegen 7 Tiere aus dem Martelltal und 3 
aus Vermiglio vor (leg. W. Arnscheid). In beiden Stichproben finden 


1.13 


sich nebst Tieren, die eindeutig zu melampus zu stellen sind (das heißt 
also, daß zumindest ein Merkmal außerhalb der Streubreite sämtlicher 
momos-Populationen liegt), auch Tiere, die zweifellos zu momos 
gehören (also zumindest ein Merkmal außerhalb der Streubreite sämt- 
licher melampus). Obwohl es sich um sehr wenig Tiere handelt, ist 
klar, daß hier eine Vermischung von melampus und momos vorliegt. 
Eine Analyse größerer Serien aus diesem Gebiet wäre dringend erfor- 
derlich, um eine eventuelle Hybridisation feststellen zu können. 


sudetica-momos 


Aus den Ostalpen sind bisher keine sudetica-Populationen bekannt. 
E. momos und sudetica sind also, soweit wir wissen, völlig allopatrisch. 


sudetica-melampus 


E. sudetica und melampus treffen an zwei Stellen in den Alpen auf- 
einander : im Berner Oberland und in den französischen Alpen. Ein 
sympatrisches Vorkommen ist in der Literatur mehrmals erwähnt 
worden, und zwar bei Grindelwald (Bretherton, 1966; Forster & 
Wohlfahrt, 1976 ; Réal, 1988). Es fehlen aber Originalmeldungen oder 
Hinweise darauf. Wahrscheinlich gehen diese Angaben auf Warren 
zurück, der zunächst (1936) aus Grindelwald nachdrücklich E. melam- 
pus nennt und später (1949) sowohl melampus als sudetica aus den 
Berner Alpen erwähnt, was auch richtig ist. Nach Bretherton (1966) 
wird die Kohabitation von Sonderegger (1963) bestätigt, indem er sagt : 
“Unter meiner Grindelwaldner Ausbeute brachte ich auch zwei E. 
melampus ssp. tigranes mit nach Hause.” Die Tiere wurden aber in 
einem Biotop (trockenen Straßenrand, östlich des Dorfes) gefangen, 
wo E. sudetica nicht fliegt (Sonderegger, in litt.). Trotz intensiver Suche 
hat Sonderegger später an dieser Stelle nie mehr E. melampus beob- 
achtet ; auch von anderen sind weder Fundmeldungen noch Belegstücke 
bekannt. Er nimmt an, daß die Tiere beim Straßenbau verschleppt 
wurden. Auf jeden Fall ist melampus dort nicht bodenständig. Neulich 
wurde auch in der Schweizer Literatur der vermeintlichen Kohabitation 
von melampus und sudetica deutlich widersprochen : “E. sudetica 
kommt in der Schweiz nur in der Umgebung von Grindelwald vor 
und ist gegenwärtig weitgehend von E. melampus getrennt” (Lepidop- 
terologen-Arbeitsgruppe der Schweiz, 1987). 


Auch aus den französischen Alpen ist ein Fall von Sympatrie gemeldet 
worden : in einer Serie von 85 Stück E. sudetica, die vor dem letzten 
Weltkrieg im Massiv de Belledonne gesammelt wurde, fand Réal (1988) 
(nebst einigen zweifelhaften Weibchen) ein einziges melampus-Männ- 


114 


chen. Eine melampus-Population wurde aber nie gefunden. In den 
letzten Jahren konnte in diesem Gebiet nur sudetica nachgewiesen wer- 
den. Ein sympatrisches Vorkommen an dieser Stelle ist daher unbe- 
wiesen. Etwas östlicher fand Savourey (in litt.) eine Kontaktstelle von 
melampus und sudetica. Obwohl es im Gelände keine natürliche 
Barriere gibt, vermischen die beiden sich nicht (7). Es muß also fest- 
gestellt werden, daß bis jetzt keine einzige Stelle bekannt ist, an der 
melampus und sudetica unter Beibehaltung ihrer spezifischen Merkmale 
ständig zusammenfliegen. Es fehlt also bislang der Beweis für einen 
vollendeten Speziationsprozeß. 


Geographische Verbreitung und Variation 
Die melampus-Gruppe 


Das melampus-Areal umfaßt die Alpen westlich der Grenze von 
momos, ausgenommen die sudetica-Enklave bei Grindelwald und die 
französischen Westalpen (Abb. 6). Allgemein werden zwei ssp. aner- 
kannt : nominotypische melampus (locus typicus : Graubünden), und 
ssp. tigranes Fruhstorfer, 1910 (l.t.: Chamonix). Letztere ist größer 
und reicher gezeichnet. Inwiefern sie sich geographisch trennen lassen, 
ist aber ungenügend bekannt (). Auf jeden Fall gibt es intermediäre 
Populationen, die von Fruhstorfer (1910) sogar als ssp. augurinus 
beschrieben wurden. 


Die momos-Gruppe 


Diese Gruppe bewohnt die Ostalpen; fehlt offensichtlich in den 
Karawanken und den Julischen Alpen, in den Venezianischen Alpen 
und den Lessinischen Alpen, und im Dachsteingebiet. Innerhalb dieser 
Gruppe gibt es große Unterschiede im Habitus. Sehr auffallend ist der 
bereits erwähnte Unterschied zwischen den Populationen aus dem 
Stubaital, den Gurktaler Alpen und den Niederen Tauern auf der einen 
Seite, und den übrigen Populationen auf der anderen Seite. Obwohl 
genitaliter kein Zweifel daran besteht, daß alle der momos-Gruppe 
angehören, haben die Flügelmerkmale der ersten Gruppe eine so starke 
sudetica-Prägung, daß die Fruhstorfer’schen momos-Merkmale auf sie 
nicht mal zutreffen (“Klein, Längsbinde schmal und sehr dunkel, 


(2) Ein ähnliches Verhalten ist die Regel bei den alpinen Vertretern der E. tyndarus- 
Gruppe (DE Lesse, 1952 ; Lorkovic, 1957 ; Lorkovic & DE LESSE, 1954). 

(2) Sogar Warren hatte seine Mühe damit : die Populationen aus dem Berner Oberland 
(Kandersteg und Niesen) stellte er zuerst zu melampus (WARREN, 1936), später zu 
tigranes (WARREN, 1949). 


115 


\ 


_ 


Se E. sudetica Wf melampus-Gruppe W momos-Gruppe 


Y, N 
Ubergangszone Uz EN Grenze unsicher 


Abb. 6. Verbreitung der drei Formen innerhalb der Alpen. Teilweise nach Willien 
(1986), Gonseth (1987), Verity (1953). 
Gestrichelte Linie : Grenze des Alpenraumes. 


unterseits mit unbedeutenden gelblichen Makeln”) (siehe Abb. 7). Sie 
wird hier deshalb als neue Subspezies betrachtet. 


Erebia melampus momos Fruhstorfer, 1910 


Die ssp. momos ist verbreitet in den Dolomiten, den Karnischen Alpen 
und den Hohen Tauern, und nach Westen hin, allerdings weniger 
typisch, in den Otztaler Alpen. 

Erebia melampus semisudetica ssp.n. (Abb. 7) 

Hororypus : &, “Neustift-Tirol, Austria. Elferhiitte, 1800 m., 6.VIIL.91. 
F. Cupedo leg.”, in Coll. F. Cupedo. 


116 


PARATYPEN : 40 66 und 10 29, gleiche Funddaten, in Coll. F. Cupedo, 
und Coll. Institut für taxonomische Zoologie (Zoologisches Museum), 
Amsterdam. 


Die äußerst lebhaft gefärbte und reich gezeichnete Form der Stubaier 
Alpen. Genital von E. melampus momos nicht zu unterscheiden ; die 
Ocelle in Zelle 6 der Vorderflügeloberseite häufig vorhanden (>35% 
der Tiere) ; Fleck in Zelle Ic der Hinterflügelunterseite oft vorhanden 
(17-25% der Tiere). 


VERBREITUNG : Stubaier Alpen, Osterreich. Zu diesem Taxon sind nach 
derzeitiger Kenntnis auch die Populationen der Gurktaler Alpen und 
der Niederen Tauern zu rechnen. Die Frage, ob wirklich eine Areal- 
disjunktion vorliegt, muß dahingestellt bleiben, weil im Moment kein 


if Abb. 7. Erebia melampus semisudetica ssp.n. 
Neustift (Österreich, Tirol), 1800 m, 6.VIII.91, 3 46 (links) und 3 99 (rechts). Oben 
links : Holotypus. 


117 


Material aus den verbindenden Gebirgszügen (Zillertaler, Kitzbühler 
und Salzburger Alpen) vorliegt. 


Die sudetica-Gruppe 


Das Areal von sudetica ist größtenteils extra-alpin und stark disjunktiert. 
Unser Wissen beziiglich ihrer Verbreitung erweitert sich standig. Fol- 
gende Ubersicht ist aktuell. Nur von einem Fundort (auf der polnischen 
Seite der Sudeten) besitze ich keine Belegstiicke. 


E. sudetica sudetica Staudinger, 1861 


Nur in den Sudeten (Gesenke oder Jesenik). In Tschechien heute nur 
noch an fünf Stellen im Altvatergebirge (Hruby Jesenik) : Aitvater 
(Praded), GroBe Kessel (Velky Kotel), Karelsbrun (Karlova Studanka), 
Hohe Haide (Rejviz) und Schweizerei (Skritek) (Stiova, 1988). Fliegt 
auch auf der polnischen Seite der Sudeten : Ladek Zdroj (Krzywicki, 
1982 ; Kudrna, 1986). Alte Angaben aus der Tatra konnten nicht 
bestätigt werden : das vorhandene Material erwies sich als E. epiphron 
(Moucha, 1959). 


E. sudetica radnaensis Rebel, 1915 


In den Nordkarpaten nur auf der Südseite des Rodnaer Schneegebirges ; 
in den Südkarpaten bekannt aus dem Retezatgebirge und aus dem 
Ciucasgebirge (König, 1975). Von Hormuzaki (1901) zitiert alte Funde 
aus dem Bucegigebirge. Neuere Nachweise fehlen. 


E. sudetica liorana de Lesse, 1947 (*) 


Bewohnt ein engbegrenztes Gebiet im französischen Zentralmassiv 
(Plomb du Cantal, Puy Mary) (Bretherton, 1966a ; Burton, 1977; 
Perceval, 1976 ; Willien, 1990). 


E. sudetica inalpina Warren, 1949 


Fliegt an den Südhängen des Faulhornmassivs (Grindelwald, Berner 
Alpen, CH) ; wurde neulich auch im Lauterbrunnental, bei Stechelberg 
entdeckt (M. Delnoy leg.) 


(*) Von DE Lesse (1947) als ein Quadrinomen beschrieben : Erebia melampus ssp. 
tigranes race lioranus. Weil aber de Lesse nachdrücklich betont, mit “race” sei das 
Aquivalent des englischen “form” gemeint, ist liorana, gemäß Artikel 45g-ü-l der Code 
(ICZN, 1985), als von subspezifischem Rang zu werten, da der Name vor 1985 als 
verfügbar behandelt und als Name einer Subspezies anerkannt worden ist (WARREN, 
1949 ; 1961). 


118 


E. sudetica belledonnae ssp.n. (Abb. 8) 


Hororypus : 6, “Le Rivier d’Allemont (F-38), Les Chaumes, 1650 m, 
13.V111.87, F. Cupedo leg.” Zweites Etikett : “Gen. prep. nr. 769, F. 
Cupedo”. In Coll. F. Cupedo. 

PARATYPEN : 35 G& und 4 Q9, gleiches Etikett, in Coll. F. Cupedo 
und in Coll. Institut für taxonomische Zoologie (Zoologisches Museum), 
Amsterdam. 


Abb. 8. Erebia sudetica belledonnae ssp.n. 
Le Rivier d’Allemont (Frankreich, Isère), 1650 m, 13.V111.87, 3 && (links) und 3 29 
(rechts). Oben links : Holotypus. 


Die Falter sehen denen der ssp. inalpina ähnlich, unterscheiden sich 
von diesen durch die schmaleren Postdiskalbinden mit weniger und 
beträchtlich kleineren Punkten (siehe Tabelle 10). Das verleiht ihnen 
auf den ersten Blick ein düsteres Aussehen. Aus diesem Grund und 


119 


Tabelle 10 
Die Zahl der schwarzen Punkte auf der Vfl.Os. 


N 0 


wegen ihres völlig isolierten Vorkommens betrachte ich sie als eine 
distinkte Subspezies. 


VERBREITUNG : Diese Subspezies bewohnt drei von tiefen Tälern ge- 
trennte Gebirgsstöcke in Isère und Savoie (französischen Alpen): 
1. Taillefer, 2. Belledonne - Arvan - Villards, 3. Le Grand Arc - La Lau- 
ziere - le Cheval Noir (Cupedo, 1987 ; Savourey, 1989). Ich nenne sie 
belledonnae ssp.n. nach dem Belledonnemassiv, wo ich 1981 die ersten 
Exemplare fing. 


VARIABILITAT : Die Tiere von La Toussuire sind etwas kleiner und 
schwächer gezeichnet als die der anderen Fundorte. Da aber Genau- 
stausch mit der Belledonne-Population möglich ist, sind die Unterschiede 
offensichtlich ökologisch bedingt. Bei la Toussuire fliegt die Art in 
offenen, windexponierten Biotopen bis über 2200 m, in La Morte und 
le Rivier d’Allemont dagegen in windgeschützten Lichtungen unterhalb 
der Baumgrenze (1600 m) (°). 


Es ist durchaus möglich, daß es in den französischen Alpen weitere, 
isolierte Vorkommen von E. sudetica gibt. Ich sah ein Exemplar aus 
Pralognan (M. Waber leg.), und zwei von Nevache (F. Borde leg.), 
konnte aber an beiden Stellen bisher keine Population auffinden. 


Diskussion 


Nach diesen Untersuchungen bildet der Erebia melampus-Komplex 
einen Formenkreis aus drei weitgehend allopatrische Gruppen. Nur 
melampus und momos treffen über die ganze Breite der Alpen auf- 
einander, wobei sich eine relativ schmale Übergangszone gebildet hat. 
Dies ist die übliche Situation, wenn zwei nah verwandte Taxa nach 
einer (glazialen) Isolation in sekundären Kontakt geraten (Bourgogne, 
1952 ; Ford, 1962 ; Huxley, 1942 ; de Jong, 1972). 


(5) Die Population von la Toussuire ist das morphologische Äquivalent der ssp. 
melampus innerhalb der melampus-Gruppe. Es ist also denkbar, dass dem Unterschied 
zwischen ssp. melampus und ssp. tigranes in ähnlicher Weise ökologische Bedingungen 
zugrunde liegen. 


120 


Auffallend ist der weitgehend intermediäre Charakter der momos- 
Gruppe. Ein naheliegender Gedanke ist, daß es sich um eine Misch- 
population zwischen melampus und sudetica handelt. Diesen zu über- 
prüfen, ist schwer möglich ; seine Wahrscheinlichkeit läßt sich aber 
einigermaßen einschätzen : 


Im Prinzip könnte eine solche Vermischung glazialen oder postglazialen 
Ursprungs sein. Letztere Möglichkeit ist aber aus verschiedenen Grün- 
den sehr unwahrscheinlich : 1. Wenn nämlich melampus vom Westen 
her, und sudetica vom Osten her die Ostalpen besiedelt und sich 
gemischt hätten, müßten die Frequenzkurven von momos (Abb. 5) den 
Charakter von Additionskurven haben, also eine größere Streubreite 
aufweisen und dadurch flacher sein. Beides ıst nicht der Fall. Gerade 
die häufigsten Varianten von melampus und von sudetica sind bei 
momos stark untervertreten ; die Extremvarianten (vor allem von 
melampus) fehlen sogar ganz, auch in den westlichsten momos-Popu- 
lationen. 2. Auch müßte man in dem Falle statt einer so charakte- 
ristischen Kontaktzone Merkmalgradienten finden. 3. Schließlich wäre 
es nur schwer verständlich, wie in den Ostalpen eine so innige Ver- 
mischung stattfinden konnte, während im Berner Oberland und in den 
französischen Alpen eine so strenge Vikarianz aufrechterhalten blieb. 


Eine Vermischung während des jüngsten Glazials ist dagegen nicht ganz 
auszuschließen. Wenn, wie heute, die präglaziale melampus in den 
Alpen beheimatet war und sudetica ın den Karpaten, könnte ın einem 
gemeinsamen Teil ihrer Refugien (im Periglazialraum zwischen Alpen 
und Karpaten) eine innige, langfristige Vermischung ihrer Genpools 
zustandegekommen sein. Für das Fehlen der Extremvarianten können 
wegen des langen Zeitraums sowohl Selektionsprozesse als genetische 
Drift verantwortlich sein. Eine Voraussetzung wäre allerdings, daß 
melampus und sudetica damals schon weitgehend morphologisch diffe- 
renziert, dagegen nicht oder kaum reproduktiv isoliert waren. 


Auf jeden Fall weist alles darauf hin, daß momos entweder als eine 
Mischform oder als eine selbständige phylogenetische Einheit spätestens 
während des Würmglazials entstanden ist. 


Bei der sudetica-Gruppe geht die Zersplitterung des Areals und die 
daraus erfolgte morphologische Differenzierung zum Teil schon auf eine 
würmglaziale Disjunktion zurück (Cupedo, unveröffentlicht). Die Ab- 
trennung der sudetica-Gruppe im ganzen muß also erheblich früher 
stattgefunden haben. 


Um die drei Gruppen taxonomisch einstufen zu können, müßte ihr 
reproduktiver Isolationsgrad bekannt sein. Durch die fast vollständige 


121 


Allopatrie ist das unmöglich. Nur Hybridisationsversuche können dar- 
über Aufschluß geben. Beim Genus Erebia gibt es davon schon ein- 
drucksvolle Beispiele (Lorkovic, 1961). 


Zur Feststellung des genetischen Differenzierungsgrades haben sich 
enzymelektrophoretische Methoden als sehr geeignet erwiesen (Geiger, 
1988 ; Geiger, 1990 ; Lattes et al., 1994). Beim E. melampus-Komplex 
jedoch geben sie bislang geringen Halt. Für die melampus-Gruppe und 
die sudetica-Gruppe fand Loertscher (1988) eine genetische Distanz (D, 
Nei, 1972) von 0.48, die momos-Gruppe blieb jedoch auBer Betracht. 


Der Speziationsgrad, der genetische Differenzierungsgrad, und sicherlich 
die phylogenetischen Beziehungen der Glieder des E. melampus- 
Komplexes bleiben somit weitgehend unklar. Deshalb sollte man aus 
rein praktischen Griinden nomenklatorisch jetzt alles beim alten 
belassen, das heißt : melampus und sudetica als zwei Arten und die 
ssp. der momos-Gruppe als ssp. von melampus behandeln. Nicht weil 
das richtig wäre, sondern weil jede Alternative im Moment genau so 
spekulativ wäre und unnôtigerweise nomenklatorische Verwirrung 
verursachen würde. 


Dankwort 


Einen Teil des Materials, das dieser Arbeit zugrunde liegt, verdanke ich der 
Sammeltätigkeit meiner Kollegen. Von einigen Fundorten hätte ich kein oder 
weitaus weniger Material gehabt ohne die Hilfe von : Marcel Prick und Wim 
Hoen (Niederlande), Wilfried Arnscheid und Dr. Günther Achenbach (Deutsch- 
land), Prof. Dr. Vladimir Sterba und Dr. Lubomir Stiova (Tschechien) und 
Prof. Andreas Szabö (Rumänien). Dank der Bewilligung von Ben Brugge 
und Wim Hogenes konnte ich das umfangreiche Material der Sammlung des 
Zoologischen Museums Amsterdam studieren. Dr. Mathias Loertscher (Bern, 
Schweiz) gab mir wichtige Hinweise für die statistische Bewertung der Daten. 
Besonders erkenntlich bin ich Dr. Peter Roos (Sprockhövel, Deutschland) 
und Prof. Dr. Zdravko Lorkovic (Zagreb, Kroatien), deren sachverständliche 
Kritik die endgültige Fassung des Manuskripts mitbestimmt hat, und Marcel 
Prick, ohne dessen Bemühungen der deutsche Text voller Fehler geblieben 
wäre. 


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125 


Nota lepid. 18 (2) : 126 ; 29.11.1996 ISSN 0342-7536 


Dirk HAMBORG à (1957-1995) 


„Die Metamorphose ist ein grundlegendes Phänomen bei der Betrachtung von Faltern. 
Zugleich symbolisieren sie Zartheit und Vergänglichkeit...“ 
Dirk Hamborg 


Die Nachricht vom Tod unseres Kollegen und Freund Dirk Hamborg hat uns 
wie ein Schlag getroffen. In einer depressiven Kurzschlußhandlung hat er sich 
am 21. November 1995 in Bleckede bei Lüneburg das Leben genommen. 


Dirk ist am 7. Juli 1957 in Hamburg geboren. Er ist in Wolfsburg aufgewach- 
sen, wo er sehr früh sein großes Interesse der Nachtfalter verfolgte. 1978 publi- 
zierte er ein faunistisches Werk über die Nachtfalter Wolfsburgs. Dirk war auch 
Künstler und fing im Oktober 1981 ein Studium an der Gesamtschule Kassel 
im Fachbereich Visuelle Kommunikation, mit Schwerpunkt Freie Graphik, an. 
Dieses Studium unterbrach er von Juni bis Oktober 1985 um sich der freien 
Landschaftsmalerei in der Steiermark zu widmen. Zu dieser Zeit hat er sich auch 
stärker mit dem Thema „Nachtschmetterlinge und Kunst“ auseinandergesetzt. 
Vor allem die Noctuidae erschienen oft in seinen Bildern. Im Februar 1987 hat 
er seine Abschluß Prüfung zum Thema ’Nachtfalter, scheinbar und unscheinbar’ 
gemacht. 


Er hat seine Werke auch im Oktober 1994 anläßlich der Eröffnung der neuen 
Arbeitsund Sammlungsräume des Tiroler Landesmuseum Ferdinandeum in 
Innsbruck ausgestellt, wo ich ihm und seine Kunst erstmals kennenlernte. Im 
Winter war er Künstler, im Sommer ein hervorragender Feldbiologe. Er hat 
nicht nur die Nachtfalter, besonders die Noctuidae, sondern auch die Sesiidae 
studiert und fotografiert. Im Auftrag des Bundeslandes Steiermark hat er die 
Sesien Steiermarks erforscht und viel Neues entdeckt. 1996 hatte er vor, ver- 
mehrt die Schweizer Sesienfauna zu forschen. Leider wird es nicht mehr dazu 
kommen. 


Steven WHITEBREAD 


126 


Nota lepid. 18 (2) : 127-137 ; 29.11.1996 ISSN 0342-7536 


Endothermic heat production in three species 
of Nymphalidae (Lepidoptera) 


C. MAIER (1) & T. G. SHREEVE 


School of Biological and Molecular Sciences, Oxford Brookes University, Gipsy Lane Campus, 
Headington, Oxford OX3 OBP, UK 


Summary 


Endothermic heat production is often described as an uncommon mechanism 
in butterflies, although shivering has been reported in a number of species. 
Here, we detail initial observations of endothermic heat production by three 
Nymphalidae species (subfamily Nymphalinae) ; the Painted Lady, Cynthia 
cardui (L.) ; Peacock, Inachis io (L.) and Red Admiral, Vanessa atalanta (L.). 
Closed wing shivering occurred over the whole experimental ambient tem- 
perature range of 14.5-23°C. All species and individuals tested showed in- 
reased thoracic temperatures while shivering. The maximum excess tempe- 
ratures and the rates of warming recorded for the three species were T., = 
16.5°C at 3°C min! for C. cardui (T, = 19°C); T., = 18°C at 4°C min! 
for 7. io (T, = 16°C) and T,, = 13°C at 4.3°C min for V. atalanta (T, 
= 17°C). We suggest that endothermic heat production may contribute to 
the widespread distribution and generalist habitat associations of these long 
lived butterflies. We also suggest that the role of endothermic heat production 
in butterflies needs further examination. 


Résumé 


La production de chaleur endothermique est souvent considérée comme un 
mécanisme peu commun chez les papillons, bien que le «frissonnement» ait 
été signalé chez plusieurs espèces. Ici, nous détaillons des observations préli- 
minaires sur la production de chaleur endothermique chez trois espèces de 
Nymphalidae (sous-famille Nymphalinae). Le «frissonnement» ailes fermées 
s’est produit à toutes les températures expérimentales de 14,5 à 23°C. Toutes 
les espèces et tous les individus testés ont présenté une température thoracique 
élevée pendant le «frissonnement». Les températures maximales en excès et 
les taux de chauffage observés pour les trois espèces ont été T., = 16,5°C 
a 3°C min! pour C. cardui (T, = 19°C); T., = 18°C a 4°C min! pour 
1. io (1, = 16°C) et T., = 13°C à 4,3°C min! pour V. atalanta (T, = 17°C). 


(1) Corresponding author. 


127 


Nous suggérons que la production de chaleur endothermique pourrait contri- 
buer à la grande extension de la répartition géographique et aux associa- 
tions à habitat généraliste de ces papillons qui ont une grande longévité. De 
surcroît, nous estimons que des études supplémentaires sur le rôle de la pro- 
duction de la chaleur endothermique chez les papillons mériteraient d’être 
entreprises. 


Introduction 


Flight is essential for most Lepidoptera. In the majority of temperate 
species of butterfly the required thoracic temperature (T,,) for flight 
is above that of ambient air temperature (T,). A number of studies 
of the temperatures required for sustained voluntary flight indicate that 
species with moderate to fast wing beat frequencies require T;} of 28- 
40°C, with vigorous flight restricted to the range of 33-38° C (Kingsolver, 
1985). For species with slow, gliding flight, which do not require the 
same power and frequency of muscle contraction, flight may be initiated 
at lower thoracic temperatures ; for example 10°C for overwintering 
monarch butterflies, Danaus plexippus L. (Nymphalidae) (Chaplin & 
Wells, 1982) and 17-18°C for Parnassius phoebus F. (Papilionidae) 
(Guppy, 1986). 


There are two possible options for heat gain during pre-flight warm- 
up ; physiological (generating internal metabolic heat) or behavioural 
(choice of microhabitats, orientation and posture relative to the sun) 
(Heinrich, 1981). For butterflies, the most common method of raising 
body temperature is to use some form of basking. Endothermic heat 
production is often described as an uncommon mechanism in butterflies 
(Kingsolver 1985 ; Shreeve 1992 ; Heinrich, 1993), although shivering 
has been reported in a number of species, including Papilionidae, 
Hesperiidae and Nymphalidae (Findlay, Young & Findlay, 1983 ; 
Dennis, 1993). 


Endothermic heat production by butterflies is described as energetically 
expensive and inefficient because the generally narrow body and poor 
insulation leads to rapid heat loss (Kingsolver, 1985). Where it has 
been recorded in the butterflies Danaus plexippus (Kammer, 1970) and 
Papilio polyxenes (Rawlins, 1980) this mechanism of shivering only 
raises thoracic temperatures 4°C above ambient at the most , with 
a maximal recorded heating rate of 1.3°C min-!. In two species of 
Nymphalidae greater excess temperatures have been found ; 18°C in 
Vanessa atalanta (Krogh & Zeuthen, 1941) and 8-11°C in Nymphalis 
antiopa (Douglas, 1986). In contrast, endothermic heat production is 
well documented in many species of moth, chiefly Sphingidae and 


128 


Saturniidae, which may elevate body temperature to at least 20°C 
above ambient by prolonged shivering (Heinrich & Bartholomew, 
1971 ; Bartholomew & Casey, 1973). 


Here, we detail initial observations of endothermic heat production 
by three Nymphalidae species (subfamily Nymphalinae) ; the Painted 
Lady, Cynthia cardui (L.) ; Peacock, Inachis io (L.) and Red Admiral, 
Vanessa atalanta (L.). The data presented here, taken with that of 
previously published work on Nymphalinae species (V. atalanta, Krogh 
& Zeuthen, 1941 and N. antiopa, Douglas, 1986) indicate that in some 
butterfly species pre-flight warm-up using endothermic heat production 
may be both rapid and produce a large excess temperature over am- 
bient. 


Materials and methods 


Specimens of C. cardui (N = 8), I. io (N = 6) and V. atalanta (N 
= 2) were caught in an Oxfordshire garden between 10 August and 
15 September 1994, or were from captive stock reared from eggs of 
wild females caught during the same period. Prior to measurements 
of endothermic heat production, individuals were kept for a maximum 
of 21 days, in a 50 cm X 50 cm X 60 cm flight cage, with free access 
to fresh cut flowers and honey-water. Trials took place in an unheated 
room over the temperature range 14.5-23°C, under conditions of low 
illumination («<5 watts m). Butterflies were immobilised by placing 
them in a jar with ethyl acetate. As soon as they stopped moving 
vigorously (approximately 10-15 seconds) they were removed from the 
jar and restrained on a setting board, with the wings held open by 
paper strips. Immobilising specimens by placing them in a killing jar 
for a few seconds is a method recommended by Friedrich (1986) for 
hand pairing nymphalid butterflies. This technique was found to be 
preferable to cooling or to no immobilisation as it facilitated thermistor 
implantation with minimal damage. A 0.4 mm diameter micro ther- 
mistor probe (Skye Instruments : SKTS 200U/M, time constant <1.0 
sec) was shallowly inserted under the dorsal thoracic cuticle and held 
in position by water soluble glue (Gloy!M). The butterfly was released 
from the setting board when the glue had dried (approximately ten 
minutes). The leads from the thermistor probe consisted of a 1 m long, 
0.13 mm diameter wire, leading to a lightweight screened cable con- 
nected to a data logger (Skye Instruments : SDL 2960). This arran- 
gement allowed the butterfly free wing movement when settled, together 
with limited flight at the end of the tether. Butterflies treated in this 
way lived for up to ten days after the implant was removed, behaving 


129 


in an apparently normal fashion (e.g. flying and feeding), suggesting 
that minimal internal damage was caused by the experimental procedure. 


Temperatures were recorded at 30 second intervals. This time period 
was used in preference to shorter periods for ease of comparison with 
other studies presenting data on this time scale (e.g. Kammer 1970). 
In addition to butterfly thoracic temperature, simultaneous data were 
recorded from a bare thermistor probe (control probe) placed adjacent 
to the butterfly, and an integral temperature sensor with radiation shield 
to record ambient air temperature. For the first three trials two further 
controls were used ; freshly dead butterflies of the same species as the 
test specimen, one with wings set closed, the other with wings open 
(basking posture), with thermistors implanted in the same way as for 
the test specimen. As thoracic temperatures of these butterflies showed 
no difference from that of the control probe, they were not included 
in subsequent trials. Ambient and control probe temperatures never 
differed by more than 1°C. 


Results 


When butterflies were released from the setting board, the typical 
pattern of behaviour was to settle with wings closed. Shivering occurred 
either spontaneously or in response to mild disturbance (e.g. gently 
flicking the wings). Butterflies shivered while standing still with wings 
closed, shivering being manifested by high frequency, low amplitude 
vibrations of the wings. Shivering occurred over the whole experimental 
ambient temperature range of 14.5-23°C. All species and individuals 
tested showed increased thoracic temperatures while shivering (Tab. 1). 
After a few minutes (mean = 4.8 mins, S.E. = 0.28, N = 28 shivering 
bouts) they usually attempted to fly (flying vigorously for a few seconds 
at the end of the tether), after which they would settle again with their 
wings closed. 


At the lower ambient temperature range (14.5 + 6°C) the mean thoracic 
temperature when flight was attempted by C. cardui was 24.4 + 1.1°C 
(N = 6). For I. io, at T, of 15-16°C, the mean thoracic temperature 
for attempted flight was 30.7 + 2.3°C (N = 3). At the upper ambient 
temperature range, mean thoracic temperatures at flight initiation were 
higher for C. cardui (mean = 31.1°C, at T, of 19-20°C, N = 2) but 
not for /. io (mean = 30.0°C, at T, of 20-23°C, N = 3). Mean thoracic 
temperature at flight initiation for V. atalanta was 29°C at T, of 17- 
17.5°C (N = 2). After attempted flight, individuals usually returned 
to a posture of standing with closed wings as they cooled. This pattern 


130 


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was often repeated with subsequent bouts of shivering followed by 
an attempt at flight. Fig. 1 illustrates increased thoracic temperatures 
achieved by shivering, for a peacock and painted lady butterfly tested 
simultaneously. Fig. 2 shows a similar pattern for a red admiral. 


The maximum excess temperatures and the rates of warming recorded 
for the three species were T., = 16.5°C at 3°C min for C. cardui 
(T, = 19°C); T., = 18°C at 4°C min! for 7. io (T, = 16°C) and 
T., = 13°C at 4.3°C min for V. atalanta (T, = 17°C). When butterflies 
were quiescent (before shivering and after cooling down from a 
shivering bout), thoracic temperatures were not significantly different 
from that of the control probe (t = 0.7, P= 0.49, N = 24). 


Discussion 


The maximum excess temperatures and the rates of warming recorded 
were higher than those found for P polyxenes (Rawlins, 1980) and 
D. plexippus (Kammer, 1970) which belong to a different family and 
subfamily (Papilionidae and Danainae respectively) from that of the 
species tested here. Our results are directly comparable with two earlier 
studies of members of the Nymphalidae, subfamily Nymphalinae ; 
Vanessa atalanta (Krogh and Zeuthen, 1941) and Nymphalis antiopa 
(Douglas, 1986) which may achieve a T,, of 8-18°C by shivering. 
Thoracic temperatures recorded at flight initiation for the three 
Nymphalinae species we tested are similar to those found for voluntary 
flight initiation of other species of butterfly which do not engage in 
gliding flight (Kingsolver, 1985 ; Shreeve, 1992). 


Our data, taken with that of Krogh & Zeuthen (1941) and Douglas 
(1986), representing four species of Nymphalidae, subfamily Nympha- 
linae, may indicate that there is a phylogenetic component to the 
shivering posture adopted and temperature excess achieved by these 
butterflies. Further investigation of the occurrence of endothermy in 
different butterfly taxa would help to clarify this. The ability to use 
endothermic means of heat production may be particularly advanta- 
geous to these generally long lived butterflies and contribute to their 
overall widespread distributions and generalist habitat associations. 


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diapause as adults. The ability to generate heat by shivering would 
enable overwintering adults to warm-up and move from (or within) 
cool, dark hibernation sites, when behavioural means of warming up 
are not an option. Douglas (1986) reports that N. antiopa, which also 
overwinters as an adult, raises its thoracic temperature by shivering 


182 


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to enable it to move from tight, dark hibernation roosts to open sunlit 
areas where it can bask. A similar function for shivering has been 
suggested for the black swallowtail, P polyxenes, which uses endo- 
thermic heat production as a subsidiary warm-up mechanism under 
conditions of low solar radiation and ambient temperature. Under these 
circumstances shivering probably generates enough heat to allow 
minimal activity for activities such as changing habitat or regaining 
a roosting site if dislodged (Rawlins & Lederhouse, 1978). Shivering, 
followed by flight, has been observed in an overwintering /. io when 
dislodged from its roost (C. Maier pers. obs.). 


Prior to entering diapause and immediately afterwards overwintering 
adult butterflies spend considerable time feeding to accumulate and 
replace stored energy reserves respectively (Pullin, 1986). In autumn 
and early spring ambient temperatures may be low and solar radiation 
of low intensity. At these times of the year these energy sources may 
also be extremely variable, both within and between days. In such 
marginal circumstances the ability to employ endothermy may allow 
individuals to maximise the time they can spend feeding. 


Cynthia cardui does not enter winter diapause and is probably not 
a permanent resident within Europe, its main permanent centres being 
in North Africa, Arabia and Asia (Emmet & Heath, 1989). Vanessa 
atalanta is also a temporary resident in northern latitudes with 
permanent centres within southern Europe. Both are strongly oppor- 
tunistic and engage in extensive migration. Endothermic heat production 
may facilitate this strategy, allowing summer survival in extreme 
northern areas including locations within the Arctic Circle. It may also 
account for observations (Emmet & Heath, 1989) of active adults of 
both species during warm winter days in December and January within 
the British Isles. 


All three species are fairly brightly coloured on their dorsal wing surface 
but the underside is cryptic against usual resting backgrounds. All can 
adopt an open winged (dorsal) basking posture, but basking in 
conditions of low solar radiation, when body temperature is too low 
for efficient flight, may make individuals vulnerable to potential preda- 
tor attack. Several studies suggest that butterflies are most at risk from 
predation in marginal temperature and weather conditions when 
effective thermoregulation and active flight are difficult (Rawlins & 
Lederhouse, 1978 ; Bowers, Brown & Wheye, 1985 ; Kingsolver, 1987). 
The ability to raise thoracic temperature endothermically, with closed 
wings, may in certain circumstances enable flight temperature to be 


135 


achieved without compromising the protection afforded by the cryptic 
pattern of the hind wings. 


We suggest that the role of endothermic heat production in butterflies 
needs further examination, in respect of actual temperatures achieved 
and in relation to the energetic cost of this warming method. We further 
suggest that for nymphalid species there is a case for further research 
into how periodic endothermy may contribute to adult overwintering 
success and what role it might play in migratory strategies. 


Acknowledgements 


Andrea Turner kindly translated the abstract into French. 


References 


BARTHOLOMEW, G. A. & Casey, T. M.,1973. Effects of ambient temperature 
on warm-up in the moth Ayalophora cecropia. J. exp. Biol. 58 : 503- 
SOM: 

Bowers, M. D., Brown, I. L. & WHEYE, D., 1985. Bird predation as a 
selective agent in a butterfly population. Evolution 39 : 93-103. 

CHAPLIN, S. B. & Weııs, P. H., 1982. Energy reserves and metabolic 
expenditures of Monarch butterflies over wintering in southern California. 
Ecol. Ent. 7 : 249-256. 

Dennis, R. L. H., 1993. Butterflies and Climate Change. Manchester 
University Press, Manchester. 

Douac as, M. M. 1986., The Lives of Butterflies. University of Michigan Press, 
Michigan. 

Emmet, A. M. & HEATH, J. (Eds.), 1989. The Moths and Butterflies of Great 
Britain and Ireland. Vol. 7, Part 1. Hesperiidae-Nymphalidae. Harley 
Books, Colchester. 

FINDLAY, R., Young, M. R. & FINDLAY, J. A., 1983. Orientation behaviour 
in the Grayling butterfly : thermoregulation or crypsis * Ecol. Ent. 8: 
145-153. 

FRIEDRICH, E., 1986. Breeding Butterflies and Moths. (English edition). Harley 
Books, Colchester. 

Guppy, C. S., 1986. The adaptive significance of alpine melanism in the 
butterfly Parnassius phoebus F. (Lepidoptera : Papilionidae). Oecologia 
70 : 205-213. 

HEINRICH, B., 1981. Insect Thermoregulation. John Wiley & Sons, New York. 

Hernricu, B., 1993. The Hot Blooded Insects : Strategies and mechanisms 
of thermoregulation. Springer-Verlag, Berlin. 

HEINRICH, B. & BARTHOLOMEW, G.A., 1971. An analysis of the pre-flight 
warm up in the sphinx moth, Manduca sexta. J. exp. Biol. 55 : 223- 
239) 


136 


KAMMER, A. E., 1970. Thoracic temperatures, shivering and flight in the 
Monarch butterfly, Danaus plexippus (L.). Z. vergl. Physiol. 68 : 334- 
344. 

KINGSOLVER, J. G., 1985. Butterfly thermoregulation : organismic mechanisms 
and population consequences. J. Res. Lep. 24 : 1-20. 

KINGSOLVER, J. G., 1987. Predation, thermoregulation, and wing color in 
Pierid butterflies. Oecologia 73 : 301-306. 

KROGH, A. & ZEUTHEN, E., 1941. The mechanism of flight preparation in 
some insects. J. exp. Biol. 18 : 1-10. 

Purım, A.S., 1986. Life History Strategies of the Butterflies, /nachis io and 
Aglais urticae, Feeding on Nettle, Urtica dioica. Ph.D. Thesis (CNAA), 
Oxford Polytechnic. 

RAwLINS, J. E., 1980. Thermoregulation by the Black Swallowtail butterfly, 
Papilio polyxenes (Lepidoptera: Papilionidae). Ecology 61: 345-357. 

RAwLINS, J. E. & LEDERHOUSE, R. C., 1978. The influence of environmental 
factors on roosting in the black swallowtail, Papilio polyxenes asterius 
Stoll (Papilionidae). J. Lep. Soc. 32 : 145-159. 

SHREEVE, I. G., 1992. Adult Behaviour. Jn Dennis, KR. L. H. (Ed.): The 
Ecology of Butterflies in Britain. pp. 22-45. Oxford University Press, 
Oxford. 


137 


Nota lepid. 18 (2) : 138 ; 29.11.1996 ISSN 342-7536 


Book reviews - Buchbesprechungen - Analyses 


The Nepticulidae of eastern Europe and Asia. Rimantas PUPLESIS. 
291 pp. + 840 Figs, 16.8 x 24.7 cm, hardback. Backhuys Publishers, 
1994. ISBN 90-73348-29-3. Obtainable from : Backhuys Universal Book 
Services, P.O. Box 321, NL-2300 AH Leiden, The Netherlands. Price : 
fl 210, excl. postage. 


The family Nepticulidae comprise the smallest moths in the world, the majority 
ranging from about 2.5 to 6.5 mm in wingspan. Because of their small size, 
they have not been very well studied in the past. However, over the last 20 
years there has been a veritable boom in the study of the world’s Nepticulidae 
fauna. Major treatments of the Nepticulidae of South Africa, North America, 
Japan and Europe have appeared. The present book is the first taxonomic 
revision of the family from the rest of the Palaearctic Region, that vast expanse 
between Japan and Europe. The author is to be congratulated on this tremen- 
dous work, especially considering that much of the material was collected 
personally by him in the field, often under difficult conditions. 


In all, 221 species are treated, 114 of which have been described by the author. 
For each species, adults, and male and female genitalia are described, where 
known, and figured in very good line drawings. Full details of the biology 
are also given, including a description of the mine and the species’ known 
distribution is summarised. Unfortunately, keys to species are not given. Of 
general interest are the additional chapters on the methods used, adult 
morphology, phylogeny and classification, keys to the Holarctic genera, and 
remarks on bionomics and biogeography, including the biogeographic divisions 
of the Palaearctic recognised by the author. 


The text is generally well written, clear and concise, but the publishers should 
have had the English checked more thoroughly. Inexcusably, an index is not 
provided. The title of this book is rather misleading, as eastern Europe is 
generally taken to include countries such as Rumania and Bulgaria. This is 
here not the case and the vast majority of material studied has come from 
the western and southern parts of the former Soviet Union. Relatively little 
has been collected from the vast areas of central, northern and eastern Russia. 


The book will be of interest to all students of the Nepticulidae. Unfortunately, 
the high price (approx. £80) will deter many individuals from buying it. 


Steven WHITEBREAD 


138 


Nota lepid. 18 21) : 139-159 ; 29.11.1995 ISSN 0342-7536 


Révision des Scythrididae paléarctiques. 

VI. Les types de Scythrididae du Natural History 
Museum de Londres, du Muzeul de Historia Naturala 
«Grigore Antipa» de Bucarest et du Zoologisches 
Museum der Humboldt-Universität de Berlin. 
(Première partie) (1) 

(Lepidoptera : Gelechioidea) 


Pietro PASSERIN D’ENTREVES 


Dipartimento di Biologia Animale, Universita di Torino, Via Accademia Albertina 17, I-10123 
Torino (Italie) 


Summary 


The type material of some nominal species of Scythrididae, e.g. S. canetella 
Car., S. hyalinella Car., S. grandipennis porrectella Car., S. subflabella (Stt.), 
S. trinummulata Meyr., S. parachalcha Meyr., S. balanophora Meyr., S. 
taurella Car., S. triochrias Meyr., S. aristidella Rbl., S. erminea Wlsm., S. 
subaerariella (Stt.), S. tenuivittella (Stt.), S. insulella (Stgr.), S. compsias Wism. 
has been studied. The material is preserved at the Natural History Museum 
of London, at the Muzeul de Historia Naturala «Grigore Antipa» of Bucarest 
and at the Zoologisches Museum der Humboldt-Universitat, Berlin. Lectotypes 
of S. hyalinella Caradja, S. grandipennis porrectella Caradja, Butalis sub- 
flabella Stainton, S. parachalca Meyrick, S. erminea Walsingham, S. subae- 
rariella Stainton, B. tenuivittella Stainton, B. insulella Staudinger and S. 
compsias Walsingham are designated. B. subflabella Stainton, 1867 is a junior 
synonym of S. inclusella (Lederer, 1855), S. taurella Caradja, 1920 junior 
synonym of S. balanophora Meyrick, 1916 and S. erminea Walsingham, 1907 
junior synonym of S. tessulatella Rebel, 1903. The known geographical 
distribution of the species treated is given. 


Riassunto 


Vengono trattate le seguenti specie nominali appartenenti aglı Scythrididae : 
S. canetella Car., S. hyalinella Car., S. grandipennis porrectella Car., S. 


(!) Travail réalisé dans le cadre de la recherche «Biosistematica dei Lepidotteri Scythri- 
didae» MURST 60%. 


139 


subflabella (Stt.), S. trinummulata Meyr., S. parachalcha Meyr., S. balanophora 
Meyr., S. taurella Car., S. triochrias Meyr., S. aristidella Rbl., S. erminea 
Wilsm., S. subaerariella (Stt.), S. tenuivittella (Stt.), S. insulella (Stgr.) e S. 
compsias Wlsm. Il loro materiale tipico € conservato presso il Natural History 
Museum di Londra, il Muzeul de Historia Naturala «Grigore Antipa» di 
Bucarest e presso lo Zoologisches Museum der Humboldt-Universität di 
Berlino. Il materiale à rivisto sia dal punto di vista della nomenclatura che 
della tassonomia. I Lectotypi di S. hyalinella Caradja, S. grandipennis 
porrectella Caradja, Butalis subflabella Stainton, S. parachalca Meyrick, S. 
erminea Walsingham, S. subaerariella Stainton, B. tenuivittella Stainton, B. 
insulella Staudinger e S. compsias Walsingham sono scelti qui per la prima 
volta. Sono inoltre state riconosciute le seguenti nuove sinonimie: B. 
subflabella Stainton, 1867 € sinonimo recente di S. inclusella (Lederer, 1855) ; 
S. taurella Caradja, 1920 é sinonimo recente di S. balanophora Meyrick, 1916 ; 
S. erminea Walsingham, 1907 & sinonimo recente di S. tessulatella Rebel, 1903. 
Vengono anche fornite alcune indicazioni sulla distribuzione geografica. 


Zusammenfassung 


Diese Arbeit befaBt sich mit einigen Arten aus der Gattung Scythrididae, 
nämlich S. canetalla Car., S. hyalinella Car., S. grandipennis porrectella Car., 
S. subflabella (Stt.), S. trinummulata Meyr., S. parachalcha Meyr., S. 
balanophora Meyr., S. taurella Car., S. triochrias Meyr., S. aristidella Rbl., 
S. erminea Wlsm., S. subaerariella (Stt.), S. tenuivitella (Stt.), S. insulella 
(Stgr.) und S. compsias Wlsm. Das Typenmaterial dieser Arten wird im 
Natural History Museum in London, im Muzeul de Historia Naturala 
“Grigore Antipa” in Bukarest und im Zoologischen Museum der Humboldt- 
Universitat in Berlin aufbereitet. Zahlreiche Lektotypen werden festgelegt und 
einige neue Synonymien aufgezeigt. Soweit bekannt, wird die geografische 
Verbreitung angegeben. 


Introduction 


Au cours de trois différentes visites au Natural History Museum de 
Londres, entre 1979 et 1986, j'ai pu étudier la plus grande partie du 
matériel typique des espèces décrites par Zeller, Meyrick, Stainton, 
Walsingham et Turati qui y sont conservées. Et encore, grace a la 
courtoisie de M. Aurelian Popescu-Gorj du Muzeul d’ Historia Naturala 
«Grigore Antipa» de Bucarest, Roumanie, et de M. H. J. Hannemann 
du Zoologisches Museum der Humboldt-Universität de Berlin, Alle- 
magne, J'ai reçu et étudié depuis longtemps le matériel typique des 
taxa nominaux décrits par Aristide Caradja et par Otto Staudinger. 


140 


En ce qui concerne les matériaux conservés au Natural History Museum 
de Londres, les Lectotypi ont été choisis en accord avec K. Sattler 
(voir aussi Sattler, 1976: 89), responsable, à l’époque, de la Section 
«Microlépidoptères». En particulier pour les taxa nominaux décrits par 
Walsingham, on a suivi le critère suivant ; ayant remarqué que l’auteur 
anglais avait l’habitude d’étiqueter en décrivant une nouvelle espèce 
sur plus d’un exemplaire des deux sexes, l’un comme Typus & et un 
autre comme Typus Q, en les indiquant aussi dans la description ori- 
ginale, on a choisi comme Lectotypus l’exemplaire qui avait été publié 
le premier comme Typus. Celui-ci est du reste toujours reconnaissable 
dans la collection, étant marqué d’un numero d’ordre qui correspond 
à celui du travail original. 


Pour les espèces nominales décrites par Meyrick, celui-ci avait l’habitude 
d'indiquer, dans la publication originale, où il avait déposé le Typus, 
même s’il avait a disposition des Syntypi. Lorsqu’on a trouvé l’indication 
«Typus at the B.M.», le seul exemplaire ainsi indiqué éventuellement 
présent dans les collections de ce Musée a donc été considéré comme 
étant l’Holotypus. 


Une bonne partie des synonymies indiquées dans ce travail ont été 
personellement reconnues depuis longtemps pour la première fois, 
d’après l'étude du matériel typique de la plus grande partie des taxa 
nominaux de Scythrididae, conservés dans les principaux Musées et 
Collections européens. Toutefois dans quelques cas, leur publication 
a été faite par d’autres auteurs auxquels on les avait communiquées 
à plusieures reprises pour la rédaction de catalogues régionaux ou 
nationaux, ou encore de monographies (par ex. Leraut, 1980 ; Vives 
Moreno, 1984 ; Bengtsson, 1984 ; 1991). 


Pour abréger, on n’a pas cité ici en entier les dénominations des 
differents Musées et des Collections où les matériaux objet de la 
présente note sont conservés ; leurs sigles sont les suivants : 


BMNH.: Natural History Museum, Londres, Angleterre 

MGAB: Muzeul de Istoria Naturala «Grigore Antipa», Bucarest, Roumanie. 
MHNP : Muséum National d’ Histoire Naturelle, Paris, France 

NHMV : Naturhistorisches Museum, Vienne, Autriche. 

TMB : Termeszettudomanyi Muzeum Allattara, Budapest, Hongrie. 

UMO: University Museum, Oxford, Angleterre. 

ZMHB: Zoologisches Museum der Humboldt-Universität, Berlin, Allemagne. 


Scythris ventosella Chrétien, 1907 
Scythris ventosella Chrétien, 1907, Le Naturaliste, 29: 179. Locus typicus : 
Canet, Languedoc, France. Lectotypus G : «Canet 9.7.06»; «TYPE» ; 


141 


«ventosella» ; «Lectotypus Scythris ventosella Chrét. teste Passerin d’E. 
1975» ; «Prep. genitale 380 3 Passerin d’E. 1975». Choisi et publié par 
Passerin d’Entréves, 1976 : 30. MNHN. 


Scythris herniariae de Joannis, 1908, Bull. Soc. ent. France, 1908 : 248. Locus 
_ typicus : Plouharnel, St. Pierre Quiberon, Morbihan, France. Lectotypus 
@ : «TYPE» ; «Plouharnel 28 mai» ; «exemplaire dessiné» ; «Scythris her- 
niariae Joann.» ; «teste no. 182 H.» ; «Coll. L & J De Joannis Muséum 
Paris» ; «Lectotypus Scythris herniariae De J. teste Passerin d’E. 1975» ; 
«Scythris herniariae Joan. Bull. Soc. Ent. France 1908, p. 248». Choisi 
et publié par Passerin d’Entreves, 1976 : 30. MNHN. Synonymie établie 

et publiée par Passerin d’Entreves, 1976 : 30. 

Scythris canetella Caradja, 1920, D. ent. Z. Iris, 34: 144. Locus typicus : 
Canet, Pyrenées Or., France. Lectotypus & : «Canet 9.7.06» ; «Butalis 
Pyren. or.» ; «Lectotype & Scythris canetella Car. Des. A. Popescu- 
Gorj» ; «Prep. Genitale 2414 4 Passerin d’E.» ; «Scythris ventosella Chrét 
det. Passerin d’Entréves 1982». Choisi par Popescu-Gorj, publié ici pour 
la première fois. MGAB. Synonymie établie par Passerin d’Entrèves et 
publiée par Vives Moreno, 1984: 111. 

Scythris jugicolella Zerny, 1936, Mem. Soc. Sci. nat. phys. Maroc, 42 (1935) : 
148. Locus typicus : Tizi n° Tachdirt, Grand Atlas, Maroc. Lectotypus 
& : «Marokko, Gr. Atlas Tizi n° Tachdirt 3100-3200 m 21.-25.VII.33 
Zerny» ; «Scythris jugicolella Zerny Type 4»; «in coll. Hpt S. Wien 
ist jugicolella Zerny Type & E. Jäckh 1974 Prap. Ja 8276» ; «Lectotypus 
Scythris jugicolella Zerny Passerin d’E. 1980»; choisi et publié par 
Passerin d’Entréves, 1980 : 58. MNHN. Synonymie établie et publiée 
par Passerin d’E., 1976 : 30. 


ARMURE GÉNITALE MALE: Voir Passerin d’Entreves, 1976, Tab. 2a-c. 
ARMURE GENITALE FEMELLE : voir Passerin d’Entreves, 1976, Tab. 12c. 


Notes: Sur la base de la structure des genitalia des deux sexes, S. 
ventosella appartient au groupe «cicadella» tel qu’il a été défini par 
Passerin d’Entreves, 1986 : 220-221. Aux espèces indiquées dans cet 
article il faut à présent ajouter encore S. iconiensis Rebel. 


RÉPARTITION GÉOGRAPHIQUE : Espagne, France, Maroc (Haut-Atlas). 


Scythris hyalinella Caradja, 1920. 


Scythris hyalinella Caradja, 1920, Dt. ent. Z. Iris, 34: 146. Locus typicus : 
Kasikoparan, Turquie. Lectotypus © : «Kasikoparan Korb 1901» ; «4098 
Wlsm, 1903» ; «Scythris sp named by Wlsm»; «Lectotype ® Scythris 
hyalinella Car. Des. A. Popescu-Gorj» ; «Prep. genitale 2416 © Passerin 
d’E.» ; «Scythris hyalinella Car. det. Passerin d’E. 1982». Choisi par 
Popescu-Gorj et publié ici pour la première fois. MGAB. 


142 


ARMURE GÉNITALE MÂLE : inconnue. 
ARMURE GÉNITALE FEMELLE : Fig. 4A. 


Notes: Il est impossible, à présent, d’attribuer S. hyalinella Car. à 
l’un des groupes naturels déjà reconnus dans le genre Scythris. Il faudra 
attendre l'étude d’un matériel plus abondant comprenant aussi des 
mâles. 


RÉPARTITION GÉOGRAPHIQUE : Turquie. 


Scythris grandipennis (Haworth, 1828). 


Porrectaria grandipennis Haworth, 1828, Lepidoptera Britannica, 4: 536. 
Locus typicus : Angleterre. Lectotypus 4: «Lectotype» ; «Lectotype» ; 
«grandipennis» ; «Hope». Choisi et publié par Bradley, 1966 : 137. UMO. 


Scythris (Butalis) cuencella Rebel, 1901, Iris, 13 : 175. Locus typicus : Cuenca, 
Espagne. Lectotypus @: «Cuenca 1900 Stgr; «Prap. Ja 8205»; «In 
coll Hpt Wien als cuencella Rbl. ist grandipennis Hw.». Choisi et publié 
par Jackh, 1977 : 263. NHMW. Synonymie établie et publiée par Jackh, 
IST 265$: 

Scythris unicolorella De Joannis, 1909, Bull. Soc. ent. France: 90. Locus 
typicus : environs de Vannes, Morbihan, France. Lectotypus 4 : «TYPE», 
«Vannes 30 mai» ; «var. unicolorella Stgr. 1.1.» ; «Scythris gallicella var. 
unicolorella Joann. Bull. Soc. Ent. Fr., 1909, p. 90» ; «1920-1932 Coll. 
L. & J. De Joannis Muséum Paris» ; «Lectotypus Scythris unicolorella 
De J. st. nov. teste Passerin d’E. 1975» ; «Prep. genitale 399 À Passerin 
d’E. 1975». Choisi et publié par Passerin d’Entreves, 1976 : 40. MHNP. 
Synonymie établie et publiée par Passerin d’Entreves, 1976 : 40. 

Scythris grandipennis porrectella Caradja, 1920, Iris, 34 : 143. Locus typicus : 
Alfacar, Espagne. Lectotypus @ : «Sierra Alfacar» ; «Lectotype Scythris 
grandipennis var. porrectella Car. des Dr. A. Popescu-Gorj» ; «Prep. 
Genitale 1469 & Passerin d’E.». Choisi par Popescu-Gorj et publié ici 
pour la première fois. MGAB. 


ARMURE GÉNITALE MALE: voir Passerin d’Entréves, 1976, Tab. 6d-f. 
ARMURE GÉNITALE FEMELLE : voir Jäckh, 1977, Tab. 21, fig. 35. 

Notes: S. grandipennis (Haworth) appartient au groupe naturel 
«grandipennis» tel qu’il a été défini par Jäckh (1977: 262) et qui 


comprend dans l’état actuel de nos connaissances aussi S. nevadensis 
Passerin d’Entreves, en plus des espèces indiquées par l’auteur allemand. 


RÉPARTITION GÉOGRAPHIQUE : Angleterre, Espagne, France. L’espece 
a été signalée aussi d’Allemagne, de Suisse et du Portugal. 


143 


Fig. 1. A, B, C : armure génitale mâle de S. inclusella (Lederer) (PG 905 PE); D, 
E, F : armure génitale mâle de S. trinummulata Meyrick (PG 802 PE). A — complexe 
uncus-tegumen-gnathos-valve droite ; D — complexe uncus-tegumen-gnathos-valvae ; 
B, E — VIII": sternite ; C, F — édéage. 


145 


Scythris inclusella Lederer, 1855. 


Butalis inclusella Lederer, 1855, Verh. zool.-bot. Ges. Wien, 5 : 231. Locus 
typicus : Liban. Lectotypus : pas encore choisi n’ayant pu retrouver 
aucun exemplaire sûrement attribuable à la série typique de S. inclusella 
ede 

Butalis subflabella Stainton, 1867, The Tineina of Syria and Asia Minor : 
48. Locus typicus : Beyrouth, Liban ; Hebron, Palestine. Lectotypus ¢ : 
«Lectotype» ; «Beirut Hebron Palestine Cambridge 1865 Tristan 1897 
9213 Cotype & (1/3)»; «BM Genitalia Slide @ 22101»; «Lectotypus 
Butalis subflabella Stt. Passerin d’E. 1993». Choisi par Passerin d’Entreves 
et publié ici pour la première fois. BMNH. Syn. nov. 


ARMURE GENITALE MALE : Figs 1A-C. 
ARMURE GENITALE FEMELLE : Fig. 4B. 


Notes : Du moment que S. inclusella est bien connue, on peut établir 
ici la synonymie avec S. subflabella (Stt.), méme si on n’a pas encore 
pu étudier son matériel typique. Sur la base des caractéristiques des 
genitalia males et femelles, S. inclusella (Led.) appartient au groupe 
naturel «scopolella» qui comprend aussi: S. scopolella (F.), S. apicalis 
(Z.), et S. flabella (Mann). Ce groupe naturel est caractérisé, au niveau 
des genitalia, par les caractères (probablement synapomorphiques) 
suivants : tegumen subtriangulaire ; uncus bien développé, courbé cau- 
dalement ; gnathos très bien développé, avec un processus basal pointu, 
assez long ; valves symétriques, sauf chez S. scopolella ; édéage bien 
développé ; VIII*™* sternite portant une incision en profondite à l’extré- 
mité caudale. Femelle : le VIII" segment porte sur sa face ventrale 
une pièce sclérifiée. Antrum plus ou moin sclérifié. Chez quelques 
espèces, on trouve la lamella antevaginalis bien sclerifiee. 


RÉPARTITION GÉOGRAPHIQUE : Liban, Jordanie, Syrie, Israël. 


Scythris trinummulata Meyrick, 1922 


Scythris trinummulata Meyrick, Exotic Microlep., 2: 554. Locus typicus : 
Haifa, Palestine. Holotypus © : «Holotype» ; «Haifa Palestine V.1921 
P.A. Buxton 1922-107»; «Scythris trinummulata Meyr. Allotypus 9»; 
«BM Genitalia Slide 9 22099» ; «Holotypus Scythris trinummulata Meyr. 
Passerin d’E. 1981». BMNH. 


ARMURE GENITALE MALE: Figs 1 D-F. 
ARMURE GENITALE FEMELLE : Fig. 4C. 


NOTES : On a pu étudier aussi un Paratype mâle de S. trinummulata 
(BMNH, BM Genit. slide 22098). S. trinummulata Meyrick appartient 


146 


au groupe naturel «seliniellay dont font partie, à présent, aussi les 
espèces suivantes : S. seliniella (Z.), S. subseliniella (Hein. & Wocke), 
S. clavella (Z.), et S. villari Agenjo. Le groupe «seliniella» est caractérisé, 
au niveau des genitalia des deux sexes, par les caractères (probablement 
synapomorphiges) suivants : mâle : tegumen cylindrique, ou bien sub- 
cylindrique ; uncus dilobé, bref ; gnathos le plus souvent en forme de 
coupe, fondu avec le tegumen, avec un processus ventral, bref, crochu ; 
valves symétriques, séparées ou bien fondues médialement à la base : 
édéage cylindrique, allongé chez une lignée d’espèces, bref, trapu et 
robuste chez une autre lignée ; VIII*™ sternite bien developpé, laminaire, 
arrondi ou à peine incisé caudalement. Femelle : antrum plus ou moins 
allongé et sclérifié ; du VIII"® sternite s'élève ventralement un processus 
membraneux évaginable plus ou moins développé se terminant par une 
plaquette arrondie. 


RÉPARTITION GÉOGRAPHIQUE : Palestine. 


Scythris parachalca Meyrick, 1916. 


Scythris parachalca Meyrick, 1916, Exotic Microlepidoptera, 2: 14. Locus 
typicus: Alma Dagh, Asie Mineure. Lectotypus 4: «Lectotype» ; 
«Type»; «Alma Dagh Asia Minor J 06»; «Meyrick Coll. BM 1938- 
290» ; «Scythris parachalca Mey. Holotype 4»; «Lectotypus Scythris 
parachalca Mey. Passerin d’E. 1981». Choisi par Passerin d’Entreves 
et publié ici pour la première fois. BMNH. 


ARMURE GÉNITALE MÂLE : Figs 2A-C. 
ARMURE GÉNITALE FEMELLE : voir Jäckh, 1977, Tab. 23, Fig. 43. 


Notes : Les genitalia femelles de S. paracalcha ont été publiés pour 
la première fois par Jäckh, 1977, Tab. 23, Fig. 43, sous le nom de 
Scythris cupreella var. Staudinger. L’auteur allemand a toutefois re- 
connu qu'ils n’appartenaient pas à cette espèce, mais plutôt à une espèce 
du groupe «aerariella». Et en effet S. paracalcha appartient, sur la base 
des caractères des armures génitales des deux sexes, au groupe naturel 
«aerariella», défini par Passerin d’Entreves en 1982 : 80, qui comprend 
non seulement les espèces indiquées dans l’article, mais encore : S. corsa 
Passerin d’E., S. /hommei Bengtsson & Passerin d’E. et S. jaeckhi 
Bengtsson. 


RÉPARTITION GÉOGRAPHIQUE : Turquie. 
Scythris balanophora Meyrick, 1916. 


Scythris balanophora Meyrick, Exotic Microlep., 1916, 2 : 15. Locus typicus : 
Alma Dagh, Turquie. Lectotypus 3 : «Lectotype» ; «Type» ; «Alma Dagh 


147 


Asia Minor J. 06» ; «Meyrick Coll BM 1938-290» ; «Scythris balanophora 
Mey Holotype 4»; «BM Genitalia Slide & no 22100» ; «teste Passerin 
d’E. 1981». Choisi par Passerin d’Entreves et publié ici pour la première 
fois. BMNH. 

Scythris taurella Caradja, 1920. Dt. ent. Ztschr. Iris, 34 : 144. Locus typicus : 
Zeitun, Turquie. Lectotypus & : «Zeitun» ; «Lectotypus Scythris taurella 
Car. Des. A. Popescu-Gorj». Choisi par Popescu-Gorj et publié par 
Jackh, 1978 : 81. MGAB. Syn. nov.! 


ARMURE GÉNITALE MÂLE : voir Jäckh, 1978, Tab. 12, Figs 47,48. 
ARMURE GÉNITALE FEMELLE : voir Jäckh, 1978, Tab. 19, Fig. 76. 


Notes : Sur la base des caractéristiques des genitalia mâles et femelles, 
S. balanophora Meyr. appartient au groupe naturel «basistrigella» tel 
qu'il a été défim par Jackh (1978 : 80) et qui comprend à présent S. 
basistrigella (Stgr.) et S. balanophora Meyr. 


REPARTITION GEOGRAPHIQUE : Turquie. 


Scythris triochrias Meyrick, 1916. 


Scythris triochrias Meyrick, 1916, Exotic Microlepidoptera, 2: 15. Locus 
typicus : Taurus, Asie Mineure, Turquie. Holotypus ® pour monotypie : 
«Q»; «Taurus m. Asia minor 06»; «Holotypus Scythris Triochrias 
Meyr.». BMNH. 


Notes: L’Holotypus, le seul exemplaire attribué a cette espèce 
nominale qu’on ait pu étudier, est une femelle sans abdomen. C’est 
pourquoi on ne peut illustrer ici les genitalia des deux sexes. L’habitus 
de l’adulte ne permet malheureusement pas de résoudre ce problème, 
étant donné que beaucoup d’espèces de Scythris présentent les mêmes 
caractéristiques extérieures que S. triochrias, c’est-à-dire surtout une 
ligne horizontale et un point jaune, ou jaunätre, sur les ailes antérieures 
(voir Jäckh, 1978, tab. 1-28). 


Scythris aristidella Rebel, 1902. 


Scythris aristidella Rebel, 1902, Dt. ent. Z. Iris, 15 : 119, n° 19. Locus typicus : 
Ak Chehir, Turquie. Holotypus @ pour monotypie : «Ak Chehir» ; 
«Holotypus Scythris aristidella Rbl. Passerin d’E.» ; «Prep. genitale 4001 
& Passerin d’E.». MGAB. 


ARMURE GÉNITALE MALE: Tab. 2d-f. 
ARMURE GÉNITALE FEMELLE : Tab. 4d. 


Notes: S. aristidella appartient, sur la base des caractéristiques des 
armures génitales des deux sexes, au groupe «inertella» ainsi caractérisé : 


148 


> 
ges CG Ti = es 


Fig. 2. A, B, C: armure génitale mâle de S. parachalca Meyrick (PG 22097 BM) ; 
D, E, F: armure génitale mâle de S. aristidella Rebel (PG 1302 PE). A — complexe 
uncus-tegumen-valvae ; D — complexe uncus-tegumen-gnathos-valve droite ; B, E — 
VITE sternite ; C, F — édéage. 


149 


mâle : tegumen globeux, plus allongé le long de l’axe cephalo-caudal ; 
uncus bref, dilobé, asymétrique, fondu avec le tegumen, très sclérifié ; 
gnathos allongé, subcylindrique, se terminant par un croché dirigé 
ventralement ; valves laminaires plus ou moin dilatées ; édéage bref, 
tronconique, ou allongé, subcylindrique. VIII" sternite avec bords 
arrondis, laminaire, dilaté. Femelle : /amella antevaginalis bien déve- 
loppée, plus ou moins subtriangulaire, sclérifiée à l’apex. 


Dans l’état actuel de nos connaissances, le groupe ne comprend que 
S. inertella (Zeller) et S. aristidella Rebel. 


REPARTITION GEOGRAPHIQUE : Turquie. 


Scythris tessulatella Rebel, 1903. 


Scythris Tessulatella Rebel, 1903, Verh. zool.-bot. Ges. Wien, 53 : 414. Locus 
typicus : Guer-el-Alia, Hammada, Sahara, Algérie. Holotypus & pour 
monotypie: «Krauss Sahara Guer-el-Alia Hammada Nachtfana 
25.4.1893» ; «Scythris Tessulatella Rbl Type»; «in coll. Hpt S. Wien 
ist tessulatella Rbl & Type E. Jäckh 1974 Präp. Jä 8232». Choisi et 
publié par Passerin d’Entreves, 1980 : 49. NHMV. 

Scythris erminea Walsingham, 1907, Ent. mon. Mag., 43 : 8. Locus typicus : 
Biskra, Hammam-es-Salahin, Algérie. Lectotypus &: «Type H.T.»; 
«Biskra Algeria 30.1V.1903 Wlsm 96518» ; «Walsingham Coll. 1910-427» ; 
«Scythris erminea Wlsm Ent. Mo. Mag 1907 Type 6» ; «BM Genitalia 
Slide & 22105». Choisi par Passerin d’Entreves et publié ici pour la 
première fois. BMNH. Syn. nov. 


ARMURE GENITALE MALE: voir Passerin d’Entreves, 1980, Figs 4a-e. 
ARMURE GENITALE FEMELLE : inconnue 


REPARTITION GEOGRAPHIQUE : Algérie, Iran, Lybie, Tunisie. 


Scythris subaerariella (Stainton, 1867). 


Butalis subaerariella Stainton, The Tineina of Syria and Asia Minor, 1867: 
55, n° 5. Locus typicus: Bosz Dagh, Turquie. Lectotypus 4: «Bos 
dagh» ; «Coll. Led.» ; «Origin» ; «in coll Berlin als subaerariella Stt. 4 
det E. Jäckh 1975»; «Prep. genitale 2262 @ Passerin d’E. 1980» ; 
«Lectotypus Butalis subaerariella Stt. Passerin d’E. 1992». Choisi par 
Passerin d’Entreves et publié ici pour la première fois. ZMHB. 


ARMURE GENITALE MÂLE : Figs 3A-C. 
ARMURE GÉNITALE FEMELLE : jusqu’a présent inconnue. 


NOTES : S. subaerariella Stt. pourrait apartenir au groupe «obscurella», 
mais cette attribution nécessite un matériel plus abondant et appartenant 
aux deux sexes. 


150 


RÉPARTITION GÉOGRAPHIQUE : Turquie. 


Scythris tenuivittella (Stainton, 1867). 


Butalis tenuivittella Stainton, 1867, The Tineina of Syria and Asia Minor, 
1867: 55, n° 6. Locus typicus: Brussa, Turquie. Lectotypus 9: 
«Brussa» ; «Origin» ; «Lectotypus Butalis tenuivittella Stt. Passerin d’E. 
1993» ; «Prep. genitale 2261 Passerin d’E. 1980». Choisi par Passerin 
d’Entreves et publié ici pour la première fois. ZMHB. 


Butalis leucogaster Mann, 1872, Verh. zool.-bot. Ges. Wien, 22, 1-2 : 40. Locus 
typicus : Sangraben, Schneeberge, Sonnenschein, Autriche. Lectotypus 
& : «Type 1872»; «Butalis leucogastrella Mann»; «Holotypus Nr.» ; 
«Mus. Vind. Gen. Präp 4186 199 H.» ; «Lectotypus Butalis leucogaster 
Mann Passerin d’E. 1980» ; choisi et publié par Passerin d’Entreves, 
1980 : 46. NHMV. Synonymie établie par Passerin d’Entreves et publiée 
par Vives Moreno, 1984 : 111. 

Scythris constantella Dumont, 1925, Enc. Entm. Lepidoptera, I : 31. Locus 
typicus : Ile d'Oléron, Charente-Maritime, France ; Capbreton, Landes, 
France. Lectotypus & : «TYPE»; «Cistus salvifolius Ch. Capbreton 
20.5.85 souflée coll.on Ecl. 21.7.85»; «1911 Coll. Lafaury Muséum 
Paris» ; «Scythris constantella Dum. Enc. Entm. Serie B, III Lepidoptera, 
I, 1925, p. 31 (P. Viette VI. 1951)» ; «Lectotypus Scythris constantella 
Dum. teste Passerin d’E. 1975»; «Prep. genitale 412 & Passerin d’E. 
1975». Choisi et publié par Passerin d’Entreves, 1976: 41. MHNP. 
Synonymie avec leucogaster établie et publiée par Passerin d’Entréves, 
1976 : 42. Synonymie avec tenuivittella établie par Passerin d’Entreves 
et publiée par Vives Moreno, 1984 : 111. 

Scythris ruehli Gozmany, 1960, Annls hist.-nat. Mus. natn. hung., 52: 423. 
Locus typicus : Büyük Ada, Mer de Marmara, Turquie. Holotypus © 
pour désignation originale: TMB. Synonymie établie par Passerin 
d’Entreves et publiée par Leraut, 1980 : 73. Synonymies avec /eucogaster 
et tenuivittella établies par Passerin d’Entreves et publiées par Leraut, 
1980 : 73 et par Vives Moreno, 1984: I11. 


ARMURE GENITALE MALE : voir Passerin d’Entreves, 1976, Tab. 7a,b,c 
(sub. constantella). 
ARMURE GENITALE FEMELLE : voir Passerin d’Entreves, 1976, Tab. 14c 
(sub. constantella). 


NOTES : S. tenuivittella (Stt.) appartient au groupe naturel «fuscoaenea» 
tel qu’il a été défini par Bengtsson, 1991 : 405, et qui comprend, dans 
l’état actuel de nos connaissances les espèces suivantes : S. fuscoaenea 
(Hw.), S. parafuscoaenea Bngtss., S. vartianae Kasy, S. tenuivittella 
(Stt.), S. vittella (Costa) (= restigerella Z.), S. dissimilella (H.-S.), S. 
derrai Bngtss. et S. traugotti Bngtss. 


151 


152 


Fig. 3. A, B, C : armure génitale mâle de S. subaerariella (Stainton) (PG 2262 PE) ; 
D, E, F: armure génitale mâle de S. compsias Walsingham (PG 790 PE). A, 
D — complexe uncus-tegumen-gnathos-valve droite; B, E — VII" sternite; C, 
F — édéage. 


153 


154 


0,1 mm 
0,2mm 


Fig. 4. Armure génitale femelle de: A — S. hyalinella Caradja (PG 2416 PE) ; B — S. 
inclusella Lederer (PG 1494 PE) ; C — S. trinummulata Meyrick (PG 22099 BM) ; 


D — S. aristidella Rebel (PG 1495 PE). 


155 


RÉPARTITION GÉOGRAPHIQUE : Espagne, France, Italie, Sardaigne, 
Autriche, Roumanie, Crète, Turquie. 


Scythris insulella (Staudinger, 1859). 


Butalis Insulella Staudinger, Stett. ent. Ztg., 20 : 249, n° 108. Locus typicus : 
Chiclana, Andalusie, Espagne. Lectotypus @: «Erica Chiclana m.» ; 
«Origin» ; «teste H 108»; «Lectotypus Butalis insulella Stgr. Passerin 
d’E. 1990». Choisi par Passerin d’Entreves et publié ici pour la première 
fois. ZMHB. 

Scythris onychopyga Le Marchand, 1945, Rev. Frang. Lépid., 10 : 160. Locus 
typicus : Gazinet, Cestas, Gironde, France. Holotypus & par désignation 
originale teste Viette, 1954 : «TYPE» ; «Gazinet 12.VII.30» ; «Gironde» ; 
«onychopyga Le Md. Type» ; «TYPE» ; «Scythris onychopyga Le Md.» ; 
«Scythris onychopyga Le March. Rev. Franc. Lépid. 10, 1945, p. 160» ; 
«Muséum Paris Coll. S. Le Marchand 1953»; «Prep. genitale 355 @ 
Passerin d’E. 1975». Publié par Viette, 1954: 210 et par Passerin 
d’Entreves, 1976 : 43. MHNP. Synonymie établie par Passerin d’Entréves 
et publiée par Leraut, 1980 : 73. 


ARMURE GÉNITALE MALE: voir Passerin d’Entreves, 1976, Tab. 8a,b,c 
ARMURE GÉNITALE FEMELLE : jusqu’a présent inconnue. 


Notes: S. insulella (Stgr.) appartient au groupe naturel «insulella» 
qui, au niveau des caractères des genitalia est défini par les synapo- 
morphies suivantes. Mâle : VIII" sternite et VIIIwe tergite articulés 
latéralement pour former un complexe unique. Le VII" sternite, 
laminaire, bien développé, se présente incisé en V dans sa portion 
caudale, tandis que céphaliquement il présente deux zones articulaires 
avec le tegumen bien évidentes et renforcées par deux processus dirigés 
céphaliquement ; tegumen globeux fondu avec les valves ; uncus bien 
développé, dilobé ; gnathos très allongé, cylindrique, avec bras évidents, 
porte caudalement un processus crochu dirigé ventralement ; édéage 
bref, déprimé, dilaté dans sa portion medio-caudale ; valves subtrian- 
gulaires fondues médialement à la base et avec le tegumen. Femelle : 
Lamella postvaginalis considérable en forme de triangle avec l’apex 
dirigé caudalement. Le VIIème segment porte deux processus symétriques. 
Le groupe «insulella» comprend actuellement S. insulella (Stgr.) et S. 
ericivorella (Rag.). 


REPARTITION GEOGRAPHIQUE: Espagne, France. Signalée aussi du 
Portugal. 


156 


Scythris compsias Walsingham, 1907. 


Scythris compsias Walsingham, 1907, Entomologist’s mon. Mag., 43 : 9. Locus 
typicus : Biskra, Algérie. Lectotypus & : «Lectotype»; «Type H.T» ; 
«Biskra Algeria 16.1V.1903 Wlsm 96519» ; «Walsingham collection 1910- 
427» ; «Scythris compsias, Wlsm Ent. Mo. Mag 43 p 9 Type @ descr.» ; 
«Lectotypus Scythris compsias Wlsm Passerin d’E. 1986». Choisi par 
Passerin d’Entréves et publié ici pour la première fois. BMNH. 

Necrothalassia argilosella Amsel, 1935, Veröff. dt. Kolon.- u. Ubersee-Mus. 
Bremen, 1: 212. Locus typicus: Palestine. Holotypus‘: on n’a pas 
encore su retrouver l’Holotype, ni dans la collection Amsel, LNK, ni 
ailleurs. Synonymie établie et publiée par Amsel lui-même, 1940 : 52. 


ARMURE GÉNITALE MÂLE : Figs 3D-F. 
ARMURE GÉNITALE FEMELLE : Fig. 5. 


Notes : En l’absence du matériel typique de Necrothalassia argilosella 
Amsel, on considère ici comme valable la synonymie proposée par 


I 0,1 mm. 1 


\ \ 


Fig. 5. Armure génitale femelle de S. compsias Walsingham (PG 1806 PE). 


159. 


Amsel. Toutefois, dans l’attente d’une meilleure définition du genre 
Scythris, on continue à considérer compsias comme faisant partie du 
genre Scythris (cfr. aussi Landry, 1991 : 30). 


RÉPARTITION GÉOGRAPHIQUE : Algérie, Palestine. 


Bibliographie 


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Toten-Meer-Gebietes, nebst Beschreibung neuer palastinensischer Macro- 
und Microlepidoptera. Veröff. Dt. Kolon. u. Uebersee Mus. Bremen 
1 (2) : 203-221. 

AMSEL, H.G., 1940. Ueber alte und neue Kleinschmetterlinge aus dem 
Mittelmeer-Gebiet. Veröff. Dt. Kolon. u. Uebersee Mus. Bremen 3 (1): 
37-56. 

BENGTSSON, B. A., 1984. The Scythrididae (Lepidoptera) of Northern Europe. 
Fauna ent. scand. 13, 137 pp. 

BENGTSSON, B. A., 1991. Review of the fuscoaenea species-group of Scythris 
(Lepidoptera : Scythrididae). Entomologica scand. 21 : 405-414. 

BRADLEY, J. D., 1966. Type specimens of Microlepidoptera in the University 
Museum, Oxford, described by Haworth. Entomologist’s Gaz. 17 : 129- 
140. 

CARADJA, A., 1920. Beitrag zur Kenntnis der geographischen Verbreitung der 
Mikrolepidopteren des palaearktischen Faunengebietes nebst Beschrei- 
bung neuer Formen. Dt. ent. Z. Iris 34 : 75-179. 

CHRÉTIEN, P., 1907. Lépidoptères du Languedoc. Naturaliste 29 : 178-179. 

DE Joannis, J., 1908. Une nouvelle espèce de Scythris (Microlép.) des environs 
de Vannes. Bull. Soc. ent. France 1908 : 248-250. 

DE Joannis, J., 1909. Note sur une espèce de Scythris des environs de Vannes. 
(Lepidopt. Elachistidae). Bull. Soc. ent. France 1909 : 89-90. 

Dumont, C., 1925. Description de deux Lépidoptères nouveaux de France 
(Geometridae, Scythrididae) et d’une Pyrale tératologique (Phycitidae). 
Encycl. ent. Ser. B, II, Lepidoptera, I : 29-34. 

GozMANY, L., 1960. Records on Microlepidoptera. Annls hist.-nat. Mus. natn. 
hung. 52 : 423-428. 

HAWORTH, A. H., 1809-1828. Lepidoptera Britannica. Londini. 

JACKH, E., 1977. Bearbeitung der Gattung Scythris Hübner (Lepidoptera, 
Scythrididae). 1. Die “grandipennis-Gruppe”. Dt. ent. Z. 24 (1-3) : 261- 
2H IMDb) 

JAckH, E., 1978. Bearbeitung der Gattung Scythris Hübner (Lepidoptera, 
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Dt. ent. Z. 25 (1-3) : 71-89, 21 Tab. 

LANDRY, J.-F, 1991. Systematics of Nearctic Scythrididae (Lepidoptera : 
Gelechioidea) : Phylogeny and classification of supraspecific taxa, with 
a review of described species. Mem. ent. Soc. Can. No. 160, 341 pp. 

LEDERER, J., 1855. Beitrag zur Schmetterlings-Fauna von Cypern, Beirut und 
einem Theile Klein-Asiens. Verh. zool.-bot. Ges. Wien 5 :177-234, 5 Taf. 


158 


LE MARCHAND, S., 1945. Un nouveau Scythris de France: Scythris ony- 
chopyga n. sp. (Scythrididae). Revue fr. Lépidopt. 10 (1944) : 158-161. 

LERAUT, P., 1980. Liste systématique et synonymique des Lépidoptères de 
France, Belgique et Corse. Alexanor, suppl., 334 pp. 

Mann, J., 1872. Beschreibung sieben neuer Arten Microlepidopteren. Verh. 
zool.-bot. Ges. Wien 22 (1-2) : 35-40 

Meyrick, E., 1912-1937. Exotic Microlepidoptera. 5 vol. Marlborough. 

PASSERIN D’ENTREVES, P., 1976. Revisione degli Scitrididi (Lepidoptera, 
Scythrididae) paleartici. II. — I tipi di Scythris del Muséum National 
d’Histoire Naturelle di Parigi. Boll. Mus. Zool. Univ. Torino 3 : 27-70. 

PASSERIN D’ENTREVES, P., 1980. Revisione degli Scitrididi (Lepidoptera, 
Scythrididae) paleartici. V. — I tipi di Scythris del Naturhistorisches 
Museum di Vienna. Boll. Mus. Zool. Univ. Torino 5 : 41-60. 

PASSERIN D’ENTREVES, P., 1982. Note su alcuni Scitrididi della fauna italiana 
(Lepidoptera, Scythrididae). Boll. Mus. Zool. Univ. Torino 6 : 79-86. 

PASSERIN D’ENTREVES, P., 1986. Quatre nouvelles espèces paléarctiques du 
genre Scythris (Lepidoptera, Scythrididae). Nota lepid. 9 (3-4) : 216-224. 

REBEL, H., 1901. Neue palaearctische Tineen. Dt. ent. Z. Iris 13 (2): 161- 
188. 

REBEL, H., 1902. Neue Pyraliden, Pterophoriden und Tineen des palaeark- 
tischen Faunengebietes. Dt. ent. Z. Iris 15 : 100-126. 

Rese, H., 1903. Eine Heteroceren-Ausbeute aus der Sahara II. Verh. zool.- 
bot. Ges. Wien 53 : 404-415. 

SATTLER, K., 1976. A Taxonomic revision of the genus Ornativalva Gozmany, 
1955 (Lepidoptera : Gelechiidae). Bull. Br. Mus. nat. Hist. (Ent.) 34 (2): 
87-152. 

STAINTON, H. T., 1867. The Tineina of Syria and Asia Minor. 84 pp. Van 
Voorst, London. 

STAUDINGER, O., 1859. Diagnosen nebst kurze Beschreibung neuer Andalu- 
sischer Schmetterlinge. Stettin. ent. Ztg. 1859 : 211-259, n° 108. 

VIETTE, P. E. L., 1954. Les travaux de S. Le Marchand. Revue fr. Lépidopt. 
15-17 : 204-210. 

Vives Moreno, A., 1984. Lista actualizada de la Familia Scythrididae Rebel, 
1901, de España y Portugal. SHILAP Revta. lepid. 12 (46): 109-113. 

WALSINGHAM, Lord, 1907. Algerian Microlepidoptera. Entomologist’s mon. 
Mag. 18 (43) : 6-10. 

ZERNY, H., 1936. Die Lepidopteren-fauna des Grossen Atlas in Marokko und 
seiner Randgebiete. Mém. Soc. Sci. nat. phys. Maroc 42(1935) : 1-163. 


159 


Nota lepid. 18 (2) : 160 ; 29.11.1996 ISSN 0342-7536 


Book reviews - Buchbesprechungen - Analyses 


Die Schmetterlinge Baden-Württembergs. Band 3 & 4, Nachtfalter I 
& II. Gunter EBERT (Ed.). 518 pp., 344 col. Figs, 166 diagrams, 64 
distribution maps (Vol. 3) + 535 pp., 488 col. Figs, 204 diagrams, 112 
distribution maps (Vol. 4), 18 x 24.5 cm, hardback. Verlag Eugen 
Ulmer, 1994. ISBN 3-8001-3472-1 & 3-8001-3474-8. Price/ Vol. : DM 79. 


This series deals with the butterflies and moths of Baden-Wiirttemberg in 
south-west Germany. The butterflies, including the skippers, were covered in 
volumes | & 2, published in 1991. The series is now continued with the larger 
moths, Vol. 3 treating the families Hepialidae, Cossidae, Zygaenidae, Lima- 
codidae, Psychidae and Thyrididae, and Vol. 4 the Bombycidae, Endromidae, 
Lasiocampidae, Lemoniidae, Saturniidae, Sphingidae, Notodontidae, Dilobi- 
dae, Thaumetopoeidae, Drepanidae, Thyatiridae, Lymantriidae, Nolidae and 
Ctenuchidae. The Arctiidae and Sesiidae will follow in Vol. 5 and the Noc- 
tuidae are planned for Vols 6 & 7. 


No serious European lepidopterist should be without this series in his library, 
even if his knowledge of German is weak or non-existent (a few pages of 
explanations in English and French are included). I know of no other work 
giving so much detail for most of these families. Information is given on 
distribution (global, regional — with map, and vertical), phenology (adult and 
early stages), ecology (biotope, foodplants of the larva and adult, microhabitat, 
behaviour and parasites) and red list status. The colour photographs are all 
of a very high standard and depict living adults and early stages of most 
species, their habitats, foodplants, parasites etc. 


Introductory chapters in Vol. 3 include the following interesting topics : 
Methods for observing moths (the word “collecting” is carefully avoided !), 
aspects of competition for food among moths, categories and lists of 
endangered species. An index to taxa is provided for each volume, but the 
comprehensive list of references is given at the end of vol. 4 only. A subject 
index is not given. 


This is not an identification work and no descriptions of the adults or larvae, 
or colour identification plates will be found. However, in a few cases, dis- 
tinguishing characters of close species are illustrated, including a few genitalia 
figures. The classification and nomenclature used by the various authors are 
very heterogeneous, but this does not detract from the work. 


The work provides an excellent basis, so urgently required, for the further 
study of these families, not only in Baden-Wiirttemberg, but also in the rest 


of Europe. 
Steven WHITEBREAD 


160 


Nota lepid. 18 (2) : 161-177 ; 29.11.1996 ISSN 0342-7536 


The distribution of the genus Tarucus Moore, [1881], 
in the Maghreb States of Morocco, Algeria and Tunisia, 
with notes on species identification 

(Lepidoptera, Lycaenidae) 


John TENNENT 


1 Middlewood Close, Fylingthorpe, Whitby, North Yorkshire YO22 4UD 


Summary 


As part of a wider study of butterflies in the Maghreb States of Morocco, 
Algeria and Tunisia, a large amount of material of the genus Tarucus Moore, 
[1881], was examined with a view to determining the distribution of 7: 
theophrastus (Fabricius, 1793), T. rosaceus (Austaut, 1885), and 7: balkanicus 
(Freyer, 1845), in the Maghreb and to assess whether there were any external 
features which might allow the separation of species. Results indicate that 
published information relating to distribution, identification and host-plant 
association is often unreliable. The apparently local and scarce T. balkanicus 
may be identified with a high degree of certainty from wing markings, but 
T. theophrastus and T: rosaceus cannot be reliably separated other than by 
examination of the male genital armature. Examination of a large number 
of male genitalia established that a high percentage of populations sampled 
contained both TZ: theophrastus and T. rosaceus. It is suggested that the two 
species fly together more often than they do not, utilising whichever Zizyphus 
or Paliurus (Rhamnaceae) host-plants are available. 


Résumé 


En vue d’une meilleure connaissance des Lépidoptères des pays maghrébins 
du Maroc, d'Algérie et de Tunisie, une quantité substantielle de matériel du 
genre Tarucus Moore, [1881], a été réunie et examinée dans le but de préciser 
les répartitions de 7! theophrastus (Fabricius, 1793), T. rosaceus Austaut 1885 
et T: balkanicus (Freyer, 1845), au Maghreb et de savoir si quelques caractères 
externes pourraient permettre une différenciation spécifique. Les résultats 
montrent que les données déjà publiées quant à leurs distributions, identi- 
fications et plantes-hötes ne sont pas toujours fiables. 7: balkanicus, appa- 
remment localisé et rare, peut-être identifié avec forte certitude par ses dessins 
alaires, alors que 7: theophrastus et T. rosaceus ne peuvent être différenciés 
que par l'étude de l’armature génitale male. Un grand nombre de genitalia 
ont été examinés et il en ressort qu’un fort pourcentage des populations 
étudiées contient a la fois les deux espèces T° theophrastus et T: rosaceus. 


161 


Ces deux espèces volent donc conjointement plus fréquemment qu’on ne le 
pense, recourant tout autant au Zizyphus qu’au Paliurus (Rhamnaceae) 
comme plantes nourricières certaines. 


Zusammenfassung 


Als Teil einer umfassenden Studie der Tagfalterfauna der Maghreb Staaten 
von Marokko, Algerien und Tunesien, wurde auch sehr umfangreiches 
Material von Arten der Gattung Tarucus Moore, [1881], zusammengetragen. 
Dieses soll nun vorab untersucht und analysiert werden um die Verbreitung 
der Arten T: theophrastus (Fabricius, 1793), 7: rosaceus (Austaut, 1885) und 
T. balkanicus (Freyer, 1845), in der Maghreb-Zone aufzuzeigen. Darüber 
hinaus soll festgestellt werden ob es irgendwelche äußerliche Merkmale gibt, 
die es erlauben, diese Arten danach zu unterscheiden. Die gewonnenen 
Ergebnisse deuten darauf hin, daß die bisher publizierten Angaben, die 
Verbreitung, die Identifizierung der Arten und deren Futterpflanzen betreffend, 
oft unglaubwürdig erscheinen. Die offensichtlich lokale und seltene T. bal- 
kanicus kann, mit einem großen Maß an Sicherheit, aufgrund der Flügel- 
zeichnung, bestimmt werden. T. theophrastus und T. rosaceus hingegen, 
können nicht verläßlich, ohne Berücksichtigung ihrer Genitalien, voneinander 
getrennt werden. Eine große Zahl männlicher Genitalien wurde untersucht, 
wodurch gezeigt werden kann, daß ein hoher Prozentsatz der herangezogenen 
Populationen sich aus T. theophrastus und T. rosaceus zusammensetzt. 
Vermutlich fliegen diese beiden Arten öfters zusammen als getrennt, wobei 
sie entweder, je nach dem Vorkommen, Zizyphus oder Paliurus (Rhamnaceae) 
also Ablagepflanze annehmen. 


Introduction 


The Lycaenid genus Tarucus comprises ca 25 species distributed from 
north-western Africa to South East Asia. Males of many species are 
blue on the upperside (ups) and the females are brown with a variable 
pattern ; the underside (uns) of both sexes is creamy white with a 
pattern consisting of brown or black spots, streaks and lines. The 
separation of species is notoriously difficult using external features. In 
India and the Middle East, the butterflies inhabit very dry localities 
and the genus reaches its western limits in the semi-desert regions of 
south-western Morocco. In the Maghreb States three species fly: T. 
theophrastus (Fabricius, 1793), 7: rosaceus (Austaut, 1885), and 7: 
balkanicus (Freyer, 1845). 


Bethune-Baker (1918) published a revision of the genus world-wide. 
He did not give the origin of material examined but confined himself 
to general comments on distribution, describing 7: mediterraneae, a 
synonym for T: rosaceus. In 1955, Evans revised the Zarucus of Europe, 


162 


North Africa and Asia, examining 4000 specimens, mainly in the 
collection of The British Museum (Natural History), London 
(BM(NH)), and listed the provenance of the material examined. The 
only other report dealing specifically with Tarucus in the Maghreb 
States was that of Baz (1988), who drew certain conclusions regarding 
the distribution of 7: theophrastus in Morocco based on material 
collected at nine sites during two weeks in May 1987. In the light 
of the results of the present study, it would be surprising if many of 
these ‘theophrastus’ are not in fact rosaceus. 


One of the few contemporary butterfly books comprehensively incor- 
porating north-western Africa is Higgins & Riley’s “Butterflies of 
Britain and Europe” which 1s, as the author has commented previously 
(Tennent, 1993b : 257), often flawed so far as the North African fauna 
is concerned. In that book, T! theophrastus was described as flying 
around the host-plant, Zizyphus vulgaris (Rhamnaceae), usually in 
lowlands near the coast of North Africa ; T. rosaceus and T. balkanicus 
were also said to be found in the lowlands, probably to the foothills 
[of the Atlas mountains], flying near the host-plant, Paliurus spinachristi 
(Rhamnaceae). Maps accompanying Higgins & Riley’s text showed 
the North African distribution of all three species limited to the 
Mediterranean coastal region, with a further isolated population of 
T. rosaceus in western Morocco. 


In endeavouring to determine the distribution of Tarucus species in 
North Africa for a comprehensive work (Tennent, 1996 in prep.), the 
author experienced practical difficulty in separating T. theophrastus 
and T° rosaceus in the field. The suspicion that the authenticity of 
some previously published records was in doubt and the knowledge 
that Tarucus species extend to considerable altitudes in the High Atlas 
mountains as well as deep into the Sahara desert, prompted the present 
review. 


Material examined 


During field work in 1991-1994, a total of 303 male and 208 female 
Tarucus specimens was collected from 46 different localities in Morocco, 
Algeria and Tunisia. The study was seriously curtailed by the political 
situation in Algeria. The loan or gift of a further 14 male specimens 
was received from the Zoologisk Museum, Copenhagen (DK) and from 
entomologists in Marktleuthen (D), Bonn (D), San Gimignano (I) and 
Cannes (F). The comprehensive collections in the BM(NH), London, 
were also examined but dissection of specimens not forming part of 
the author’s accumulated material was deemed unnecessary. 


163 


Collecting was arbitrary and the number of Tarucus specimens obtained 
from a locality is not indicative of abundance. The combined total 
of 317 male Tarucus from 58 localities in the Maghreb available for 
genitalic examination provided the data on which this study is based ; 
unless otherwise stated, material is in the author’s collection. 


Localities from which Zarucus material was available (see Fig. 1) 


T1 — 566 — Tafraoute, Anti-Atlas (M), 1300 m, 13.9.91 

T2 — 1766, 798  — Sidi M’Zal (M), ca. 1300 m, 3-6.4.92 / 10.5.92 / 18.4.94 

T3 — 264 — Agadir (M), 80 m, 19.4.94 

T4 — 664,492  — Ait Moussa, 29 km E of Agadir (M), 100 m, 28.5.93 

T5 — 988,422  — Taroudannt (M), 250 m, 12.9.91 

T6 — 664,399  — Ait-lazza, E of Taroudannt (M), 300 m, 11.5.92 / 
26.5.93 

T7 — 584,292 —4 km N of Olad Berhil (Taroudannt) (M), 620 m, 
26.5.93 

T8 — 544 — 12-14 km W of Aoulouz (between Taroudannt and 


Taliouine) (M), 740 m, 26.5.93 / 31.3.94 


T52ee T53 


Fig. 1. Location of sites T1-T58. 


164 


T9 — 19 — Tizi-n-Test (M), 1100 m, 4.6.92 


T10 — 18 — nr Asni (M), 1050 m, 6.9.91 

T11 — 18, 299 — Oukaimeden (M), 2400 m, 3,5.9.91 / 14.6.94 
T12 — 446 — Barrage de Lalla-Takerkoust (M), 600 m, 31.5.94 
113 — 244 — Amizmiz (High Atlas) (M), 900 m, 1.6.94 

T14 — 788, 722 — Ouriki, SE of Marrakech (M), 820 m, 24.6.93 
T15 — 546, 19 — Marrakech (M), 500 m, 31.5.94 


T16 — 488,229 — El Kelaa-des-Sraghna (M), 600 m, 1.9.91 

T17 — 1288, 999 — Khemis-des-Oulad-Ayad (El Kelaa-des-Sraghna) (M), 
780 m, 10-11.7.93 

T18 — 438 — 2 km S of Nid de Cigogne (El Kelaa-des-Sraghna) (M), 
420 m, 10.7.93 / 27.5.94 

T19 — 5188, 1599 — Bzou (ca. 30km E of EI Kelaa-des-Sraghna) (M), 
400 m, 9-10, 20.4.94 

T20 — 488,222 —3 km SW of Ouaoumara, S of Khenifra (M), 820 m, 
19.6.93 

T21 — 366,289 —R.P. 33, 11 km E of junction with R.24, (Khenifra) 
(M), 1050 m, 19.6.93 

T22 — 744, 622 — Khenifra (M), S outskirts of town, 900 m, 3.6.93 

T23 — 544,722 — Tizi-n-Bachkoum (M), 1400 m, 10.9.91 / 14.3.94 


T24 — 14 — nr. Bou-Azzer (Anti-Atlas) (M), 1320 m, 26.3.94 
T25 — 1644, 299 — Meski (Er Rachidia) (M), 1100 m, 1.4.92 / 25.5.93 
T26 — 14, 19 — N outskirts of Er Rachidia, 1150 m, 25.5.93 


T27 — 388,222 — 30 km N of Er Rachidia (M), 1340 m, 25.5.93 
T28 — 668,499 — 12km S of Rich, Ziz valley (M), 1400 m, 25.5.93 


T29 — 14 — Tafilalt, vic. Aoufouss (M), 1000 m, 3.6.90, W. ten 
Hagen les. 

T30 — 14 — R.P. 24, 22 km S of Azrou (M), 1100 m, 3.6.93 

T31 — 18, 19 — nr Sefrou (M), 900 m, 27.8.91 


T32 — 1038, 1189 — 9 km S of Mechra-Bel-Ksiri (M), 100 m, 6.6.93 
133 — 2288, 799 — nr Ouezzane (M), 120 m, 23.8.91 


T34 — 538 — 8 km W of Guercif (M), 480 m, 13.6.93 

T35 — 388 — 20 km E of Guercif (M), 600 m, 24.4.92 

T36 — 438 — 7 km NE of El Aïoun (M), 650 m, 13.6.93 

T37 — 388, 19 — 6 km E of Driouch (Nador) (M), 380 m, 25.5.93 

138 — 14, 19 — 10 km W of Berkane, N slopes of Monts de Beni- 
Snassen (M), 380 m, 14.5.93 

T39 — 244, 19 — Berkane (M), 300 m, 14.6.92 

T40 — 238 — near El Aïoun, Oujda (M), 500 m, 22.5.82, F. Fabiano 
(coll. Fabiano) 

T41 — 14 — Commune d’el Aouinet, 24 km S of Oujda (M), 880 m, 
13.5.93 

T42 — 844, 299 — 10 km SW of Relizane (A), 300 m, 11.5.93 

143 — 14 — Gorges de la Djemaa, env. d’Alger (A), [ ? alt], 30.06.55, 
G. Barragué leg. 

T44 — 14 — Rassauta, dep. d’Alger (A), [ ? alt], 13.9.57, G. Barragué 
leg. 


165 


T45 — 14 — Lalla Marnia (A), [ ? alt], 15.5.15, Dr H. C. Nissen 
leg. (coll. Zool. Museum Copenhagen) 

T46 — 238 — Guelt-es-Stel (A), [ca. 1100 m], 26.8.51, G. Barragué 
leg. (coll. Barrague) ; 14.9.17, [Dr H. C. Nissen leg.] 
(coll. Zool. Museum Copenhagen) 

T47 — 16 — Ain Sefra (A), [ ? alt], 17.5.13, [Dr H. C. Nissen leg.] 
(coll. Zool. Museum Copenhagen) 

T48 — 3288, 1199 — EI Kantara (A), 6-900 m, 30.6.92 / 17-20.4.93 


T49 — 18 — Col de Sfa [N of Biskra] (A), [ ? alt], 15.4.09, Dr. H. C. 
Nissen leg. (coll. Zool. Museum Copenhagen) 

T50 — 14 — 3-5km NW of Biskra (A), [ ? alt.] 17.3-5.4.84, H. 
Falkenhahn leg. [ex. entomologisches museum Dr. Ulf 
Eitschberger] | 

TS1 — 14 — Ghardaia (A), [ ? alt], 20.4.11, Dr H.C. Nissen leg. 
(coll. Zool. Museum Copenhagen) 

T52 — 14 — 12 km E of Source Chapuis, Hoggar mountains 
(Tamanrasset) (A), 1540 m, 28.4.93 

T53 — 744, 19 — Source Chapuis, Hoggar mountains (Tamanrasset) 
(A), 1540 m, 27/28.4.93 

T54 — 14 — 3-5 km SW of Gafsa (T), 400 m, 4.3.84, H. Falkenhahn 
leg. [ex. entomologisches museum Dr. Ulf Eitschberger] 

T55 — 288, 19 — Dj. Hachichina, SE of Gafsa (T), 100 m, 5.4.93 

T56 — 664, 422 —Dj. Sif El Leham, 12 km N of T55 (1), 300 m, 6.4.93 

T57 — 288 — El Amayem, 16 km E of El Guetter (1), 30.3.93 / 
7.4.93 

T58 — 14 — 15km E of Matmata (T), 100 m, 6.4.91, W. ten Hagen 
leg. 


The distribution of Tarucus species in the Maghreb 


The primary aim of this study was to determine the distribution of 
the three Tarucus species within the political boundaries of Morocco, 
Algeria and Tunisia. As will be discussed, although balkanicus may 
usually be readily identified using external features (1.e., wing markings), 
theophrastus and rosaceus may not and it was necessary to examine 
the genitalia of 170 male theophrastus and rosaceus in order to gain 
knowledge of their distribution. The results are reproduced in Tab. 1. 


Species confirmed (theophrastus and rosaceus) from the 38 localities 
from which more than one male was available are recorded in Tab. 2. 


Although theophrastus and rosaceus were found flying together in 
almost two-thirds of the localities from which Tab. 2 was produced, 
it does not necessarily follow that, where only one species was identified, 
the other does not also occur. Indeed, in 26 localities sampled from 
where six or less males were available, both species were found, 


166 


Table 1 


Male T. theophrastus and T: rosaceus 
identified from genitalic examination. 


Locality Males examined rosaceus theophrastus 


[1 GAANDe NN DD 


— 
Ii NO m1 
mi mn © DB BR = ON RRS NNR KI QI ND NRK 


Un nu (99 nu pt nt pd mt | 
| 


— 
— 


Ln mm I 


— 
ı O1 mm NO En ni ni | ©) À nu nu © Jr 1! 


— 


3 
4 
2 
2 
3 
3 
5 
3 
l 
l 
3 
2 
2 
2 
3 
9 
2 
6 
4 
3 
2 
2 
1 
2 
l 
3 
2 
l 
l 
l 
7 
0 
2 
À 
4 
3 
l 
2 
2 
1 
8 
l 
1 
1 
2 
1 
l 
1 


167 


Table 1 (cont.) 


Locality Males examined rosaceus theophrastus 


PoreeoenNnoenee ft 


BNO HI ı mm 


1 
1 
1 
7 
1 
2 
6 
2 
1 


including 3 out of the 7 localities from where only two males were 
available. The main finding of this study is that the two species fly 
together in most localities and it may be inferred that they do so in 
most if not in all localities. Higher altitudes of the Moroccan High 
Atlas and desert regions of southern Algeria may prove to be an 
exception to this assumption. All 8 males examined from the Hoggar 
mountains in southern Algeria, proved to be theophrastus and the 
author knows of no confirmed record of rosaceus from that region. 
Speidel & Hassler (1990 : 115-116), in briefly discussing Tarucus species 
in the Hoggar, listed both theophrastus and rosaceus but accepted that 
only the former was confirmed from there. There are several balkanicus 
from southern Algeria in the BM(NH) collections and Stempffer (in 
Chopard & Villiers, 1950 : 269) recorded both mediterraneae |rosaceus] 
and balkanicus from the Air region of northern Niger, directly south 
of the Hoggar. 


Judging from the results of this study and from other material available, 
balkanicus appears to be a very local butterfly in the Maghreb and 
its true distribution is still not clear. It obviously flies in Mediterranean 
coastal regions (although the author has not actually seen material 
from there) as well as deep into the Sahara desert. Data incorporated 
in Fig. 2 for this species has been gleaned from material in the BM(NH), 


Table 2 


Presence of 7: theophrastus and T. rosaceus in localities 
from where more than one male was available. 


Localities Percentage 


Both species present 23 60.5 


T. theophrastus only 10 26.3 
T. rosaceus only 5 1622 


168 


including that examined by Bethune-Baker and Evans in their respective 
generic reviews, confirmed by them through genitalic examination, and 
should not be taken as an attempt to provide a comprehensive review 
of the distribution of that species in North Africa. The genitalia of 
none of these specimens have been examined by the author. By 
comparison, both theophrastus and rosaceus are widespread throughout 
the region up to 1300-1400 m, wherever suitable host-plants occur. 
Above that altitude, Zarucus spp. are less frequent, presumably due 
to a lack of host-plant availability. Geographical localities of specimens 
identified during this study are plotted on Fig. 2. 


The identification of Tarucus species in north-western Africa 


The secondary aim of this study was to assess the usefulness of external 
features in allowing species identification with certainty, without the 
necessity for genitalic examination. Previous authors have differed on 
the importance of reported features for the identification of specimens 
(see Bethune-Baker, 1918 ; Evans, 1955). The present author’s results 
show that, although 7. balkanicus may be identified with a degree 


Ak GS ie A 


T. theophrastus/ T. rosaceus flying together 


T. theophrastus only 
T. rosaceus only 
T. balkanicus (in BM (NH) etc.) 


Fig. 2. Identification of Tarucus spp. from study sites. 


169 


of success from wing markings alone, external features (1.e., wing 
colouration and markings) are unreliable for separating T. theophrastus 
and 7: rosaceus in North Africa and are, at best, only moderately 
useful as a guide. Additionally, since the two species apparently fly 
together more often than they do not, host-plant association is even 
more unreliable as a means of species identification. The only certain 
method of separating theophrastus and rosaceus is by genitalic exa- 
mination. The features most frequently given for identification are 
critically examined. 


External features 


Overall size 


Forewing length (fwl), measured from the wing apex to the base, of 
more than 200 male Tarucus specimens examined from North Africa 
is given in Tab. 3; it should be noted that overall size is variable in 
relation to local ecological conditions with, as might be expected, eremic 
populations being generally slightly smaller. The average fwl of 7 male 
theophrastus from the Hoggar mountains in southern Algeria is 
10.12 mm. It will be seen that overall size does not vary to any 
significant degree and is of no value in species identification. 


Table 3 
Male forewing length (fwl) of Tarucus species in North Africa. 


Species Males examined Smallest Largest Mean + SD 


T. balkanicus 
T. theophrastus 
T. rosaceus 


Upperside wing markings 


It transpired that the author’s material from north-western Africa 
contained only T. theophrastus and T. rosaceus and it was apparent 
from examination of material contained in the collections of the 
BM(NH) that 7: balkanicus is the only one of the three species under 
consideration which may be identified with any certainty using external 
features. The most obvious feature is the presence of prominent black 
postdiscal (pd) and discoidal spots on the upperside forewing (upf) 
and the author has not seen any specimens of balkanicus in which 


170 


this feature is lacking, although Larsen (1983 : 398) noted that these 
spots are sometimes absent in material from the Middle East. Conver- 
sely, 1 male theophrastus and 4 male rosaceus from localities T6, T21, 
T38, T50 and T57 displayed (aberrant) upf black markings similar to 
those of T! balkanicus and a number of other specimens appeared 
to display these features, although on closer examination it was found 
that this was due to underside (uns) markings showing through to the 
upperside (ups). 


Higgins & Riley (1983), also gave the size and shape of the upf discoidal 
spot as a useful guide to separation, stating (1983 : 57-58) that in 
theophrastus the spot is “dark and oblong”, “more linear and less 
conspicuous” in rosaceus and “obvious and dark” in balkanicus. The 
size and shape of the upf discoidal spot in North African material 
examined is immensely variable and is of no value for identification 


purposes. 


Male upperside colour 


The original description of 7: rosaceus (Austaut, 1885 : 141-142) was 
based on “a dozen” males from “high situations” in Algeria and 
Morocco which, among other features, were said to be smaller than 
theophrastus, rosy-blue on the ups rather than dark-blue violet and 
with slightly diaphanous wings allowing the uns markings to show 
through to the ups. Bethune-Baker (1918 : 277, 281, 286) stated that 
the male ups ground colour (gc) is “deep lustrous violet” in balkanicus, 
“Tilac-blue” in [rosaceus] and “violet-blue or lilac-blue” in theophrastus. 
D’Abrera (1993 : 470) stated that rosaceus maybe distinguished from 
its congeners by the fact that the ups purple colour has “hints of pink 
in it” and Higgins & Riley (1983 : 58) stated that rosaceus “resembles 
theophrastus but with a faint pinkish tint”. Larsen (1983 : 397) stated 
that, in the case of rosaceus (in Saudi Arabia), “the lilac-violet ground 
colour distinguishes it from [theophrastus and balkanicus]’ and a 
number of other authors have made similarly conflicting statements. 


It is true that some Tarucus do have a rather pinkish or lilac hue 
when seen in certain lights, but it is equally clear that, in North Africa 
at least, this is not a feature which is helpful in separating the species. 
In the case of mixed rosaceus | theophrastus populations examined, 
no constant difference in ups coloration has been discerned between 
the species and it may be that the ‘rosy hue’ is due to some other 
factor. Whatever the case, assessment of perceived minor differences 
in colour must be subjective and, in North Africa, are of little value 
in species determination. 


Al 


Underside wing markings 


Recent authors, including Higgins & Riley (1983 : 57-58), Larsen (1983 : 
397) and D’Abrera (1993 : 470), have given certain differences in uns 
wing markings as features by which the three species may be separated ; 
the author has examined all these features in relation to accurately 
determined North African material and has found that, although some 
features often give good indication of a species, they are just as often 
wrong and are, in most cases, unreliable. The features most often 
presented are : 


1. The short streaks/spots which combine to form the uns pd band 
are Joined to form an irregular (jagged) line in balkanicus but remain 
as individual spots or streaks in theophrastus and rosaceus. 


COMMENT. This holds good for almost all specimens of balkanicus 
examined (in a few individuals, the pd line is ‘broken’ by the veins), 
but applies equally to a moderate proportion of rosaceus and to several 
theophrastus examined ; combined with the upf markings already 
discussed, it apparently provides the means to positively identify male 
balkanicus in a large majority of cases, but is by no means definitive. 


2. The underside forewing (unf) black submarginal and pd lines of 
theophrastus are broken by the veins which remain pale coloured from 
the cell to the wing margin, whereas in rosaceus and balkanicus there 
are normally no gaps in the lines in the apical area; the veins are 
black where they cross the line(s). 


COMMENT. This is not a reliable feature in North Africa; a small 
number of well-marked (i.e., heavily marked) theophrastus examined 
have an unbroken pd line and a high proportion of rosaceus have 
the pd line broken by the veins. The size, shape and density of all 
uns markings are very variable. 


3. The unf dark transverse ‘stripe’ common to all three species consists 
of 2 streaks of unequal length. In balkanicus and theophrastus, these 
streaks are in line ; in rosaceus the streaks are turned away from each 
other at the central point near the cell where they might otherwise 
meet. 


CoMMENT. When the author first began to examine assembled material 
it was noted that the transverse line varied in this manner and 
observations based on this feature formed the basis for the first ‘batch’ 
of genitalic examinations carried out. However, it soon became 
apparent that it did not hold true in a significant percentage of cases. 
In the case of well-marked theophrastus the two streaks, which may 


172 


be of different breadths and lengths, are often joined to form a straight 
line ; in rosaceus the lines are less likely to be joined and more likely 
to have the ‘ends turned away’ from each other, but in some cases 
the reverse is true. 


The character is very variable and butterflies displaying transitional 
features are common ; although it may be said that individuals with 
a ‘straight’ transverse line or lines are likely to be theophrastus and 
that those with the central ends strongly turned away are quite likely 
to be rosaceus, individual variation is so great that the feature is no 
more than a useful guide. In a number of specimens these markings 
are asymmetrical and indicate theophrastus on one side and rosaceus 
on the other! 


Identification of females 


The primary aim of this study was to establish the distribution of the 
three Tarucus species in the Maghreb and this was achieved without 
the necessity of addressing the question of whether it was possible to 
separate females. Some authors (e.g., Higgins & Riley), have made 
an effort to provide a guide by evaluating the extent of female ups 
blue/ white scaling ; the author believes that the considerable amount 
of individual variation makes this impossible. Underside markings will 
generally allow identification of female balkanicus, using features 
already discussed for the male, but for theophrastus and rosaceus, other 
than uns features common to both sexes which provide an (unreliable) 
indication of species, the author knows of no certain way to separate 
the females. 


Host-plant association 


Higgins & Riley (1983 : 57-58) gave Zizyphus vulgaris (Rhamnaceae) 
as the host-plant of theophrastus, and Paliurus spinachristi (Rham- 
naceae) as the host-plant of both balkanicus and rosaceus ; the unstated 
inference, since all Tarucus are notably sedentary and seldom stray 
more than a few metres from their host-plant, being that theophrastus 
and rosaceus are unlikely to be sympatric but that balkanicus and 
rosaceus may fly together. It transpires that the first of these inferences 
is far from the truth. 


It is not altogether clear on what Higgins & Riley (1983) based these 
comments, since there are considerable data available concerning 
Tarucus host-plants, particularly in the Middle East, clearly suggesting 
that the three species routinely utilise the same host-plants. Selected 


173 


published records concerning Zizyphus and Paliurus host-plant asso- 
ciations of the three Tarucus species are summarised in Tab. 4. 


In many Middle Eastern localities, as Tab. 4 makes clear, all three 
Tarucus species utilise whatever Zizyphus or Paliurus host-plants are 
available ; it is also quite clear that rosaceus and theophrastus often 
fly together and that in fact all three Tarucus species may sometimes 
be sympatric. For example, Larsen (1983: 397) reported finding 
theophrastus and rosaceus on the same Z. spinachristi trees in Jordan 
and said that all three species were found in one Saudi Arabian locality. 


Table 4 


Selected published host-plant records of T. balkanicus, T: rosaceus and T. theophrastus. 
All host plants belong to the genera Zizyphus or Paliurus 


Host plants recorded for 


Reference T. theophrastus T. balkanicus 
Graves, 1925 (Arabia) Er] Z. spinachristi 


Higgins & Riley, 1983 Z. vulgaris P. spinachristi | P. spinachristi 
(Europe / N. Africa) 

Manley & Allcard, 1970 Z. lotus 
(Spain) 


Larsen, 1974 Z. vulgaris 
(Lebanon) Z. spinachristi 
P. spinachristi 

P. vulgaris 


Larsen, 1980 (Oman) 


Larsen & Nakamura, 1983 Z. spinachristi | Z. spinachristi 
(E. Jordan) Paliurus sp. 


Larsen, 1983 Z. spinachristi Z. spinachristi | Z. spinachristi 
(Arabia/ Dhofar) Z. leucodermis Z.  leucoder- 

mis Z. leucodermis 

Z. nummularia 


The same appears to be the case in North Africa. Not initially 
appreciating the significance of misleading published data concerning 
host-plant associations, the author neglected to take a sample of host- 
plant from each locality where Tarucus was collected, but since the 
only samples taken (of Z. lotus from T19 and P spinachristi from 
T25) each supported both rosaceus and theophrastus, it seems likely 
that in most North African localities the two species are sympatric 
regardless of which host-plant is utilised. Observations at 2400m below 
Oukaimeden in the Moroccan High Atlas (T11), suggest that host- 


174 


plants other than Zizyphus or Paliurus may occasionally support T. 
theophrastus. 


Genitalia 


Although external features seldom give more than an indication of 
the identity of theophrastus and rosaceus, all three Tarucus species 
in the Maghreb are easily identifiable from examination of the male 
genital armature. It is unnecessary to look further than the valve, which 
in theophrastus tapers to an apex terminating in three deeply divided 
and very distinctive spines or teeth, quite unlike any other member 
of the genus. The harpe is short, approximately half the length of the 
valve. In rosaceus the shape of the apex of the valve appears quite 
variable, but generally tapers abruptly to a pointed apex ; the harpe 
is long and gently curved, reaching almost to the apex of the valve, 
whilst in balkanicus the shorter and more strongly curved harpe rises 
from the rounded edge of the valve and extends well beyond the valve 
apex. Since the author’s material comprises only theophrastus and 
rosaceus, a number of male balkanicus from Jordan were examined 
for the purpose of comparison. 


The valves have been illustrated by several authors (Bethune-Baker, 
1918: pl. 16, pl. 18 ; Stempffer, 1942 : 121 ; Higgins, 1975 : 116-117; 
Larsen, 1983 : 398) and of these, Stempffer and Larsen provided the 
most accurate and useful figures. Surprisingly, Higgins’ figure is of the 
least practical value and gives a misleading impression of the valve, 
particularly in the case of theophrastus. The primary differences 
between the valves of the three species are shown in Fig. 3. 


Conclusions 


The conclusions of this study may be summarised as follows : 


1. Wing markings will allow identification of both sexes of 7: balkanicus 
in the great majority of cases. Only genitalic examination will allow 
separation of male theophrastus and rosaceus with any certainty ; the 
author knows of no method of separating the females of these two 
species. 


2. T. balkanicus is apparently widespread but very local in the Maghreb. 
T. theophrastus is widespread and often common throughout the region 
up to 1400 m and may occasionally be found up to 2400 m or more ; 
it has been recorded from relatively high levels of the Moroccan High 
Atlas mountains and from the Hoggar mountains of southern Algeria. 


175 


Fig. 3. Valves of 1 — 7: balkanicus ; 2 — T. rosaceus ; 3 — T. theophrastus. 


T. rosaceus ıs also widespread and is apparently sympatric with 7. 
theophrastus in many (or most) suitable localities at low or moderate 
altitudes. 


3. All three species probably utilise whatever Zizyphus or Paliurus host- 
plant species are available in the Maghreb. 


Acknowledgements 


Thanks are due to a number of people, acknowledged here in alphabetical 
order. Mr. Phil Ackery (GB-London), of the British Museum (Natural 
History), London, allowed access to the unrivalled collections therein ; Mon- 
sieur Guy Barragué (F-Cannes), loaned material from his collection ; Madame 
Hnia Bencheikh (MA-Rabat), Chef de la Division de la Coopération, Ministére 
de l’Agriculture et de la réforme Agraire, gave authority to collect and study 
butterflies in Morocco; Dr. Ulf Eitschberger (D-Markleuthen) provided 
material from his collection and translated the Zusammenfassung ; Senor 
Filipo Fabiano (I-San Gimignano), loaned material from his collection ; Dr. 
Wolfgang ten Hagen (D-Bonn), provided material from his collection ; Dr. 
Abderrahman Jerraya (TUN-Tunis), l’Institut National Agronomique de 
Tunisie, gave authority to collect and study butterflies in Tunisia; Dr. Ole 
Karsholt (DK-Copenhagen), loaned material from the Zoologisk Museum, 
Copenhagen ; Mr. Michel Tarrier ([F] E-Myas) translated the résumé ; my 
wife Julie prepared the maps and figure which accompany this paper. 


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1197 7 


Nota lepid. 18 (2) : 178 ; 29.11.1996 ISSN 0342-7536 


Tagungsankündigungen 
OEG-Fachtagungen 1996 + 1997 


Kolloquium der Osterreichischen Entomologischen Gesellschaft 1996 : 
Wien, Institut für Zoologie, Universität für Bodenkultur, 23. Marz 1996 
(10.00 17.00 Uhr). Doz. Dr. E. Christian, Gregor Mendelstraße 33, 
A-1180 Wien ; Tel. : 0222/476543231 


Fachgespräch der Osterreichischen Entomologischen Gesellschaft 1996 : 
Illmitz, Biologische Station, 19. Oktober 1996. Thema : ”Die Erfassung 
und Erforschung der Entomofauna in Osterreich - eine Standortbes- 
timmung“. HR Doz. Dr. A. Herzig, Biologische Station, A-7142 Ilmitz, 
Tel. : 02175 / 23280. 


Kolloquium der Osterreichischen Entomologischen Gesellschaft 1997 : 
Innsbruck, Institut fiir Zoologie (Abt. Terrestr. Okologie und Taxo- 
nomie) der Universitat, 15. Marz 1997. Doz. Dr. K. Thaler, Techni- 
kerstraße 25, A-6020 Innsbruck, Tel. : 0512/748/5354. 


Fachgespräch der Osterreichischen Entomologischen Gesellschaft 1997 : 
Wien, Institut fiir Zoologie der Universitat, 19. Oktober 1997. Thema : 
”Struktur Funktion Evolution : aktuelle Trends in der morphologischen 
Erforschung der Insekten“. Doz. Dr. Günther Pass, Althanstr. 14, A- 
1090 Wien, Tel. : 0222/31336/ 1348. 


Auskünfte : J. Gepp, ÖEG-Geschäftsführer, Institut für Naturschutz, 
Heinrichstraße 5, A-8010 Graz. 


178 


Nota lepid. 18 (2) : 179-180 ; 29.11.1996 ISSN 0342-7536 


Short communication — Kurze Mitteilung — En bref 


Zur Lebensweise der Raupe von Euchalcia emichi 
(Rogenhofer, 1873) in Zentralanatolien (Türkei) 
(Lepidoptera : Noctuidae, Plusiinae) 


Hans L6seL, Wilhelm Külz Str. 11, D-99706 Sondershausen 
Thomas DRECHSEL, Hufeisenstr. 14, D-17034 Neubrandenburg 


Summary 


Larvae of Euchalcia emichi (Rogenhofer, 1873) were found on a yellow- 
flowered Nonea sp. (Boraginaceae) in Central Anatolia (Turkey). The larvae 
and their biology are compared with the closely related species E. consona 
(Fabricius, 1787) from Central Europe and E. taurica (Osthelder, 1933) from 
the Near East. 


Résumé 


Description des chenilles de Euchalcia emichi et de leur vie sur la Borraginacée 
Nonea sp. a fleurs jaunes, en se basant sur des observations de terrain en 
Anatolie centrale (Turquie). Comparaison avec Euchalcia consona d’Europe 
centrale et E. taurica d’Asie Mineure. Les traits communs relevés permettent 
de conclure à une très proche parenté entre E. emichi, E. consona et E. taurica. 


Anläßlich einer entomologischen Studien- und Sammelreise während 
der 3. Aprildekade 1995 in die Türkei fielen den Autoren in der Umge- 
bung der zentralanatolischen Stadt Goreme (Kappadocien, Provinz 
Nevsehir) reichliche Bestande einer gelbblühenden Napfkrautart (Nonea 
sp., Boraginaceae) auf. Diese Pflanze gleicht weitgehend dem heimischen 
Braunen Napfkraut Nonea pulla, unterscheidet sich aber neben der 
gelben Blütenfarbe durch die mehr laubgrüne Färbung der Blätter (bei 
N. pulla deutlicher blaugrün). Die Bestimmung dieser Pflanze muß 
einem Botaniker vorbehalten bleiben. 


Da 1992 im Pontischen Gebirge bei Ilgaz (1800 m üNN) von Gelbrecht, 
Drechsel & Löbel bereits Raupen der Euchalcia taurica (Osthelder, 
1933), die als Falter der heimischen E. consona (Fabricius, 1787) stark 
ähnelt, an blaublühendem Napfkraut gefunden wurden, sollte versucht 
werden, diese Art auch hier als Raupe nachzuweisen. Mit den 


179 


Erfahrungen bei der Suche der Raupen und deren Lebensweise von 
E. consona und E. taurica wurden von den Autoren am 23. April 
1995 in kurzer Zeit zahlreiche Euchalcia-Raupen auch an dieser Pflanze 
gefunden. Der Fundort lag am östlichen Rand des Dorfes Cavusin. 


Bei einer Parasitierung von etwa 40% gelang die Zucht im Beutel auf 
der eingefrischten Futterpflanze weitgehend verluBtlos. Die Mehrzahl 
die Tiere war bis zum Riickflug am 30. April bereits verpuppt. Zwei 
kleinere, leider auch parasitierte Raupen, konnten in Deutschland 
zunächst mit Nonea pulla weitergefüttert werden. 


Die Falter schlüpften zwischen dem 10. und 18. Mai 1995. Zu unserer 
Uberraschung handelte es sich aber nicht um Euchalcia taurica, sondern 
um E. emichi (Rogenhofer, 1873). Diese Art wurde durch die Autoren 
bereits 1991 und 1992 bei Göreme und an weiteren Fundorten in der 
Türkei als Falter am Licht nachgewiesen. 


Die Lebensweise der Raupen von Euchalcia consona, taurica und 
emichi sind nahezu identisch. Wie bei E. consona und E. taurica, lebt 
die junge Raupe der E. emichi in zusammengesponnenen Spitzentrieben 
und Blütenständen des Napfkrautes. Auch in Habitus und Aussehen 
gleicht sie den anderen genannten Arten: sie ist ebenso wie diese von 
der Grundfarbe der Futterpflanze (licht blaugrün) mit zahlreichen 
dunklen Punktwarzen. 


Die erwachsene Raupe ın letzten Stadium lebt zwischen locker gehef- 
teten Blättern oder frei an der Futterpflanze, meist aber verborgen an 
unteren Blättern. Sie hat die auffälligen Punktwarzen verloren, ist etwas 
heller laubgrün und trägt eine schwache, helle Seitenlinie. In der helleren 
Grundfarbe unterscheidet sie sich von E. consona geringfügig. Die 
Verpuppung erfolgt wie bei E. consona und taurica in einem weichen 
Gespinst zwischen bodennahen Blättern der Futterpflanze. 


Das Aussehen und die Biologie der Präimaginalstadien von E. emichi 
unterstreichen deren Zugehörigkeit zur Gattung Euchalcia und ihre 
enge Verwandtschaft zu E. consona und E. taurica, auch wenn diese 
beiden Arten nach der Grundfarbe und Zeichnungsanlage stärker E. 
variabilis Piller, 1783 und E. modesta Hübner, 1786 als der kleineren, 
schokoladenbraunen E. emichi ähneln. 


180 


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Nota lepidopterologica 


Vol. 18 No. 3/4 Basel, 13.V.1996 ISSN 0342-7536 
Editor : Steven E. Whitebread, Maispracherstrasse 51, P.O. Box 27, CH-4312 


Magden, Switzerland. FAX : + 41-61-841.22.38. 


Assistant Editors : Emmanuel de Bros (Binningen, CH) 
Dr. Roger Dennis (Wilmslow, GB) 
PD Dr. Andreas Erhardt (Binningen, CH) 
Dr. Yves Gonseth (Neuchâtel, CH) 
Dr. Alexander Pelzer (Wennigsen, D) 


Contents — Inhalt — Sommaire 


ER ee aa ma errant ne ne mi EHER NEE à 10 vou 
BENYAMINI, D. : Pupal summer diapause in Chilean Pieris brassicae (Lin- 
LAEUS, SO) (AE ee ee TT NET 
FAUCHEUX, M. J. : Sensilla on the ovipositor of the carpet moth, Tricho- 
Diresarntaperzelale (imeidae) ere ee ie nn 
FiBIGER, M., HACKER, H. & MoBERG, A. : Notes on the Orthosia rorida 
(Frivaldsky, 1835) species group, with the description of a new species 
from Crete : Orthosia sellingi sp. n. (Noctuidae, Hadeninae) ............. 
Gaskin, D. E. : Hesperioidea and Papilionoidea of the Ionian island of 
Kefalonia, Greece : Additional species and a review of faunal compo- 
MENS RP PE Oe anced Mane RENNER 
HATTENSCHWILER, P.: Dahlica wehrlii (Müller-Rutz, 1920) wieder ge- 
funden. Beschreibung des Weibchens und Ergänzungen zur Kenntnis 
des Männchens und der Ökologie (Psychidae) ................................ 
HÄTTENSCHWILER, P.: Eine neue Montanima — Art aus dem Altai- 
Gebiet(Bsychidae) ..................... ne nn ne moe de 
HATTENSCHWILER, P. : Eine neue, im Februar-März fliegende Ptiloce- 
phala — Art aus dem südlichen Spanien (Psychidae) .......................- 
HAUSER, E. : Vergleichende Analyse der Zönosen tagaktiver Schmetter- 
Isehim Senssengebirge (Oberösterreich) ................22......... 0... 
MoLinA, J. Ma. & PALMA, J. M. : Butterfly diversity and rarity within 
selected habitats of western Andalusia (Spain) (Papilionoidea and 
alle SCI OI GLE DMR Re sodbves sede vandctelivadenvon 
Owen, D. F.: Larval food shortage and adult dispersal in Callimorpha 
AOA GM (Mee) CN CUM AC) MR Rene tec dececcsenbuosdeeiecsbedavarseacsess 
SCHOLZ, À. : Zur Identität von Epermenia falciformis (Haworth, 1828) 
(pemienidae) Mere hes RES ARR RER 


184 


193 


203 


213 


TAMMARU, T., VIRTANEN, T., RUOHOMAKI, K. & KAILA, L. : Eupithecia 
veratraria Herrich-Schäffer, 1850 new to Scandinavia, with the re- 
description of E. veratraria ssp. arctica Viidalepp, 1974 (Geometridae) 297 


Short communications — Kurze Mitteilungen — En bref 


LOELIGER, E. A. : Rubiginous larvae of Hyles hippophaes (Esper, 1793) 


an autosomal recessive variety (Sphingidae) .....................cccesceeseeeee 304 
LOELIGER, E. A. : Additional experiments to unravel the enigma of Hyles 

hybrid paul Mory (Sphingidac) nn... 306 
Book reviews — Buchbesprechungen — Analyses .................. 202, 308, 309 
Symposium über Entomofaunistik ....................................................... 224 
INSECTA ’% 5,050 SR M A SN EE RES 266 
Vol. 18 — 1995 
Dates of publication — Publikationsdaten — Dates de publication 310 
Contents — Inhalt — Sommaire 2... en 310 
New taxa described in Vol. 18 — Neue Taxa in Band 18 beschrieben — 

Nouveaux taxa décrits dans le VOL RE ycsen-s es eee 312 


182 


Nota lepid. 18 (3/4) : 183 ; 13.V.1996 ISSN 0342-7536 


Editorial 


As many of you will have read (SEL-News 26), this is my last issue 
of Nota as Editor. Over the last year or two, I have found it increasingly 
difficult to find sufficient time for my editorial duties and this has been 
reflected in the late appearance of the last few issues, but not, I hope, in 
their quality. I therefore informed Council of my decision to resign, as 
soon as a new Editor could be found. Alain Olivier, Luitenant Lippens- 
laan 43 b 14, B-2140 Antwerpen, Belgium, agreed to take over the 
editing of volume 19 so that I could concentrate on finishing volume 18. 
I will remain responsible for the Congress Proceedings Supplements 
so that these can be published more rapidly than in the past. 


I have been involved with Nota since 1980, when Emmanuel de Bros 
took over as Editor and invited me to join his Editorial Committee. 
In 1986, I was elected Ordinary Council Member of SEL and this was 
the same year that ‘Nota’ introduced the manuscript review process. 
In 1990, at the 7th SEL Congress (Lunz-am-See, Austria), I was elected 
as Editor. 


I set myself three main aims : To improve punctuality of publication, 
to improve the quality (content and style) and to publish the Congress 
Proceedings as supplements. Unfortunately, I found that the last two 
goals conflicted with the first. To achieve quality took time and time 
became scarcer. I was never able to reach the first goal, but judging 
from the letters I have received, I feel that I have been successful with 
the second. Three Proceedings issues have been issued and the next 
is well on its way. 


I would not have been able to produce ‘Nota’ without the help of many 
people. I would sincerely like to thank my Co-Editors : Emmanuel de 
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Pelzer and Roger Dennis, the numerous reviewers, and last but not least 
the authors, for their immense support, cooperation and patience. 


Steven WHITEBREAD 


183 


Nota lepid. 18 (3/4) : 184-192 ; 13.V.1996 ISSN 0342-7536 


Pupal summer diapause in Chilean Pieris brassicae 
(Linnaeus, 1758) (Lepidoptera, Pieridae) 


Dubi BENYAMINI 


91 Levona str, Bet-Arye, D.N. Modiin 71947, Israel 


Summary 


The European butterfly pest Pieris brassicae (L.) has been known from Chile 
since the early 1970’s, when it was probably introduced from Europe. Evidence 
is presented to suggest that it has developed a pupal summer diapause, 
previously not known in this species, to survive the dry mediterranean summer 
of central Chile. 


Résumé 


L’espèces européenne Pieris brassicae (L.), dont la chenille ravage parfois les 
plantations de choux, est connue depuis le début des années 1970 au Chili, 
ou elle a probablement été introduite d’Europe. Les constatations faites sug- 
gèrent qu'il s’est établi une diapause estivale au stade chrysalide — inconnue 
jusqu’à présent chez cette espèce — pour lui permettre de survivre à l'été de 
type méditerranéen qui règne dans la région centrale du Chili. 


Introduction 


The biology of the Palaearctic Pieris brassicae (Linnaeus, 1758) is well 
known and documented in hundreds of books and publications. This 
pest and migratory species is widespread in the whole of Europe, the 
western part of North Africa, and Asia north to about 62° of latitude. 


In Central Europe and Asia, the adults are on the wing during the 
summer from April to October in successive broods. During the winter 
it hibernates in the pupal stage (Chinery, 1989 ; Pullin & Bale, 1989 ; 
Pullin et al., 1991). In southern, warmer climates the winter pupal 
diapause is short or non-existant and during the summer the lack of 
wild crucifers forces it to switch to cultivated areas, or to other plants, 
e.g. Capparis spinosa L. (Capparaceae) in Israel (Benyamini, 1990). 


In the early 1970’s this butterfly suddenly appeared in Chile (Gardiner, 
1974). At that time, it is known that the Allende regime imported 


184 


potatoes by boat from Poland to feed the starving population. 
Specimens of P. brassicae, probably as pupae, were possibly accidentally 
imported with these ships from Poland, although other sources cannot 
be excluded. 


If the species was introduced to Chile in the pupal stage it would have 
had a relatively good chance to survive, especially during the months 
Sept.-Dec. and April-May : 


a. If European winter diapausing pupae arrived in the Chilean summer 
the switching would be natural except during the months of Feb.-March 
when the central valley of Chile is dry, not offering any green wild 
foodplant to the larvae. The only available food source would be 
cultivated crucifers. 


b. European spring pupae/adults arriving to the Chilean autumn can 
find fresh newly grown crucifers, larvae can feed up quickly and pupae 
may go immediately into winter diapause as they do in central and 
northern Europe. 


c. European summer pupae/adults arriving to the Chilean winter will 
either remain in diapause or die of cold temperatures. 


d. European autumn pupae/adults arriving to the Chilean spring will 
have the best season for acclimatisation. 


Unfortunately, no work was done to follow year by year this fascinating 
“field-experiment” of an introduced insect whose annual seasonality 
was switched suddenly by 180°, from summer to winter, from spring 
to autumn or vice-versa. Angulo & Weigert (1982) stated that “a set 
of 30 pupae hatched after 90 days, probably after staying in diapause 
or anabiotic stage”. No dates or season were given. Angulo however 
remembers that the rearing was done during the Chilean autumn and 
that the said diapause was actually a normal “European” winter 
diapause (pers. comm.). 


From late 1992 to mid 1995 the author had the opportunity to check 
the present-day situation and found that the species had fully adapted 
to the Chilean climate. 


Observations 


On 12th Dec. 1992, a female was observed flying around a yellow 
crucifer at Rio Mapocho (1100m) on the eastern outskirts of Santiago 
de Chile. The 95cm tall Brassica campestris L. that was located half 
in the shade was a perfect choice for the female to settle on and lay 


185 


its eggs. Three batches of 14, 40 and 59 eggs were layed on the underside 
of the leaves, 35 to 40 cm above the ground. The eggs were brought 
to Santiago and under a 26/17°C 14L/10D regime started to hatch 
on 15th Dec. 1992. The newly hatched larvae were partly transparent 
green turning within a few hours to light-green, similar to the colour 
of the leaves ; the head was black. 


The larvae were fed on fresh wild crucifer leaves and grew up quickly. 
Pupation started on 30th Dec. and ended on 6th Jan. 1993. A total 
of 52 pupae were obtained. The pupae were kept outside, but protected 
from rain, in Santiago. The first adult, a male, emerged on 9th Jan., 
the second, a female, on 11th Jan. Up to the 18th Jan., 36 butterflies 
had emerged. No more specimens emerged until one on 2nd Feb. The 
emergences proceeded with 2, 3 and 6 adults for March, April and 
May respectively, although 2 of the May specimens died during 
emergence. The last adult emerged on 8th May, four full months after 
pupation. Four pupae turned dark-brown and died. 


The monthly and cumulative emergences are illustrated in Fig. 1. 
Within 10 days of pupation, 68.6% of the butterflies had emerged. 
A further 23.4% emerged gradually during the summer. From the end 
of March when the temperatures started to drop, heralding the first 
autumn rains, the emergences intensified, reaching a peak in the first 
week of May when 5 of the 6 remaining live pupae produced adults 
simultaneously on the 2nd and the 3rd. Eight percent of the pupae 
died. 


In the Chilean summer of 1994-95 field observations yielded the same 
results as those of 1992-93 from the laboratory ; from mid-January 
to mid-February 1995 no adults were observed. On February 17th the 
first adult was recorded from Coronel, a coastal locality near Concepcion 
which is about 500 km south of Santiago. In Santiago the first ap- 
pearance was about two weeks later, starting March 6th and then 8th 
& 10th in El Bosque (southern Santiago, one specimen each time) and 
on the 11th in Providencia Gardens in the centre of the city (three 
specimens). The adults were active until April 17th (El Bosque), when 
a two week period of cold and rainy weather set in. May Ist was 
sunny and warm (26° at midday) and three specimens were observed 
(Collina, Santiago region). At El Bosque the following day, 11 
specimens were observed between 11:30 a.m. and 2 :00 p.m., and on 
May 3rd, 13 specimens were observed in ten minutes at 1:30 p.m. 
May 4th was partially cloudy and only three specimens were observed 
between 1 :00-2 :00 p.m., May 5th was rainy and no butterflies were 
seen. The emergences of the wild summer pupae as observed during 


186 


Monthly and Cumulative 
emergences in % 


100 % 
8 % Died 


diapause 


75 % 


50 % 


68,6 % 


7 - 10 days 
Normal 
emergence 


25 % 


ue 8,1 
January February March April May 


Fig. 1. Monthly and cumulative pupal emergences in one brood of Chilean P brassicae. 


the autumn of 1994-95 season perfectly match the laboratory results 
as shown in Fig. 1 : No observations during mid-summer and a slow 
build up of emergences during February, March until the peak in early 
May. 


In order to have a comparison with a native Pierid, rearing material 
of the legume feeder Tatochila theodice Boisduval were obtained. On 
the 15th and 19th Nov. 1994 in Villa Paulina (Valle de la Yerba Loca) 
about 20 km east of Santiago 2000 m.a.s.l. eggs, Ist and 3rd instar 


187 


larvae were found on Vicia sp. (Fabiaceae). Of the 15 eggs and larvae 
that were collected and bred in Santiago 9 pupae were obtained between 
27.11 and 8.12.1994 (3 eggs were parasitised and 3 larvae died of 
starvation). Three pupae turned dark brown or black and died during 
Jan. 1995, probably because the larvae were tested unsuccessfully for 
feeding on Trifolium repens L. Of the 6 remaining pupae 1 & emerged 
on 17.12.1994. No more emergences took place during the next two 
and a half months, but during the first fortnight of March 3 pupae 
hatched. The last two pupae went into winter diapause. Table 2 
summarizes this observation which, although based on very little 
material, suggests the same phenomenon of two seasonal pupal 
diapauses. Over summer for two and a half months and in winter 
for eight months. 


Table 1 


Emergences of Tatochila theodice Boisduval from the western Andes Precordillera 
slopes of the Santiago Metropolitan Region, Chile 1994-1995. 


Month Dec. Jan, Feb. Mar. 
1994 1995 1995 


Behaviour Ist brood LATE? Summer LG 4 Winter 
(wild) diapause 19:73 diapause 


6 53 
2nd brood 3rd brood 
(lab.) (lab.) 


Pupal 17% 50% 33% 
emergences 


Discussion 


The European annual life-cycle of P brassicae is composed of spring 
and summer continuous generations and winter diapausing pupae 
(Table 2). In southern Europe, North Africa and the Middle East the 
behaviour is slightly different (Benyamini, 1990) ; the local temperatures 
are not low enough to initiate diapause. Instead, metamorphosis is 
simply slowed down, larvae taking longer to feed up and pupae taking 
longer to hatch. In the dry summer of Israel, when no fresh wild 
crucifers exist, the adults switch to laying eggs on C. spinosa and on 
cultivated crucifers, and pupae have no summer diapause. 


In Chile we see a modified survival technique. With the lack of potential 
alternative wild summer foodplants, the pupae go into a second annual 
diapause. It has previously been shown that a short day-length, vitamin 
A and cryoprotectant accumulation are the essential factors for winter 


188 


Table 2 


Seasonal metamorphosis of P brassicae in three different regions of the world. 


Winter Spring Summer Autumn 

Nov. - April April - July July - Sept. Sept. - Nov. 
C. & N. Europe Diapausing pupae Continuous breeding Pupae 

Nov. - Feb. Feb. - May May - Nov. 
Middle East (Israel), Slower Breeding Breeding, switching to other foodplants 
southern Europe & metamorphosis 
North Africa 

June - Sept. Sept.- Jan.  Jan.- April April - June 
Chile Diapausing pupae Breeding Diapausing pupae & Breeding 


gradual emergences 


diapause (Claret et al., 1981 ; Claret, 1985 ; 1989; Claret & Volkoff, 
1992 ; Lavialle & Dumortier, 1990 ; Spieth & Sauer 1991 ; Veerman 
et al. 1985 ; 1988). Of these, only vitamin A could possibly be playing a 
role in the summer diapause. It would also suggest that while a short 
day-length induces diapause, a long day-length does not prevent it. 


During March 1995 a similar emergence pattern was seen in other 
butterfly species in the wild. On 18.3.95 at Bafio Morales, 2200m, 
Santiago Metropolitan region 6 adults of T. theodice were observed. 
In El Bosque daily more species and individuals of Colias vauthieri 
Guerin, P. brassicae L., Tatochila autodice blanchardi Butler, (Pieridae), 
Vanessa carye Hübner (Nymphalidae), Erynnis funeralis Scudder & 
Burgess and Hylephila faciolata Blanchard (Hesperiidae) were observed. 
This phenomenon of a second peak of butterfly activity (shown 
schematically in Fig. 2) is typical of Mediterranean zones where early 
autumn rains produce a second spring-like period with many plants 
having a second flowering period, side by side with the young growth 
of annuals. The emergence of P brassicae adults after the summer 
diapause perfectly coincides with this second peak of seasonal butterfly 
activity. 


Dr. Torben Larsen, who studied the butterflies of the Lebanon for 
4 years (Larsen, 1974) also found larvae switching to Capparis, but 
believed that the few adults observed by him during the summer months 
in the mountains “may be migrating specimens.. but most of the 
summer brood probably aestivates in pupal stage”. This “aestivation” 
was not proven by Larsen. If he had bred the Capparis larvae, he 
would have found that not a single pupa diapauses, as in neighbouring 
Israel (pers. obs.). Larsen & Nakamura (1983: 159) considered the 
same theory in their work on the butterflies of East Jordan: “There 


189 


N° of 
Species 
(Active 
Adults) 


Month 


TORBEN 10 EURE LS 
Fig. 2. Schematic diagram of the flight activity of butterflies in the Chilean (Mediter- 
ranean) central valley. 


is a tendency towards both hibernation and aestivation in the pupal 
stage”. In his excellent review on P brassicae, Feltwell (1981 : 222- 
225) did not find any definite evidence of a summer pupal diapause, 
but quoted Larsen (1974). The closest observation to my own is that 
of Bowden (1966) who noted that Corsican larvae that were brought 
back to England and pupated there “hatched out much later than their 
own kin, but in a shorter time than those which diapause”. Seventeen 
males and females hatched normally, but four pupae hatched three 
weeks to three months later, being 19% of the total emergences. He 
termed this “irregular diapause”. The fact that emergences took place 
in a different country under different conditions could however have 
influenced the results. Danilevskii (1965) who bred P. brassicae from 
Sukhumi (Georgia) found that when he kept the pupae at a constant 
temperature of 23°C the emergences took place within two and a half 
to three months. Bowden (1966 : 68) wrote : “Thus the more southern 
the Russian P brassicae seems to enter a shallow diapause which does 
not require cold to terminate it”. 


It is important to note that in other genera of the Pieridae inhabiting 
Mediterranean climates i.e. Euchloe, a summer pupal diapause is very 
common. 


190 


Dr. Angulo states that P brassicae has become less common since its 
sudden appearance in Chile (pers. comm.). It seems that a new ecological 
balance has been achieved. Its well-known Palaearctic parasite Apanteles 
glomeratus (Hymenoptera, Braconidae) also appeared in Chile and I 
found it in a few of the 1994-95 wild collected larvae (e.g. 28.11.94 
Rio Clario 800m, Santiago Metropolitan Region). 

Feltwell (1981 : 223) suggested “to investigate the seasonality of P 
brassicae in Chile, as it has a similar climate to that of the Lebanon”. 
I suggest checking the biology of P brassicae in Corsica and in southern 
central Asia, as both these regions have a similar climate to the Chilean 
central valley. 


Acknowledgements 


I want to thank Dr. Andres O. Angulo of Universidad de Concepcion, Chile 
who shared with me his past experience with P brassicae. Prof. Arthur M. 
Shapiro of University of California, Davis, and Dr. David G. Furth of the 
Smithsonian Institution, Washington D.C. read the manuscript and gave some 
helpful comments. Mrs Jaqueline Ruffle, Librarian of the Royal Entomological 
Society helped with the references, Senor Alejandro Barrales drew the figures, 
Avishay Benyamini typed the manuscript. Last but not least I am grateful 
to Mr. Steven Whitebread who edited the manuscript to its final form. 


References 


AnGu1o, A. ©. & WEIGERT, G. Th., 1982. Biologia y postembriologia de 
tres Lepidopteros en Chile. Brenesia 19/20 : 431-449. 

BENYAMINI, D., 1990. A field guide to the butterflies of Israel. 234pp. Keter 
Publishing House, Jerusalem. 

BowDen, S. R., 1966. “Irregular” diapause in Pieris, with a note on Corsican 
P. brassicae L. (Lep. Pieridae). Proc. S. Lond. ent. nat. Hist. Soc. 1966: 
67-68. 

CLARET, J., 1985. Two mechanisms in the biological clock of P brassicae 
L.: An oscillator of diapause induction ; an hour-glass for diapause 
termination. Experientia 41(12) : 1613-1615. 

CLARET, J., 1989. Vitamin A and photoperiodic or thermoperiodic induction 
of diapause in P brassicae. C.R. Hebd. Seances Acad. Sci. III, Paris 
308(13) : 347-352. 

CLARET, J., DUMORTIER, B. & BRUNNARIOUS, J., 1981. Evidence for a 
circadian component in the photoperiodic clock of P. brassicae (Lepi- 
doptera). C.R. Seances Acad. Sci. III, Paris 292(5) : 427-430. 

Cuinery, M., 1989. Butterflies and day-flying moths of Britain and Europe. 
320 pp. Collins new generation guide. 

DaniLevsku, A. S., 1961 (transl.1965). Photoperiodism and seasonal develop- 
ment of Insects. Edinburgh. 


191 


FELTWELL, J., 1981. Large white butterfly. the biology, biochemistry and 
physiology of Pieris brassicae (Linnaeus). 535 pp. Dr. W. Junk, The 
Hague. 

GARDINER, B. O. C., 1974. Pieris brassicae L. Established in Chile ; another 
Palaearctic pest crosses the Atlantic (Pieridae). J. Lepid. Soc. 28(3) : 
269-277. 

LARSEN, T. B., 1974. Butterflies of Lebanon. 256 pp. National Council for 
Scientific Research, Beirut. 

LARSEN, T. B. & NAKAMURA, I., 1983. The butterflies of East Jordan. 
Entomologist’s Gaz. 34 : 135-208. 

LAVIALLE, M. & Dumortier, B., 1990. Metabolic correlates in the working 
of an insect putative photoperiodic clock. J. Comp. Physiol. (A) 166(3) : 
785-789. 

Purum, A. S. & BALE, J. S., 1989. Influence of diapause and temperature on 
cryoprotectant synthesis and cold hardiness in pupae of Pieris brassicae. 
Comp. Biochem. Physiol. 94A(3) : 499-503. 

Putin, A. S., BALE, J. S. & FINTAINE, L. R., 1991. Physiological aspects of 
diapause and cold tolerance during overwintering in Pieris brassicae. 
Physiol. Ent. 16 : 447-456. 

SHAPIRO, A. M., 1975. The temporal component of butterfly species diversity. 
pp. 181-195 in: Cody, M. L. & Diamond, J. M. (Eds). Ecology and 
evolution of communities. 545 pp. Harvard Press. 

SPIETH, H. R. & SAUER, K. P., 1991. Quantitative measurement of photo- 
periods and its significance of the induction of diapause in P brassicae 
(Lep. Pieridae). J. Insect Physiol. 37(3) : 231-238. 

VEERMAN, A., SLAGT, M. E., ALDERLIESTE, M. G. J. & VEENENDAAL, R. L., 
1985. Photoperiodic induction of diapause in an insect is vitamin A 
dependent. Experientia 41(9) : 1194-1195. 

VEERMAN, A., BEEKMAN, M. & VEENENDAAL, R. L., 1988. Photoperiodic 
induction of diapause in the large white butterfly, P brassicae : Evidence 
for hourglass time measurement. J. Insect Physiol. 34(11) : 1063-1069. 


192 


Nota lepid. 18 (3/4) : 193-201 ; 13.V.1996 ISSN 0342-7536 


Sensilla on the ovipositor of the carpet moth, 
Trichophaga tapetzella L. (Lepidoptera : Tineidae) 


Michel J. FAUCHEUX 


Laboratoire d’Endocrinologie des Insectes Sociaux, Université de Nantes, 2 rue de la Houssinière, 
F-44072 Nantes Cédex, France 


Summary 


Sensilla on the ovipositor of the carpet moth, Trichophaga tapetzella L. 
(Tineidae) have been examined by scanning electron microscopy. Three rings 
of different sensilla encircle the lobes of ovipositor (3 types of aporous sensilla 
trichodea, aporous sensilla basiconica and dome-shaped sensilla. The distri- 
bution of sensilla is specific to 7: tapetzella and has been observed neither 
in other keratophagous moths, or other phytophagous moths and butterflies. 
Dome-shaped sensilla are described for the first time on the ovipositor of 
Lepidoptera. All the sensilla are probably capable of monitoring both cuticular 
stress and mechanical contact of the ovipositor from the initial through the 
final steps of penetration in the substrate, egg-laying and copulation. 


Résumé 


Les sensilles sur l’ovipositeur de la mite tapissière, Trichophaga tapetzella L. 
(Tineidae) sont examinées au microscope électronique a balayage. Trois 
anneaux de sensilles différentes encerclent les lobes de l’ovipositeur (3 types 
de sensilles trichoides sans pore, des sensilles basiconiques sans pore et des 
sensilles en forme de dôme). La distribution des sensilles est spécifique de 
T. tapetzella et n’a été observée, ni chez les autres papillons kératophages, 
ni chez les phytophages. Les sensilles en forme de dôme sont décrites pour 
la première fois sur l’ovipositeur des lépidoptères. Toutes les sensilles sont 
probablement capables d’informer l’insecte sur les déformations cuticulaires 
et les contacts de l’ovipositeur depuis le stade initial jusqu’au stade final de 
la pénétration dans le substrat, ainsi que lors du dépôt des oeufs et de la 
copulation. 


Introduction 
The carpet moth, Trichophaga tapetzella L. (Tineidae), is a widespread 


and destructive pest of fabrics, wool, hair, and all other keratinaceous 
materials (Zagulajev, 1960), which inhabits temperate regions. Native 


193 


to the Old World, it has been introduced through trade into the New 
World and the Australasian Region. In natural conditions, the larvae 
are often found in association with bird nests. As the choice of a suitable 
substrate for oviposition is of critical importance for females, we studied 
the ovipositor sensilla which are involved in this behaviour. 


In most lepidopterans, the ovipositor bear tactile mechanoreceptive and 
contact chemoreceptive sensilla (Chadha & Roome, 1980 ; Valencia & 
Rice, 1982 ; Anderson & Hallberg, 1990 ; Faucheux, 1991). Multiporous 
sensilla with a putative olfactive function have been discovered in two 
keratinophagous Tineidae (Faucheux, 1982, 1988) and a phytophagous 
moth (Reymonet & Faucheux, 1991). 


The purpose of this study was to described the types and locations 
of sensilla on the ovipositor of the carpet moth, utilising the scanning 
electron microscopy (SEM) and to compare the results with other 
studies in the literature. 


Materials and methods 


Adults of TZ: tapetzella were obtained from larvae found in the 
regurgitated pellets of the barn owl, Tyto alba L., at Limerzel and 
Rochefort-en-terre, in Morbihan (West of France). For SEM study, 
18 ovipositors were fixed with 2.5% glutaraldehyde in sodium cacodylate 
buffer, post-fixed with 1% OsO, and dehydrated in an ethanol series. 
They were either dried in air or with a critical-point dryer. Specimens 
were coated with gold-palladium and observed with a JSM 6400 F 
SEM at 7 Kv. The porous areas of sensilla were determined by crystal 
violet (Slifer, 1960) and reduced silver techniques (Schafer & Sanchez, 
1976). 


Results 


During the acts of oviposition and copulation, the ovipositor is 
protracted and appears as a relatively solid, sclerotized structure, ca. 
5.3 mm long, constituted of abdominal segments 8 and 9 (Fig. 1). In 
cross section, the circumference of the proximal portion is round, and 
the distal portion is dorsoventrally flattened. When not in use, the 
Ovipositor is kept in a retracted position within the 7th abdominal 
segment and is externally not visible, except for the tip of the lobes 
(Fig. 2). There are three more or less sclerotized sensory regions : the 
distal two lobes (L) or anal papillae, the dorsal plate (DP) and the 
genital plate (GP), both situated on the 8th segment (Fig. 1). 


194 


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ipositor 
8, 9 — abdom 


Morphology of sensilla on ovipositor lobes 


The lobes of the ovipositor bear 5 types of sensilla : 3 types of sensilla 
trichodea, sensilla basiconica and dome-shaped sensilla. The three types 
of sensilla trichodea (A, B, C) can be distinguished according to their 
lengths and location. The proximal sensilla trichodea A are the longest 
hairs, 114.3 mm long (range : 82-165 mm) and possess a pointed tip 
(TA, Figs. 1,2). They have a basal diameter of 2.8-3.3 mm and are 
articulated with the ovipositor cuticle (Fig. 3). The socket tightly sur- 
rounds the base of the hair shaft which is smooth only in the basal 
portion (Fig. 3), but has 12 longitudinal ridges with a tiled-roof structure 
(Fig. 4) in the medial and distal regions. Sensilla trichodea A are thick- 
walled with a narrow lumen (Fig. 5). No terminal pore or wall pores 
are seen and the hairs did not stain with crystal violet or reduced silver 
nitrate. These sensilla therefore may be classified as aporous sensilla 
according to Zacharuk (1985). 


The more distal sensilla trichodea B are shorter than the type A and 
measure from 20 to 38 mm long (mean: 29.3 mm) and 1.2 mm wide 
at the base (TB, Figs 6, 11). They have a bulbous base and are sharply 
tipped. Sensilla trichodea C are short sensilla, 7.6 mm long (range : 
6.6 - 10.1 mm) and blunt ; the hairs are slightly inclined distally, but 
at varying angles (TC, Fig. 6). Like the sensilla trichodea B, the type 
C is inserted in a large pit (Fig. 7). The hair-shafts of both sensilla 
trichodea B and C are smooth, without any pore, and did not stain 
with crystal violet or reduced silver nitrate. 


Sensilla basiconica are located at the apex of lobes (B, Fig. 11). The 
peg is 8.5 mm long (ranging from 5.3 to 9.8 mm) and about 1.0 mm 
wide at the base (Fig. 8) and is blunt tipped (Fig. 9). These sensilla 
are always inclined distally, no pore is visible with SEM and they did 
not stain when crystal violet is applied. 


Dome-shaped sensilla, located distally (C, Fig. 11) possess a central 
pit (Fig. 10 and inset) and measure from 1.9 to 2.7 mm in diameter 
(mean = 2.2 mm). 


Distribution of sensilla on ovipositor lobes 


The distribution of sensilla on the lobes shows unusual features. We 
can distinguish the following from the distal region proximally : 


— an apical circle of 8 sensilla (4 per lobe) comprising from distal side 
towards the ventral side of each lobe : one sensillum basiconicum, one 


196 


Figs 6-11. 6 — Sensilla trichodea types B and C, scale bar = 2 mm ; 7 — Base of sen- 
sillum trichodeum type C, scale bar = Imm ; 8 — Base of sensillum basiconicum, scale 
bar = 2 mm ; 9 — Apex of sensillum basiconicum, scale bar = 0.5 mm ; 10 — Dome- 
shaped sensillum, scale bar = 1 mm; inset. another view of dome-shaped sensillum, 
scale bar= 1mm; 11 — The two distal circles of sensilla on the lobes, scale 
bar = 10 mm. 

B — sensillum basiconicum ; C — dome-shaped sensillum ; TB, TC — sensilla tricho- 
dea types B and C. 


dome-shaped sensillum and two sensilla basiconica. The dome-shaped 
sensillum is slightly below the circle of sensilla basiconica (Fig. 11). 


— a subapical circle of 18 sensilla (9 per lobe) regularly spaced and 
comprising 3 groups of sensilla. Each group possesses one central 
sensillum trichodeum B and 2 lateral sensilla trichodea C (Fig. 11). 


— a proximal circle consisting of 4 sensilla trichodea A (2 per lobe) 
(Figs. 1 and 2). | 


— 20-24 sensilla trichodea C (about 10 per lobe) scattered on the 
proximal part of the lobes. The numbers of sensilla on the three circles 
are identical in the 18 ovipositors examined with SEM. 


Sensilla on the dorsal and genital plates 


The dorsal plate and the genital plate bear only sensilla trichodea type 
A, similar to those of ovipositor lobes, but longer. On the genital plate, 
there are two (n = 5 ovipositors examined), four (n = 11) or 5 (n= 2) 
sensilla 162 mm long (range : 127-133 mm) which are inclined distally, 
at varying angles (Fig. 11). The dorsal plate bears two sensilla (n = 8) 
or only one sensillum (n = 4), 185 mm long (range : 158-210 mm). 


Discussion 


Prior to and during oviposition, the ovipositor is extended by hydro- 
static pressure and intrinsic muscles allow it to make orientation move- 
ments including probing the substrate. The sensilla described here are 
suited to play a role in this oviposition behaviour by sensing the im- 
mediate environment. 


Presumed functions of sensilla 


The sensilla trichodea types A, B, and C of T: tapetzella are morpho- 
logically similar to the 3 types of mechanoreceptors described by 
Valencia & Rice (1982), Faucheux (1988) and Reymonet & Faucheux 
(1991). Possibly, there are differences between the bristle types in the 
force needed for deflection. The flexibility of the cuticle around the 
sockets of sensilla trichodea B and C is probably an adaptation to 
prevent damage to these sensilla when the ovipositor penetrates the 
substrate. It is probable that each group consisting of one sensillum 
trichodeum B and two sensilla trichodea C may function as a unit. 
The aporous sensilla basiconica with inflexible sockets found on the 
antenna of Locusta migratoria (L.) are known to be thermo- and 


198 


hygroreceptive (Altner er al., 1981), but the information gathered for 
the sensilla basiconica in our study is not enough to ascribe a particular 
function. 


Comparison with other Lepidoptera 


The carpet moth does not possess gustative uniporous sensilla or 
olfactive multiporous sensilla observed in other lepidopterans. It also 
differs by the presence of aporous sensilla basiconica and dome-shaped 
sensilla. Dome-shaped sensilla on the ovipositors of Lepidoptera are 
described here for the first time. The reduced numbers and distribution 
of sensilla are one of the more striking features of the ovipositor of 
T. tapetzella. In this species, there is a total of about 60 sensilla on 
the whole ovipositor, compared to 110 in Tinea pellionella L. (Faucheux 
& Chauvin, 1981), 130 in Tineola bisselliella Humm. and 87 in Monopis 
crocicapitella Clem. (Faucheux, 1987) among the keratophagous moths. 


The behavioural role of sensilla 


As in other Tineidae of the subfamily of Tineinae (Faucheux, 1987), 
under laboratory conditions, the carpet moth prefers to oviposit in 
cracks or crevices (unpublished observations) and, under natural 
conditions, it oviposits between the woollen fibres of carpets or in 
regurgitated pellets. Thus, the tactile hairs may determine extent of 
penetration of the ovipositor. The sensilla trichodea A of the genital 
and dorsal plates may contact the substrate when the 9th and 10th 
segments are retracted into the 8th abdominal segment. To explain 
how the female determines the depth of the penetration hole, we suggest 
a mechanism analogous to that described in the dipteran insect, 
Rhagoletis pomonella. Indeed, Stoffolano & Yin (1987) believe that 
mechanosensilla continue to monitor pressure on the ovipositor until 
it can go no further and, in this case, the lack of pressure would signal 
the maximum depth. Furthermore, another possible method would be 
to have tactile sensilla located along the most distal and the most 
proximal portions of the ovipositor. In the carpet moth, circling the 
8th segment and the terminal lobes of the telescoping ovipositor are 
presumed tactile sensilla trichodea A, B, and C that are believed to 
monitor the suitable depth of egg placement. 


The mechanosensilla of the ovipositor may also serve in copulation 
in providing the female moth information regarding the male position 
during copulation. 


199 


References 


ALTNER, H., RouTIL, C. & Lortus, R., 1981. The structure of bimodal che- 
mo-, thermo-, and hygroreceptors sensilla on the antenna of Locusta 
migratoria. Cell Tissue Res. 215 : 289-308. 

ANDERSON, P. & HALLBERG, E., 1990. Structure and distribution of tactile 
and bimodal taste-tactile sensilla on the ovipositor, tarsi and antennae 
of the flour moth, Ephestia kuehniella (Zeller) (Lepidoptera : Pyralidae). 
Int. J. Insect Morph. Embryol. 19 : 13-23. 

CHADHA, G. K. & RooME, R. E., 1980. Oviposition behaviour and the sensilla 
of the ovipositor of Chilo partellus and Spodoptera littoralis (Lepidop- 
tera : Noctuidae). J. Zool. Lond. 192 : 169-178. 

FAUCHEUX, M. J., 1982. Les récepteurs sensoriels impliqués dans le comporte- 
ment de ponte chez Monopis crocicapitella Clem. et autres Lépidoptères 
Tinéides à larves kératophages. Annis Cent. Reg. Doc. Pédag. Nantes, 
Biol. 1 : 65-119. 

FAUCHEUX, M. J., 1987. Recherches sur les organes sensoriels impliqués dans 
le comportement de ponte chez deux lépidoptères à larves kératophages : 
Tineola bisselliella Humm. et Monopis crocicapitella Clem. Thèse Doct. 
état, Univ. Nantes, 511 pp. 

FAUCHEUX, M. J., 1988. Multiporous sensilla on the ovipositor of Monopis 
crocicapitella Clem. (Lepidoptera: Tineidae). Int. J. Insect Morph. 
Embryol. 17 : 473-475. 

FAUCHEUX, M. J., 1991. Morphologie et organes sensoriels de l’ovipositeur 
de trois pyrales d’importance économique: Ostrinia nubilalis Hbn., 
Ephestia kuehniella (Zell.) et Homoeosoma nebulella Den. & Schiff. 
(Lepidoptera : Pyralidae). Bull. Soc. Sci. nat. Ouest Fr. 13 : 17-24. 

FAUCHEUX, M. J. & CHAUVIN, G., 1981. Les récepteurs sensoriels impliqués 
dans la recherche du support de ponte chez Tinea pellionella L. (Lepi- 
doptera : Tineidae). Archs. Zool. exp. gén. 122 : 5-16. 

REYMONET, C. & FAUCHEUX, M. J., 1991. Wall-pore sensilla on the ovipositor 
of the european sunflower moth, Homoeosoma nebulella Denis & Schif- 
fermüller (Lepidoptera : Pyralidae). Annis. Soc. ent. Fr. 27 : 119-120. 

SCHAFER, R. & SANCHEZ, T., 1976. The nature and development of sex 
attractant specificity in cockroaches of the genus Periplaneta. I. Sexual 
dimorphism in the distribution of antennal sense organs in five species. 
J. Morph. 149 : 139-158. 

SLIFER, E. H., 1960. A rapid and sensitive method for identifying permeable 
areas in the body wall of insects. Ent. News. 71 : 179-182. 

STOFFOLANO, J. G., Jr. & Yin, L. R. S., 1987. Structure and function of the 
Ovipositor and associated sensilla of the apple maggot, Rhagoletis pomo- 
nella (Walsh) (Diptera: Tephritidae). Int. J. Insect Morph. Embryol. 
16 : 41-69. 

VALENCIA, L. & Rice, M. J., 1982. Contact chemoreceptors on the ovipositor 
of the potato moth, Phthorimaea operculella (Zell.) (Lepidoptera : 
Gelechiidae). Int. J. Insect Morph. Embryol. 11 : 121-128. 


200 


ZACHARUK, R. Y., 1985. Antennae and sensilla. pp. 1-70, in Kerkut, G. A. & 
Gilbert, L. I. (Eds) : Comprehensive Insect Physiology, Biochemistry and 
Pharmacology, vol. 6. Pergamon Press, Oxford. 

ZAGULAJEV, A. K., 1960. Faune de l’U.R.S.S. Insectes Lépidoptères. Tome IV, 
N° 3, Famille des Tineidae, 3 ème partie, sous-famille des Tineinae. 
268 pp. Ed. Académie des Sciences d’U.R.S.S., Moscou, Leningrad. 


201 


Nota lepid. 18 (3/4) : 202 ; 13.V.1996 ISSN 0342-7536 


Book review — Buchbesprechung — Analyse 


An Illustrated Key to European Sesiidae (Lepidoptera). Z. LASTÜVKA 
& A. LASTÜVKA. 174 pp, numerous b/w and 8 col. pls. 15 X 21 cm, 
paperback. Faculty of Agronomy MUAF, Brno, Czech Republic, 1995. 
ISBN : 80-7157-151-2. Obtainable from : Department of Zoology and 
Apiculture, Faculty of Agronomy MUAF, Zemédélska 1, 613 00 Brno, 
Czech Republic. Price : CK 600 (= DM 34 ; postage DM 6). 


This book represents a milestone in modern clearwing research. It is the first 
work covering the entire European clearwing moth fauna since Bartel in Seitz 
(1912). Considering the price and the aim of this volume, there is little that 
can be faulted. 


After short introductory chapters covering such topics as phylogeny and 
classification, biology, collecting and rearing, keys to the genera and species 
based entirely on external characters are given, with many references to the 
figures found in the main part of the work. A short description is given for 
each of the 105 species, together with information on bionomics and habitat, 
and distribution. Diagnostic characters are illustrated in an accompanying 
plate, e.g. parts of the male and female genitalia, abdominal rings and fore- 
wings. The plate also includes a small distribution map. Adults of all species 
are illustrated in eight photographic coloured plates. 


The authors honestly state that while most species can be identified easily with 
the use of the keys, to determine other species requires considerable experience. 
For some species, resort to the genitalia will solve the problem, but for others 
even a combination of external characters and genitalia will not lead to a 
satisfactory result. In these cases one will require knowledge of the foodplant. 
Of course, to be able to use the keys at all, it is essential that specimens 
are in good condition. 


The coloured plates of adults are very good and the diagnostic features well 
presented. The distribution maps give one a good overview of where each 
species is known to occur. Synansphecia triannuliformis is not given for 
Switzerland, whereas it has been recorded from many localities in the south 
of the country. 


This book will persuade many more European lepidopterists to study this 
difficult family and it will help them to focus on the problem species and 
poorly studied areas of Europe. 


Steven WHITEBREAD 


202 


Nota lepid. 18 (3/4) : 203-212 ; 13.V.1996 ISSN 0342-7536 


Notes on the Orthosia rorida (Frivaldsky, 1835) 
species group, with the description of a new species 
from Crete : Orthosia sellingi sp. n. 

(Lepidoptera, Noctuidae, Hadeninae) 


Michael FiBiGER*, Hermann HACKER** & Arne MOBERG*** 


*Molbechsalle 49, DK-4180 Sorg, Denmark 
**Kilianstr. 10, D-96231 Staffelstein, Germany 
***Viborgskavagen 24, S-12237 Enskede, Sweden 


Summary 


Notes on species in the Orthosia rorida (Frivaldsky, 1835) group are given. 
Orthosia sellingi sp. n. is described from the Greek island of Crete. O. gani- 
metae Kornosor & Lödl, 1990 is synonymised with O. rorida. 


Zusammenfassung 


Orthosia sellingi sp. n. wird aus Kreta beschrieben. Nebst Informationen zu 
anderen Arten der Orthosia rorida (Frivaldsky, 1835) Gruppe wird das Taxon 
O. ganimetae Kornosor & Lödl, 1990 mit O. rorida synonymisiert. 


Resume 


Description d’Orthosia sellingi sp. n. de Crête. Les auteurs renseignent sur 
d’autres especes du groupe d’Orthosia rorida (Frivaldsky, 1835) et synonymisent 
le taxon ©. ganimetae Kornosor & Lödl, 1990 avec O. rorida. 


The Greek island of Crete in the eastern Mediterranean is well known 
for its many endemic species and subspecies. In this way it is similar 
to some of the other islands in the Mediterranean : Cyprus, Corsica 
and Sardinia. Cyprus and Crete are the two entomologically less well 
studied islands and new species are still being discovered. 


On a trip to Crete in 1993, Äke Selling, although unable to join the 
rest of his family, managed to equip his wife and son with some sugar 
ropes. The sugar ropes were hung up near Agra Marina Stalos and, 
among others species, seven specimens of a noctuid species which 
looked like O. rorida Frivaldsky, 1835 were captured. It was later dis- 
covered that the specimens represented a new species. Other specimens 


203 


of this new species were found in the material from Crete collected 
by Hans Malicky (now in coll. Hacker, Behounek and Thôny). 


Orthosia sellingi sp. n. 


Ho.otyPeE : | 4, Greece, Crete, west of Agra Marina Stalos, 2.1V.1993 
(gen. prep. Fibiger) (leg. Selling, coll. Fibiger). 


PARATYPES : | © (allotype) (gen. prep. Fibiger) (coll. Selling) and 6 && 
from the same locality as the holotype, 2.-14.1V.1993 (allotype 
14.1V.1993) (coll. Selling, Fibiger & Moberg) ; 3 64, Crete, Kastellakia, 
4.11.1978, 27.11.1978 (gen. prep. Hacker N 6226, 6224, 6279) (leg. 
Malicky, coll. Hacker) ; 2 44, Crete, Meso Potami, 900 m, 23.11.1982 
(gen. prep. Hacker N 6213) (leg. Malicky, coll. Hacker) ; 6 38, Crete, 
Kastellakia, 24°29’/35°22’, 18.11.-15.111.1978, dto 1 ©, 22.11.1978 (leg. 
Malicky, coll. Behounek) ; 5 88, 19, Crete, Sisses, 24752735247 
11.-22.11.1978, 1 4, 12.1.1979, (leg. Malicky, coll. Thöny) ; 2 38,3 28, 
Crete, Mt. Ida, 1600 m, 23.1V.1995; 1&, Crete, Kalami, 700 m, 
22.1V.1995 ; 1 4, Crete, Hora Spakion, 30 m, 21.1V.1995 (leg. Fibiger 
& Selling). 


Description 


MALE (Fig. 1): Wingspan 37-38 mm. Antenna strongly bipectinate. 
Labial palpı short, dark coloured. Head, thorax and ground colour 
of forewing grey. Forewing suffused with black scales. Reniform stigma 
defined by black spot, other stigmata hardly visible. Crosslines weakly 
marked, often absent. Subterminal line most often absent, but sometimes 
visible close to costa or represented by two spots between reniform 
and termen, as in Orthosia munda (D. & S.). Dorsum shaded darker 
medially in some specimens. Terminal line indicated by row of black 
dots betwen veins. Abdomen and hindwings unicolorous dark, ochreous 
brown. Terminal line absent. 


FEMALE (Fig. 2) : Wingspan 35 -37 mm. Antenna bipectinate, lamellae 
half the length of those of male. Coloration and wing-pattern as in 
male. | 


MALE GENITALIA (Figs 6, 7): Uncus short, pointed. Juxta twice as 
long as broad ; broadest basally. Valves broad, abruptly narrowing 
distally to a short pointed cucullus, directed ventrally. Clasper long, 
pointed, curved to almost 90°, not reaching tip of cucullus. Sacculus 
heavily sclerotised. Vinculum V-shaped. Aedeagus long, narrow. Carina 
heavily sclerotised, with field of short spines terminally. Everted vesica 


204 


2 


Figs 1, 2. Orthosia sellingi sp. n. 1 — Holotype, & ; 2 — Allotype, ©. Photos G. Brovad, 
Zoological Museum, University of Copenhagen. 


short, of same width as aedeagus, projecting 180° from tip of aedeagus. 
Two separate rows of long cornuti. 


FEMALE GENITALIA (Fig. 8) : Anal papillae short, broad. Both apophyses 
short, the anterior apophyses reaching beyond antrum. Antrum, ostium 
and ductus bursa heavily sclerotised. Ventral fissure in ostium, equally 
narrow at top and base. Ventral plate of seventh segment strongly 
sclerotised, broadly rounded. Ductus bursae slightly bent lateral- 
ventrally. Base of corpus bursae and appendix bursae moderately 
sclerotised. Corpus bursae with four signa-belts placed opposite each 
other. These are difficult to see when the corpus bursae contains a 
spermatophore. 


Differential diagnosis 
The known specimens of O. sellingi sp. n. exhibit small superficial 
differences to O. rorida Friv. (Figs 3-5). The ground colour of the 


forewing is pure grey in O. sellingi sp. n., whereas in the other species 


205 


Figs 3-5. Orthosia rorida (Frivaldsky, 1835). 3 —- 4, Macedonia, Katcanovo, 250 m, 
9.1V.1982. Leg. P. Jaksic, coll. M. Fibiger ; 4 — @, Greece, Kilkis, 350 m, 1.1V.1985. 
Leg. M. Fibiger ; 5 — @, Greece, Evro, Kirki, 500 m, 2-3.1V.1985. Leg. M. Fibiger. 
Photos G. Brovad. 


206 


Figs 6, 7. Male genitalia of Orthosia sellingi sp. n. 6a,b — Holotype (gen. prep. Fibiger 
1883) ; 7a,b — Crete, Kastellakia, 27.11.1978, (gen. prep. Hacker 6279). 


Fig. 8. Male genitalia of Orthosia rorida (Frivaldsky, 1835). Greece, Kilkis, 350 m, 
1.1V.1985 (gen. prep. Fibiger 1893). 


of the O. rorida group the ground colour is suffused brownish. Further, 
the terminal line of the hindwing is absent in O. sellingi sp. n., but 
almost always present in O. rorida Friv. The genitalia differences are 
marked. For comparison, the male and female genitalia of O. rorida 
Friv. are also figured (Figs 8, 10) together with the female genitalia 
of O. wolfi Hacker, 1988 (Fig. 11). The male of O. wolfi Hacker is 
unknown. 


The clasper in O. rorida Friv. is the same length as that of O. sellingi 
sp. n., but the valve is much shorter so that the tip of the clasper 
reaches the cucullus. The valve in O. sellingi sp. n. narrows abruptly 
distally, whereas it narrows only gradually in O. rorida Friv. The juxta 
in O. rorida Friv. is small, broader than long, but longer than broad 
in O. sellingi sp. n. Apart from the quite different form of the everted 
vesica between the two species, only a single field of long cornuti is 
present in ©. rorida Friv., compared to two in ©. sellingi sp. n. 


In the female genitalia the differences between the three species are 
most obviously seen in the ductus bursae and ostium. The ostium is 
a narrow slit in O. sellingi sp. n., U-shaped in O. rorida Friv. and 
broadly U-shaped in O. wolfi Hacker. The ductus bursae is bent in 
O. sellingi sp. n. and O. rorida Friv., whereas it is straight in O. wolfi 


208 


Hacker. In ©. sellingi, the sclerotised plate around the ostium the edge 
towards the corpus bursae is broadly rounded ; in ©. rorida Friv. it 
is almost straight. 


The new species is named in honour of our Swedish colleague Axe 
SELLING. 


Notes on other taxa of the O. rorida group 


The taxon Orthosia ganimetae Kornosor & Lödl, 1990 was described 
from a single male specimen from Adana, Turkey. It was compared 


Fig. 9. Female genitalia of Orthosia sellingi sp. n. Allotype (gen. prep. Fibiger 2067). 


209 


with the sympatric rorida Friv., from which it was considered to differ 
by a more weakly marked forewing and differences in the genitalia. 
However, the specimen falls within the variation of rorida and the 
apparent differences in the tips of the valves would seem to be due 
to a preparation artefact. More importantly, the everted vesica and 
the single bundle of cornuti are not different to that found in rorida. 
Orthosia ganimetae Kornosor & Lödl, 1990 is therefore considered 
to be a synonym of O. rorida Frivaldsky, 1835, syn.n. | 


Fig. 10. Female genitalia of Orthosia rorida (Frivaldsky, 1835). Greece, Alexandropolis, 
Kirki, 2.-3.1V.1985 (gen. prep. Fibiger 2084). 


210 


The taxon O. rorida mundoides Boursin, 1940 was considered by 
Kornosor & Lödl (1990) to be infrasubspecific and therefore unavailable. 
However, it was first described as a “form or sp.”, and later in the 
same paper as a subspecies. It was described from a female from the 
Lebanon and a male from Albania. The Lebanon specimen was 
considered to represent a subspecies by Ellison & Wiltshire (1939). The 
taxon could therefore be considered as available. The present authors 
have not had the possibility to examine mundoides. However, having 


Fig. 11. Female genitalia of Orthosia wolfi (Hacker, 1988). Holotype (gen. prep. Hacker 
4073). 


230 


studied the remarkable infraspecific variation of O. rorida, it is our 
opinion that the name mundoides will probably go into synonymy 
with O. rorida Friv. 


Technical note 


To be able to compare genitalia figures it is important to be consistent 
when making the preparation. The valve of the species in the O. rorida 
Friv. group is heavily sclerotised and the tips are bowed towards each 
other. When mounting under a coverslip it is necessary to unfold and 
flatten the valves, i.e. press the tips to the same horizontal position 
as the rest of the valves. The same rules on consistency of preparation 
must be followed with everted vesica preparations. It is proposed that 
figures of aedeagi with everted vesica are consistently displayed from 
the left lateral side with the apex of the aedeagus and base of the vesica 
to the right and the aedeagus seminalis preferably pointing upwards 
(however, sometimes the shape of the vesica makes this impossible). 


Acknowledgements 


Barry Goater, Chandler’s Ford, England and Mariann Fibiger are thanked for 
checking the English language and Martin Honey, Natural History Museum, 
London, for information on the taxon mundoides Brsn. 


Literatur 


Boursin, Ch., 1940. Beiträge zur Kenntnis der “Agrotidae-Trifinae” XXIII. 
I. Neue palaearktische Arten und Formen mit besonderer Beriicksich- 
tigung der Gattung Autophila Hb. Mitt. münch. ent. Ges. 30 : 474-543, 
pl. VIII-XN. 

ELLISON, R. & WILTSHIRE, E. P., 1939. The Lepidoptera of the Lebanon ; 
with notes on their season and distribution. Trans. R. ent. Soc. Lond. 
88 : 1-56. 

HACKER, H., Huser, K. & KUHNA, P., 1988. Achter Beitrag zur systematischen 
Erfassung der Noctuidae der Tiirkei. Beschreibung von sechs neuen Taxa 
und Notizen über bemerkenswerte Funde aus neueren Aufsammlungen 
(Lepidoptera). Atalanta 18 : 339-369. 

Kornosor, S. & Lopı, M., 1990. Two new species of the genera Orthosia 
Ochsenheimer, 1816 and Simyra Ochsenheimer, 1816 from Turkey 
(Noctuidae, Lepidoptera). Z. ArbGem. 6st. Ent. 42(3/4) : 68-72. 


DAD. 


Nota lepid. 18 (3/4) : 213-223 ; 13.V.1996 ISSN 0342-7536 


Hesperioidea and Papilionoidea of the Ionian island 
of Kefalonia, Greece : Additional species 
and a review of faunal components (Lepidoptera) 


David E. GASKIN 


Department of Zoology, University of Guelph, Guelph, Ontario, Canada NIG 2W1 


Summary 


Lepidoptera in some regions of Greece have been studied intensively during 
the last 20 years, but those of the large Ionian island of Kefalonia (38° 15’N, 
20°30’E) remain poorly known. The present article reports on 16 additional 
butterfly species not encountered by Gaskin and Littler (1986), bringing the 
confirmed total to 45, although this number will surely increase with further 
surveys. The recorded composition of the butterfly fauna of Kefalonia is re- 
viewed in the context of the ecology and historical biogeography of the region 
and it is speculated why some ubiquitous species have yet to be recorded. 


Résumé 


Les Lépidoptères de quelque régions de la Grèce ont fait l’objet de recherches 
intensives durant ces vingt dernières années ; ceux de l’île Ionienne de Cépha- 
lonie (38° 15’N, 20° 30’E) sont en revanche peu connus. Le présent travail signale 
16 espèces de Lépidoptères diurnes que Gaskin & Littler (1986) n’avaient pas 
trouvées, ce qui donne un total de 45 espèces. ce nombre sera certainement 
encore accru par de nouvelles recherches. La composition de la population 
des Lépidoptères diurnes connue de Céphalomie est appréciée sur le plan de 
l'écologie et de la biogéographie historique de la région. L’auteur se demande 
pourquoi certaines espèces ubiquistes n’ont pas encore été signalées. 


Introduction 


The butterfly fauna of several regions of Greece has been studied quite 
intensively. However, except for Kérkira (Corfu) (Baldock & Bretherton, 
1981), the islands of the Ionian chain have been rather neglected. I have 
found no published accounts specific to the Lepidoptera of Kefalonia, 
although four major studies of the flora have been published (Heldrich, 
1882 ; Bornmiiller, 1928 ; Cufodontis, 1936, & Knapp, 1965). Gaskin 


218 


& Littler (1986) surveyed 13 sites across Kefalonia during the first 
half of May 1983, but three large areas, the mid-south-east, mid-north 
central and the south coast could not be investigated. Most of these 
localities were re-examined and seven new sites, all in the three regions 
missed in 1983, were surveyed in the first half of June 1992. The sites 
at Lassi and Minia were found to have been drastically modified by 
developments associated with the tourist industry. 


Methods 


Each site was examined by two observers for a minimum of two hours 
and for half a day whenever possible (see Fig. 1). Priority was given 
to focal searches of each discernible type of habitat to maximize 
potential contact with all species flying at each locality. There is as 
yet no universally accepted system of anglicizing Greek place names. 
In this contribution I follow Gaskin & Littler (1986), in spelling names 
of most localities as in the Toubis series of maps of the Grecian islands, 
which are published in Athens and widely available to visitors. In cases 
where these are clearly at odds with the pronunciation of demotic 
Greek, e.g. “Chios”, I have followed the Times Atlas (Bartholomew 
et al., 1990), in this case “Khios”, which is a more accurate rendition. 
On the other hand, the Toubis version “Kefalonia” is far more often 
used than the Times Atlas “Kefallinia” ! 


Results 


The forty species observed during the first half of June 1992 are given 
below ; the 16 recorded for the first time on Kefalonia are indicated 
by asterisks. Five species were recorded only in May 1983: Papilio 
alexanor Esper, 1799, Euchloe ausonia (Hübner, 1804), Gonepteryx 
rhamni (Linnaeus, 1758), Callophrys rubi (Linnaeus, 1758), and Glau- 
copsyche alexis (Poda, 1761). 


Hesperioidea 


Spialia orbifer (Hübner, [1823]) : ENS, TZA. 

Carcharodus alceae (Esper, 1780)* : AGN, PLA.TZA. 

Carcharodus orientalis Reverdin, 1913* : AGN. 

Thymelicus acteon Rottemburg, 1775* : PDK, TZA. 

Thymelicus flavus (Brünnich, 1763) : AGN, CAG, ENS, PDK, TZA, VAV. 


214 


| Sn 38° 30’N 


38°15’ N 


20°45/E 


Fig. 1. Butterfly observation sites on Kefalonia. June 1992 (solid circles) : Ag. Eufimia 
AGE; Ag. Nikolaos AGN ; Argostöli ARG ; Castle of Ag. Giörgios CAG ; Mt. Enos 
ENS ; Frangata FRG; Perdikata PDK ; Platies PLA ; Tzanata TZA ; Vathi Avlaki 
VAV ; Zervata ZRV. Four 1986 sites were unproductive in 1992 ; Ag. Irini IRI; Lassi 
LAS ; Katavöthres KAT, and Miniä MIN. Other sites worked by Gaskin and Littler 
(1986) are shown by open circles and identified by single letters: Assos A ; Fiskardo 
F ; Kouroukläta K ; Lixouri L ; Poros P ; and Sami S. Broken lines indicate seasonal 
or subterranean water-courses. 


215 


Papilionoidea 


Papilio machaon Linnaeus, 1758 : CAG. 

Iphiclides podalirius (Linnaeus, 1758) : AGN, CAG, FRG, PDK, TZA, ZRV. 

Pieris brassicae (Linnaeus, 1758) : AGN, TZA. 

Pieris rapae (Linnaeus, 1758) : ARG, PDK, TZA. 

Aporia crataegi (Linnaeus, 1758) : AGN, ENS, PDK, TZA, ZRV. 

Colias crocea (Fourcroy, 1785) : AGN, CAG, ENS, PDK, PLA, TZA. 

Gonepteryx farinosa (Zeller, 1847): AGE, AGN, CAG, ENS, FRG, PDK, 
PLA, TZA, VAV. 

Gonepteryx cleopatra (Linnaeus, 1767) : AGN, CAG, ENS, FRG, PDK. 

Leptidea sinapis (Linnaeus, 1758) : CAG, FRG, TZA. 

Charaxes jasius (Linnaeus, 1776)* : TZA, VAV. 

Limenitis reducta Staudinger,1901 : AGN, ENS, PDK, PLA, TZA, VAV. 

Nymphalis polychloros (Linnaeus, 1758)* : TZA. 

Polygonia egea (Cramer, 1775) : CAG,TZA. 

Cynthia cardui (Linnaeus, 1758) : AGN, CAG, ENS, FRG, PLA, TZA, VAY, 
ZRV. 

Vanessa atalanta (Linnaeus, 1758) : AGE, CAG, VAV. 

Argynnis paphia (Linnaeus, 1758)* : TZA. 

Melitaea didyma (Esper, 1779)* : CAG. 

Hipparchia syriaca (Fruhstorfer, 1908) : ENS, PDK, VAV. 

Hipparchia volgensis Mazochin-Porshjakov, 1952* : AGE, AGN, CAG, ENS, 
FRG, PDK, TZA, VAV, ZRV. 

Maniola jurtina (Linnaeus, 1758): AGE, AGN, CAG, ENS, FRG, PDK, 
TZA, VAV, ZRV. 

Coenonympha pamphilus (Linnaeus, 1758) : CAG. 

Lasiommata megera (Linnaeus, 1767) : ENS, PLA. 

Lasiommata maera (Linnaeus, 1758) : CAG. 

Pararge aegeria (Linnaeus, 1758)* : CAG, TZA. 

Kirinia roxelana (Cramer, 1777)* : TZA. 

Nordmannia ilicis (Esper, 1779)* : AGN, CAG, ENS, FRG, TZA, VAV. 

Strymonidia spini (Denis & Schiffermiiller, 1775)* : AGN, CAG, FRG, PDK, 
PLA. 

Lycaena phlaeas (Linnaeus, 1761) : AGN, ENS, PDK, TZA. 

Lampides boeticus (Linnaeus, 1767)* : CAG, FRG. 

Syntarucus pirithous (Linnaeus, 1767)* : ENS. 

Celastrina argiolus (Linnaeus, 1758)* : PDK, PLA, TZA. 

Pseudophilotes vicrama (Moore, 1865) : AGN, CAG, FRG, PDK, PLA, VAV. 

Aricia agestis (Denis & Schiffermiiller, 1775) : CAG, ENS, PDK, PLA. 

Agrodiaetus thersites Cantener, 1834* : AGN. 

Polyommatus icarus (Rottemburg, 1775) : ENS, PDK, PLA. 


Discussion 


As on most Grecian islands, the extant butterfly fauna of Kefalonia 
will have been influenced historically not only by physical and climatic 


216 


factors and the composition of the flora, but also by modification of 
habitats by human settlers and their domestic animals. This pre-dates 
the classical period, as Paleolithic and Neolithic stone tools have been 
found on Kefalonia (Melas, 1985). The processes that structured the 
composition of the modern butterfly fauna of Greece are likely to be 
at least as complex as those described by Dennis (1977) for the British 
Isles. 


For a preliminary and non-rigorous zoogeographic exercise from a 
“Graeco-centric” point of view, based on 216 species recorded for 
Greece to the early 1990s, I allocated the butterfly fauna of Greece to 
four distributional groups, using the maps of Higgins & Riley (1983) 
and information from subsequent publications by Coutsis (1984, 1986), 
Coutsis et al.(1989), Coutsis & Ghavalas (1988 ; 1991), Fuchs (1987), 
Gallo (1989), Koutroubas (1991 ; 1992; 1993; 1994), Leestmans & 
Arheilger (1988), Olivier (1987 ; 1988 ; 1989a ; 1989b ; 1990 ; 1993), Van 
der Poorten (1981 ; 1985a ; 1985b ; 1990), Van der Poorten et al. (1988), 
Schmidt & Hasler (1986), Thomson (1985 ; 1987a ; 1987b ; 1990) and 
Vanholder (1993). Coutsis informed me very recently that the species 
total had now reached 223, but the increase does not modify present 
conclusions significantly. 


Group I consists of 123 species, ubiquitous at least through central 
and southern latitudes of Europe, though some are restricted to specific 
habitats or altitude zones within this range. Because of the evolution 
of marked regional endemism I omitted most species of Erebia. Group 
II comprises 30 species with essentially Mediterranean distributions, 
Group III contains 45 which occur in Greece, but are confined to south- 
eastern Europe or penetrate into that region from the east, while Group 
IV consists of 18 species endemic to the Grecian peninsula or 
archipelagos. These numbers convert to the following percentages : 
Group I - 57%, Group II - 14%, Group II - 21% and Group IV - 8%. 
Equivalent values for the known butterfly fauna of Kefalonia are: 
Group I - 71%, Group II - 18%, Group HI - 11% and Group IV - 0%. 
A statistical comparison using a x? test, with Groups III + IV lumped, 
showed that the frequencies differed significantly from those for Greece 
as a whole (x? = 6.9, p < 0.05). However, if Groups II, HI and IV 
were lumped and compared with Group I, then the differences between 
Kefalonia and Greece are not significant and x? drops to 2.8 (p > 0.05). 


The 45 butterfly species recorded from Kefalonia to date depict a fauna 
typical of larger offshore islands, i.e. depauperate in comparison to the 
adjacent mainland, yet still reasonably representative. The dominance 
of Group I (wide-spread European) species on Kefalonia is predictable. 


217 


Many of the butterfly species abundant in Greece are quite vagile, and 
some are seasonal migrants. The Ionian Islands have few central 
European elements in their floras (Turrill, 1929), but nevertheless, in 
some areas of the east coast of Kefalonia narrow belts of deciduous 
trees, more typical of northern Greece and central Europe, crowd the 
margins of a few narrow and precarious riparian corridors. Four of the 
Group I species recorded on Kefalonia in June 1992 were only in the 
riparian margins. Yela (1992) emphasized the importance of protecting 
such habitat to maintain biodiversity in Iberian ecosystems ; this should 
be a consideration in conservation strategies in Greece also. Other 
Group I species, locally abundant on the mainland, but perhaps less 
vagile, have yet to be recorded from Kefalonia, e.g. Melanargia galathea 
(Linnaeus, 1758), M. larissa (Geyer, 1828) and Hyponephele lupina 
(Costa, 1836). Such taxa best exhibit movement when suitable habitat 
is continuous (Shreeve, 1992), sa, presumably these species might be 
poor candidates for successful island-hopping. 


As one might anticipate from its geographical position, the proportion 
of Group II species (Mediterranean) reported from Kefalonia is not 
dissimilar to that of the mainland, while the under-representation of 
Group III (south-eastern) species reflects Kefalonia as an offshore island 
at the western extremity of the Grecian peninsula, relatively distant 
from the source. 


The lack of endemic elements in the Kefalonian flora noted by Heldrich 
(1982), Bornmiiller (1928), Cufondontis (1936), Knapp (1965) and 
Polunin (1980) is consistent with relatively recent and perhaps frequent 
links to the mainland during the Pleistocene. The probability of finding 
insect species endemic to Kefalonia is low in these circumstances. Many 
butterflies endemic to Greece are characteristic of and restricted to 
montane habitats, so the lack of such Group IV species on Kefalonia 
is not surprising. The central Enos (Ainos) Massif may have supported 
a true alpine fauna at some time in the past, but it is too small in 
area, too low and too far south to support one in the present climatic 
regime (see Huxley & Taylor, 1984 and Sfikas, 1987). 


Neverthless, there are still puzzles to be solved. The coastal islands 
of the eastern Aegean Sea also have a complex (and better under- 
stood) history of connections and segregations during the late Tertiary 
and Quaternary (Dermitzakis, 1990 ; Dermitzakis & Goedicke, 1977 ; 
Dermitzakis & Sondar, 1979 ; Muelencamp, 1985), yet in this region 
three endemic butterflies have been discovered recently. These are 
Maniola chia Thomson, 1987 on Khios, M. halicarnassus Thomson, 
1990 on Nissiros (also on Bodrum Peninsula) and an endemic sub- 


218 


species, Satyrium ledereri christianae Olivier, 1989, on Samos. More 
may be identified (Olivier & Coutsis, 1993 ; Thomson, 1990), despite 
isolation of the islands from the mainland of Turkey that is unlikely 
to be much older than 20,000-75,000 B.P. Yet Kefalonia (33 km from 
Pelopönnisos, 40 km from Stereä) is about twice as far from the 
mainland of western Greece as are Khios (12-19 km) and Samos 
(11 km) from the coast of Anatolia, and there is a trench of relatively 
deep water through the channel from Levkas, to the Gulf of Kiparissia 
off the west coast of the Pelopönnisos. However, if one takes into 
account the extended shelf that must have existed at various times 
in the Pleistocene around Ithaki, Kälamos and other small islands 
between Kefalonia and Levkas (19 km apart), distances were probably 
comparable. Probably there was and is, little hindrance to intermittent 
dispersal of vagile winged insect species between these islands and 
prevailing winds presumably play an important role in seasonal and 
year-to-year variations in such a flow. Perhaps some taxa are pre- 
disposed to more rapid rates of genetic segregation than others, and 
studies of morphology alone will not necessarily produce a good 
measure of this (Collins, 1991). 


Detailed studies of butterfly populations on Kefalonia will be necessary 
before the differences between the Ionian and the eastern Aegean 
regions can be fully understood. More basic surveys are also needed. 
The addition of another 10-15 species to the Kefalonian list after further 
research would not be surprising, because survey time expended on 
the island to date is still barely one month. Excursions in the spring 
should discover more pierids and the hotter months should produce 
more species of the larger satyrids of Groups II and II. Intensive 
attention to the riparian zones should be profitable in both cases. While 
discovery of Nymphalis polychloros and Kirinia roxelana on Kefalonia 
were pleasant surprises, the failure to find Anthocharis cardamines 
(Linnaeus, 1758), Pontia edusa (Fabricius, 1777) and Argynnis pandora 
(Denis & Schiffermiiller, 1775) can be ascribed perhaps to bad luck 
with timing or weather conditions. In general, however, I would predict 
that investigations will further increase the preponderant representation 
of Group I species on Kefalonia. 


Acknowledgements 


I thank Elizabeth A. Littler of Alpena, Michigan, for making her specimens 
and records from Kefalonia available for this study ; John G. Coutsis of Athens 
for numerous valuable discussions, and confirming the identity of the Hippar- 
chia sp. and worn specimens of some other species from Kefalonia. John, and 


219 


a second anonymous reviewer, also gave valuable advice on some taxonomic 
points, and provided specific suggestions for improving the manuscript. 


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225 


Nota lepid. 18 (3/4) : 224 ; 13.V.1996 ISSN 0342-7536 


SIEEC 
XV. Internationales Symposium über Entomofaunistik 
in Mitteleuropa 


22.-27. September 1996 
Iasi, Rumänien 


Das 15. Internationale Symposium über Entomofaunistik in Mittel- 
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224 


Nota lepid. 18 (3/4) : 225-232 ; 13.V.1996 ISSN 0342-7536 


Dahlica wehrlii (Müller-Rutz, 1920) wieder gefunden. 
Beschreibung des Weibchens und Ergänzungen 

zur Kenntnis des Männchens und der Okologie 
(Lepidoptera, Psychidae) 


Peter HATTENSCHWILER 
Seeblickstrasse 4, CH-8610 Uster, Schweiz 


Summary 


Dahlica wehrli (Müller-Rutz, 1920), from the high Alps in the Canton Wallis, 
Switzerland, represented by one single male specimen and not seen for over 
60 years, has been rediscovered. Additional descriptions for the male are given 
and the female, larva, case and ecological aspects are described. 


Zusammenfassung 


Die seit über 60 Jahren verschollene und nur durch ein Männchen bekannte Art 
Dahlica wehrlii (Müller-Rutz, 1920) aus den Walliser Hochalpen in der Schweiz, 
wurde wieder gefunden. Ergänzende Angaben zum Männchen werden gegeben 
und die Weibchen, Raupen, Säcke sowie die Okologie erstmals beschrieben. 


Résumé 


Dahlica wehrlii (Müller-Rutz, 1920), Psychide de haute montagne des Alpes 
valaisannes en Suisse, espèce connue par un seul exemplaire mâle et disparue 
depuis plus de 60 ans, vient d’être retrouvée. L’auteur donne de nouveaux 
renseignements sur le mâle et décrit pour la première fois les femelles, les 
chenilles et les fourreaux, ainsi que la biologie. 


Am 9. August 1919 hat Dr. E. Wehrli auf dem Trifthorn in der 
Zermatter-Gegend im Kanton Wallis auf 3730 m ü M ein Männchen 
einer Psychide gefangen, die dort in Mehrzahl flog. 1920 hat Miiller- 
Rutz dieses eine Männchen zu Ehren des Sammlers als Solenobia 
wehrlii beschrieben. Von diesem Männchen sind nur noch der Körper, 
Antennen und ein Hinterflügel vorhanden. Das Exemplar, das als Holo- 
typus zu betrachten ist, steckt in der Sammlung des Naturhistorischen 
Museums Basel. 


225 


Die Originalbeschreibung lautet : 


„Von allen schmalköpfigen Solenobia-Arten schon durch die Größe, 8,5 mm 
Lange der Vorderfliigel, zu erkennen. Diese sind schmal, sehr gestreckt, der 
Saum sehr schrag, der Vorderrand kaum erkennbar eingedriickt. Die Farbe 
ist ein mattes Graubraun, vielleicht eine Nuance dunkler als bei den übrigen 
Arten. Die hellen Flecken sind deutlich, nicht dicht stehend, sie bilden wenig- 
stens im Discus deutliche Längsreihen. Auch längs des Saumes, vom Innen- 
winkel bis um die Spitze, so weit die Vorderrandfransen reichen, stehen deut- 
liche Fleckchen, die teilweise in die Fransen übergreifen. Die Hinterflügel wenig 
schmaler als die vordern, ebenfalls viel gestreckter als bei den übrigen Arten, 
hellgrau, die Adern etwas dunkler, gut sichtbar ; IIP und IIB [m2 und m3] 
getrennt. Die Behaarung des Kopfes graubraun, der Thorax dunkel, Hinterleib 
heller braun, der Afterbusch gelbbraun, der Bauch sparsam gelbbräunlich 
behaart. Vorder- und Mittelbeine braun, die Füße ungefleckt, die Hinterbeine 
fehlen bei dem Exemplar. Die Bewimperung der Fühler nicht dicht, fast von 
doppelter Länge der Gliederbreite. 


In einem Exemplar am Gipfel des Trifthorns, 3730 m, wo das Tierchen in 
Mehrzahl flog, von Dr. E. Wehrli in Basel erbeutet, am 9. VIII. 19“. 


Seit diesem Fang auf dem Trifthorn blieb die Art verschollen, obwohl 
immer wieder Versuche unternommen wurden, Klarheit in diese Sache 
zu bringen. Den Originalfundort zu erreichen erfordert gut trainierte, 
berggewohnte Leute. Zudem sollten sie im Sachgebiet der Kleinschmet- 
terlinge versiert sein. Es wurde auch in der weiteren Umgebung des 
Fundortes gesucht. So fand am 28.7.1980 Peter Sonderegger auf dem 
Adlergletscher, VS in 3120 m ü M ein Männchen auf dem Schnee 
sitzend. Es war schon so stark unterkühlt, dass es nicht mehr wegfliegen 
konnte. Dieser Fundort liegt nur etwa 15 km Luftlinie vom Trifthorn 
entfernt. Eine gründliche Untersuchung ergab, dass es sich um das 
zweite Exemplar von D. wehrlii handeln musste. Durch diesen Fund 
wurde bestätigt, dass die Art in einem weiteren Umkreis des Trifthornes 
zu leben scheint. 


Der nächste Glücksfall ereignete sich erst 12 Jahre später, als am 15. Juli 
1992 Renato Joos und Heinz Buser auf der Täschalp, VS ın 2500 m 
ü M unmittelbar nach der Morgendämmerung sammelten. Sie fingen 
einige Dahlica Männchen, die zusammen mit den bekannten Pseudo- 
bankesia alpestrella (Heinemann) um die Felsen flogen. Der Vergleich 
der drei Männchen ergab eine gute Übereinstimmung mit demjenigen 
vom Adlergletscher. Damit war ein Fundort bekannt, der auch von 
nicht berggewohnten Sammlern erreicht werden kann. 


Am 12. Juni des nächsten Jahres konnte ich mit den Freunden Renato 
Joos und Heinz Buser ins Fundgebiet reisen um nach den noch un- 


226 


bekannten Säcken, Raupen und Weibchen zu suchen. Mit vereinten 
Kräften gelang es uns dann auch während einigen Stunden des Suchens 
16 Säcklein zu finden, die sehr versteckt in Felsritzen angesponnen 
waren. Offenbar sind die Falter dieses Jahr früher geschlüpft, das warme 
Wetter im Frühling kann das bewirken. In vereinzelten Säcklein steckten 
noch die Puppenhüllen was zeigt, dass der Schlupf vorüber war, aber 
auch, dass Säcke mit Eiern zu erwarten sind. Unsere Hoffnung erfüllte 
sich, unter den gesammelten Säcklein waren zwei mit Ei-Gelegen. 


Die Nachzucht in Uster auf nur 475 m ü M gelang recht gut und so 
erhielten wir 1994 36 Männchen und 34 Weibchen. Mit diesen um- 
fangreichen Belegen konnten weitere Vergleiche gemacht werden um die 
Zugehörigkeit zur gesuchten Dahlica wehrlii zu bestätigen. Das Zucht- 
material befindet sich in den folgenden Sammlungen in der Schweiz : 
Naturhistorisches Museum Basel, ETH Zürich, Renato Joos, Basel ; 
Heinz Buser, Sissach, BL; Willi Sauter, Illnau, ZH ; Ruedi Bryner, 
Twann, BE, sowie in meiner Sammlung. Im Ausland in den Samm- 
lungen : Franz Lichtenberger, Waidhofen, Österreich ; British Museum 
(Nat. Hist.), London. 


Freundlicherweise hat uns das Naturhistorische Museum Basel den 
Holotypus ausgeliehen und erlaubt ein Genitalpräparat zu erstellen, 
was durch Steven Whitebread ausgeführt wurde (Präp.Nr. E.569). Der 
Genitalindex (nach Sauter, 1956) des Holotypus ist 1,68. Aus den 
Beobachtungen während der Zucht ab Ei in Uster und den Vergleichen 
der Serie gewannen wir Kenntnisse die uns erlauben, einige Ergänzungen 
zur Originalbeschreibung bekannt zu geben. 


Dahlica wehrlii (Müller-Rutz, 1920) 
Solenobia wehrlii Müller-Rutz, 1920. Mitt. Ent. Zürich 5 : 348, Taf. II, Fig. 14. 


MANNCHEN : Flügelspannweite 14,5-17,5 mm, Vorderflügel schmal, ge- 
streckt, nach aussen nur wenig verbreitert, spitzer Apex, Saum sehr 
schräg, Zeichnung mit kleinen, locker, aber regelmässig angeordneten 
creme-weissen Flecken, Flügel-Deckschuppen der Klasse 2 (Sauter, 
1956), meist zweizackig. Fransen von gleicher Farbe wıe Flügelgrund, 
ohne Fleckenmuster. Hinterflügel einfarbig heller grau mit schmalen 
Schuppen der Klasse 1. Der Genitalindex ist 1,50-1,71, im Mittel 1,62. 


WEIBCHEN (Abb. 1) : Ohne Legeröhre 3-5 mm lang, 1-1,3 mm Durch- 
messer, blassgelb, Kopf dunkelbraun, die Brust- und Abdominalseg- 
mente mit braunen Dorsaplatten. Ventral mit je zwei dreieckigen, 
sklerotisierten Platten, deren Spitzen sich treffen oder getrennt sind. 


2211 


Abb. 1. Weibchen von Dahlica wehrlii von der Seite betrachtet. 


Fühler mit 17-20 Gliedern. Beine mit 4 Tarsengliedern. Die Afterwolle 
variabel in der Färbung, sie kann dunkelgrau sein oder am Körper 
dunkelgrau beginnen und nach aussen heller, fast weiss werden. 


RAUPE : 5,5 mm lang bei einem Durchmesser von 1,2-1,5 mm, weisslich- 
grau bis grau, Brustbeine braun, Kopf schwarz. 1. Brustsegment dunkel- 
braun, 2. Segment dorsal braun, seitlich je ein braunes Feld. Das 3. 
Segment nur noch mit 4 unscharfen, braunen Flecken. Rückenplatte 
des letzten Abdominalsegmentes bräunlich. Sie fressen Algen und 
Flechten die an den Felsen und Steinen wachsen seltener auch welke 
oder verwesende Pflanzenteile. Tagsüber sind sıe oft versteckt, teilweise 
in Bodennähe und steigen bei Dunkelheit an den Felsen empor um zu 
fressen. Sie sind schon bei relativ tiefen Temperaturen aktiv, so wurden 
Raupen beim Fressen beobachtet bei nur 1°C über dem Gefrierpunkt. 
Zur Überwinterung verkriechen sich die Raupen in Spalten und Ritzen 
an den Felsen, sie gehen seltener in die Bodennähe. Dieses Verhalten 
erlaubt das Schlüpfen und Paaren schon früh ım Jahr, auch wenn 
der Boden noch mit Schnee bedeckt ist. 


228 


Verschiedene Arten von kleinen Schlupfwespen parasitieren die Raupen, 
darunter auch solche aus der Gattung Gelis mit ungeflügelten Weibchen. 
Der Grad der Parasitierung dürfte jedoch nicht sehr hoch liegen. 


SACKBESCHREIBUNG : Die Säcke sind 5-7 mm lang, im Mittel 6 mm, 
bei einer Breite von 1,5-2,0 mm, im Mittel 1,7 mm. Es scheint, dass 
die Säcke der Weibchen im Mittel etwas kürzer aber leicht breiter sind 
als die angegebenen Mittelwerte. Die Seiten sind nach aussen gebaucht, 
die Rückenkanten meist nicht sehr deutlich jedoch etwas gröber ge- 
körnt. Der Belag ist fast ausschliesslich mineralisch, er besteht aus feinen 
Sandkörnchen (Abb. 2). 


eo SS 
“ort 
rc» 
+ & N a 
NE cp à 
ng eee te 
1, CHAT DA 
5 À D £a: 
3 fe: 4 [es EG 
Wee ‘2 
DES FA Wr 
EB ATS Ar. 
= i arr ex 
ES ae > EN 
25 Wr Se Ser ae 
Re a = Mr » CN: a 
ST. < BE SE 
NET: à LR CEA 
- >. AN TR = 
3 +: € Se ie R Se 
SSSR 7) x BEN. 
N =. SE DR; À à re 
2 LOTERIE: 7 © RT Mer 
> x ale: eo = 
EN. 7 CES | 
“ARS e4 ie 
~ ds GARE 
CNE Ss By 
Be ig Le QE) 
‘TL NE 
À CE ie WWE 
à od Wes 
ea ( ; YEP ES 
AIR LA 
AS N er: a 


Abb. 2. A+B Säcke von Dahlica wehrlii; C + D Säcke von Dahlica triquetrella, 
jeweils in Seitenansicht. (A + C) im Larvenstadium und (B + C) von dorsal im Puppen- 
stadium. (D. triquetrella-Sack nach Galliker, 1958, verändert). 


PUPPENBESCHREIBUNG : Die Brust-Kopfplatte der weiblichen Puppen 
mit Fühlerscheiden, die meistens wenig länger als die Beinscheiden sind, 
sie können aber auch etwa um die Scheidenbreite kürzer sein, oft sogar 
links und rechts unterschiedlich (Abb. 3). 


OKOLOGIE DER FALTER: Die geflügelten Männchen schlüpfen am 
Abend, bei bewölktem Himmel kann der Schlupf schon am Nachmittag 


229 


Abb. 3. Brust-Kopfplatte der weiblichen Puppe von Dahlica wehrlii. 


beginnen. Sie bleiben aber meistens fast unbeweglich bis zum Morgen 
sitzen und werden erst durch den Duft der lockenden Weibchen erregt. 
Die Weibchen schlüpfen erst bei Tagesanbruch. 


LEBENSRAUM: Felsen, Steine oder Mauern wo Algen oder Flechten 
wachsen ; auch wenn der Bewuchs für uns kaum sichtbar ist, finden 
die Räuplein genügend Futter. Die bevorzugten Stellen sind Felsen und 
Steinbrocken mit Spalten und Ritzen, ın die sich die Raupen zurück- 
ziehen können. Man trifft sie öfter an Felsen, die am Fuss leichten 
Grasbewuchs haben und die tagsüber besonnt sind. 


JAHRESZEITLICHE ENTWICKLUNG: Die Flugzeit beginnt nach der 
Schneeschmelze, von Ende April bis Juli, je nach Lage und Höhe. 
Obwohl an schattigen Stellen und am Fuss der Felsen der Schnee noch 
metertief liegt kann man aktive Raupen an den besonnten, schneefreien 
Felsen und Steinen erwarten. Die Falter schlüpfen aus den Puppen 
ab einer Umgebungstemperatur von nur +2°C. Die Entwicklung ist 
einjährig, die Raupen überwintern voll erwachsen und nehmen in der 
Regel nach der Winterruhe kein Futter mehr auf. Sofort nach der Über- 
winterung suchen die Raupen einen Ort zur Verpuppung in einer Ritze 
oder Nische auf und spinnen den Sack fest. Nach 2-4 Wochen schlüpfen 
die Falter. 


VERBREITUNG: Diese Art ist nur aus der Zermatter-Gegend in der 
Schweiz bekannt, dürfte aber in den angrenzenden italienischen Bergen 
auch heimisch sein. Die Höhenverbreitung reicht von 2300-2500 m ü M 


230 


auf der Täschalp, bis um 3730 m ü M auf dem Trifthorn, beide im 
Wallis. Einen weiteren Fund der sehr wahrscheinlich diese Art betrifft 
hat Steven Whitebread am 22.7.1984 auf 2170 m oberhalb Zermatt 
(Chüeberg) gemacht (einen Sack mit toter männlicher Puppe, Genital- 
index 1,64). 


Diskussion 


In den bekannten Fundgebieten von D. wehrlii sind keine Arten ge- 
funden worden, die leicht verwechselt werden könnten. Die partheno- 
genetische Form der Dahlica triquetrella Hübner hat Weibchen mit 
5 Tarsengliedern und deutlich grössere Säcke (Sauter, 1956 ; Galliker, 
1958) (Abb. 3), die oft am vorderen Ende mit Teilen von Insekten und 
Spinnen „verziert“ sind, die Säcke haben deutliche Kanten. D. wehrlii- 
Weibchen haben vier Tarsenglieder. Möglicherweise kann auch die 
parthenogenetische Form von Dahlica lichenella Linnaeus, die auch 
vier Tarsenglieder hat und von der Sackgrösse her ähnlich ist, an- 
getroffen werden. Diese Art hat jedoch lange Fühlerscheiden an der 
Brust-Kopfplatte der Puppenhülle und am Sack viele pflanzliche Teile 
wie Algen und Flechten. In der Region trifft man recht häufig die 
Säcke der Pseudobankesia alpestrella Heinemann an, sie sind auch 
grösser und weicher und oft mit weisslichen Krustenflechten belegt, 
sie haben scharfe Kanten und sind seitlich eingefallen. Alle Weibchen 
der Gattung Dahlica tragen die Afterwolle nur auf der Bauchseite, alle 
Pseudobankesia Arten hingegen haben einen Ring von Afterwolle um 
den ganzen Körper auf dem 7. Abdominalsegment. Die Männchen 
dieser Art wurden beobachtet als sie gleichzeitig mit D. wehrlii flogen. 
Die Flügelzeichnung der ? alpestrella ist viel ausgeprägter, die Flecken 
weisslich, nicht grau wie bei D. wehrlii, und erlaubt oft die Unter- 
scheidung der beiden Arten. Auffallend verschieden ist bei diesen beiden 
Arten auch der Aedeagus der männlichen Genitalapparate. Er ist stark 
gebogen bei D. wehrlii und gerade bei P alpestrella. In diesen Höhen- 
lagen ist der Einfluss des Wetters mit Rückschlägen im Frühling und 
warmen Tagen so gross, dass die Flugzeit keine Aussage über die Art- 
zugehörigkeit erlaubt. 


Verdankungen 


Ich möchte den Freunden und Kollegen Peter Sonderegger, Ruedi Bryner, 
Heinz Buser und Renato Joos ganz herzlich danken für die Hilfe beim 
Sammeln, für die Überlassung der Tierchen und für die Beratung. Den Herren 
Dr. Brancucci und Männi de Bros im Museum Basel danke ich für die Aus- 
leihung des Holotypus und Steven Whitebread für die Herstellung des Genital- 


231 


präparates. Nicht zuletzt danke ich auch meinem Freund Willi Sauter für 
die Hilfe, die Beratung und die Durchsicht des Manuskriptes und meiner Frau 
und Tochter Sereina fiir die aufopfernde Betreuung der Zuchten. 


Literatur 


GALLIKER, P., 1958. Morphologie und Systematik der präimaginalen Stadien 
der schweizerischen Solenobia-Arten. Rev. Suisse Zool. 65 : 95-183. 
MÜLLER-RUTZ, J., 1920. Aus der Welt der Kleinschmetterlinge mit Beschrei- 

bungen neuer Arten und Formen. Mitt. Ent. Zürich 5 : 334-348. 
SAUTER, W., 1956. Morphologie und Systematik der schweizerischen Sole- 
nobia-Arten. Rev. Suisse Zool. 63 : 451-559. 


282 


Nota lepid. 18 (3/4) : 233-238 ; 13.V.1996 ISSN 0342-7536 


Eine neue Montanima — Art aus dem Altai-Gebiet 
(Lepidoptera, Psychidae) 


Peter HATTENSCHWILER 
Seeblickstrasse 4, CH-8610 Uster, Schweiz 


Summary 


Montanima aurea sp. n. is described from the Altai area of the former USSR. 
The specimens were taken at light in 1983 by a group of Finnish entomologists 
collecting at 1200 m above sea level. The new species is compared with the 
type species of the genus M. karavankensis (Hôfner, 1888) and species of the 
closely related genera Bijugis Heylaerts, 1881 and Rebelia Heylaerts, 1900. 


Zusammenfassung 


Eine finnische Entomologen-Gruppe hat 1983 im Altai-Gebiet auf 1200 m ii M 
in der damaligen USSR gesammelt und beim Lichtfang eine neue Psychiden- 
Art gefunden. Diese wird als Montanima aurea sp. n. beschrieben und mit der 
Typusart M. karavankensis (Höfner, 1888) und den nahe stehenden Gattungen 
Bijugis Heylaerts, 1881 und Rebelia Heylaerts, 1900 verglichen. 


Resume 


Un groupe d’entomologistes finlandais a collectionné en 1983 dans la région 
de l’Altai (ex-URSS) a 1200 m d’alt. et découvert lors de chasses à la lampe 
une nouvelle espèces de Psychide. L’auteur la décrit et la nomme Montanima 
aurea Sp. n. Il la compare à l’espèce-type M. karavankensis (Höfner, 1888) 
et aux genres proches Bijugis Heylaerts, 1881 et Rebelia Heylaerts, 1900. 


Eine Bestimmungssendung vom „Finnish Museum of Natural History, 
Entomology Div., University of Helsinki“ in Finnland enthielt verschie- 
dene Psychiden. Darunter befanden sich nebst bekannten Arten auch 
eine Serie von 21 Männchen vom Lichtfang, die einer neuen Art an- 
gehören. Alle stammen von einer Expedition, die unter anderem auch 
ins Altai Gebirge, der damaligen Soviet Union USSR geführt hatte. 
Gesammelt wurde durch die drei Entomologen Kauri Mikkola, Heikki 
Hippa und Jukka Jalava. Letzterer hat freundlicherweise eine Beschrei- 
bung des Fanggebietes angefertigt, sie folgt im Ökologie-Teil. 


233 


Die Männchen dieser neuen Art haben golden glänzende Flügelschup- 
pen und die Behaarung des Körpers und der Beine ist auch goldgelb 
so möchte ich sie, nach dem lateinischen Wort aureus, was vergoldet 
oder goldig usw. bedeutet, Montanima aurea sp. n., nennen. Diese neue 
Art ıst der Unterfamilie Epichnopteryginae, Tribus Epichnopterygini 
zuzuordnen. 


Montanima aurea sp.n. 


Ho.totypus : Männchen 15.-19.7.1983, SW-Altai, Katun Valley, 10 km 
W. Katanda, 1200 m ü M, leg. Exp. Mikkola, Hippa, Jalava. 


PARATYPEN : 20 Männchen, gleicher Fundort : | vom 28.6.-5.7.1983, 
7 vom 6.7.-8.7.1983 und 12 vom 15-19.7.1983. 


Der Holotypus und 10 Paratypen befinden sich im Finnish Museum 
of Natural History in Helsinki, 5 Paratypen im Zoologischen Institut 
St. Petersburg und die anderen 5 Paratypen in meiner Sammlung. 


FALTERBESCHREIBUNG : Flügel-Spannweite der Männchen 16-18 mm, 
im Mittel 17 mm, Vorderflügel mit 9 Adern aus der Discoidalzelle, alle 
getrennt entspringend, nur vereinzelt r3 + r4 aus einem Punkt, mit ei- 
ngeschobener Zelle. Hinterfliigel mit 5 getrennt aus der DZ entsprin- 
genden Adern (Abb. 1). Alle Flügel einfarbig blassgelb-crème mit goldi- 
gem Glanz. Die Fliigelmembran milchig, mit feinen, goldig glanzenden 
Haarschuppen der Breitenklasse 1-2 im Vorderflügel und fast aus- 


Abb. 1. Flügelgeäder von M. aurea sp. n. 


234 


ce) 


Abb. 2. Breite-Klassen der einspitzigen (haarfôrmigen) Flügelschuppen zwischen 
Mittelzelle und Flügelapex. Auch die breiten Schuppen enden meistens in einer Spitze, 
seltener ist diese fein gezähnt (Nach Sauter und HÄTTENSCHWILER, Schlüssel zu den 
paläarktischen Psychiden-Gattungen, in Vorbereitung). 


schliesslich der Klasse 1 im Hinterflügel, schütter besetzt (Abb. 2). Die 
Flügelfransen von gleicher Farbe, im Bereich des Vorderflügel-Apex 
lang, schmal in eine Spitze auslaufend und nur ganz schwach gezähnt. 
Kopf ohne Ocellen, Augen gross, Abstand 1-1.2 mal die Augenhöhe. 
Gesicht mit langen abstehenden goldigen Haaren. Labialpalpen auf ein 
Glied mit Haarbusch reduziert. Fühler mit 19-22 Gliedern die, mit Aus- 
nahme der drei Basisglieder, Kammzähne in der Lange von etwa 2 
Geisselgliedern tragen. Die Kammzähne unbeschuppt jedoch, bewimpert 
(Abb. 3 links). Der Körper ist dunkelbraun bis schwarz, jedoch stark 
mit langen goldigen Haaren bedeckt. Die Beine ebenfalls goldgelb be- 
haart, Vorderbein mit langer Epiphyse (Abb. 3 rechts), Mittelbeine mit 
einem und Hinterbeine mit 2 Paar Tibialspornen. Das Genital etwas 
breiter als beim Gattungstypus M. karavankensis (Hôfner, 1888), das 


235 


Abb. 3. links Antenne, rechts Vorderbein mit Epiphyse von M. aurea sp. n. 


Tegumen weniger spitz zulaufend, Aedeagus stark, fast rechtwinklig 
gebogen (Abb. 4). Die Intersegmentalhäute der hinteren Abdominal- 
segmente tragen feine, nach der Kopfseite gerichtete, Dornen. Diese sind 
besonders ausgeprägt auf der Intersegmentalhaut 7/8 dorsal (Abb. 5). 
Die Dornen in diesem Feld sind sehr fein und nur mit Vergrösserungen 
ab etwa 25-fach erkennbar. 


Die Weibchen, Raupen und Säcke sind noch unbekannt. 


Abb. 4. Männliche Genitalapparate von a — M. aurea sp. n. ; b — M. karavankensis 
(Höfner). ; c — Rebelia sapho (Milliere) und d — Bijugis bombycella (Denis & Schiffer- 
müller). Der Pfeil deutet auf die Einschnürung. 


236 


opus ge Uy 
ed, Dar 


Abb. 5. Bedornung der Intersegmentalhaut 7/8 bei M. aurea sp. n., die Dornen sind 
nach der Kopfseite gerichtet. 


ÖKoLoGiE : Die Männchen fliegen am Abend und kommen zum Licht. 
Daraus ist zu schliessen, dass der späte Nachmittag oder der Abend 
auch die Paarungszeit sein dürfte. Während der Tageszeit wurden von 
den Expeditionsteilnehmern keine Falter beobachtet. Der Lebensraum 
wurde von Herrn Jukka Jalava als Steppenhang, sehr xerotherm, der 
Sonne ausgesetzt, Südhang, sehr reich an Pflanzen und Blumen be- 
schrieben (briefliche Mitteilung). Die Futterpflanzen sind nicht bekannt. 


JAHRESZEITLICHE ENTWICKLUNG: Die Flugzeit fällt in die Zeit von 
Ende Juni bis Ende Julı. 


VERBREITUNG : Über die Verbreitung kann noch nichts ausgesagt 
werden, denn alle bekannten Exemplare stammen aus der gleichen 
Gegend im Altai Gebirge. 


Diskussion 


Die Gattung Montanima Sieder, 1949 steht den Gattungen Bijugis Hey- 
laerts, 1881 und Rebelia Heylaerts, 1900 recht nahe. Bijugis hat bei 
mehreren Arten ebenfalls gelbliche Färbung, jedoch breitere Flügel und 
unterscheidet sich deutlich durch die seitlichen Einschnürungen am 
Vinculum des männlichen Genitales (Abb. 4d) und durch das Fehlen 
der dorsalen Bedornung auf der Intersegmentalhaut 7/8 (Abb. 5), die 


23) 


bei den Gattungen Rebelia und Montanima vorhanden sind. Weniger 
leicht ist die Unterscheidung der anderen beiden Gattungen. Hier ist 
der männliche Genitalapparat eine Hilfe. Das Tegumen ist bei den 
Montanima-Arten schmaler als bei den Rebelia-Arten, wobei allerdings 
zu sagen ist, dass die neue Art M. aurea schon eine Zwischenstufe 
darstellt (Abb. 4). Der Aedeagus ist bei den Montanima-Arten stark 
abgewinkelt, meist nahezu rechtwinklig, während er bei den Rebelia- 
Arten einen stumpferen Winkel aufweist. Als wichtigste Gründe für die 
Errichtung der Gattung Montanima erwähnt Sieder (1949) das schmal 
zulaufende Tegumen des männlichen Genitales und der völlıg unter- 
schiedliche Sackbau. Nachdem aber bei unseren Tieren vom Lichtfang 
die Säcke nicht bekannt sind, fehlt dieses wichtige Merkmal zur Unter- 
scheidung der Gattungen. Trotzdem scheint es beim heutigen Wissens- 
stand richtig diese neue Art der Gattung Montanima zuzuordnen. 


Verdankungen 


Den Herren Kauri Mikkola, Heikki Hippa und Jukka Jalava vom Finnischen 
Museum danke ich für die Unterstützung und die Überlassung der fünf 
Paratypen. Besonderen Dank geht an Jukka Jalava für die wertvollen Angaben 
über seine Beobachtungen und die Beschreibung des Lebensraumes. Nicht 
zuletzt möchte ich auch meinem Freund Prof. Dr. Willi Sauter für die Hilfe 
und die Durchsicht des Manuskriptes danken. 


Literatur 


SIEDER, L., 1949. Montanima gen. nov. Z. wien. ent. Ges. 34 : 2-13. 


238 


Nota lepid. 18 (3/4) : 239-246 ; 13.V.1996 ISSN 0342-7536 


Eine neue, im Februar-März fliegende Prilocephala — 
Art aus dem siidlichen Spanien (Lepidoptera, Psychidae) 


Peter HATTENSCHWILER 


Seeblickstrasse 4, CH-8610 Uster, Schweiz 


Summary 


Ptilocephala piae sp. n. is described from the coastal region of the Spanish 
province Huelva and compared with the related Ptilocephala albida Esper, 
1786. The two species are easily recognised by their very different larval cases 
and the fact that the new species flies earlier in the year, from February to 
early March. 


Zusammenfassung 


Im Februar bis Anfang Marz fliegt im Küstengebiet der Provinz Huelva im 
südlichen Spanien eine Prilocephala — Art, die als Ptilocephala piae sp. n. neu 
beschrieben und mit der nahe stehenden Prilocephala albida Esper, 1786 ver- 
glichen wird. Die beiden Arten unterscheiden sich am auffalligsten durch einen 
vollig anderen Sackbau und die frühe Flugzeit der neuen Art. 


Résumé 


De février à début mars vole dans la région côtière de la province de Huelva 
(Espagne méridionale) une Prilocephala que l’auteur décrit et nomme Ptilo- 
cephala piae sp.n. Il la compare à l’espèce proche Prilocephala albida Esper, 
1786. La différence la plus frappante entre les deux espèces est la construction 
complètement différente de leur fourreau et la période de vol précoce de la 
nouvelle espèce. 


Während eines Ferienaufenthalts Ende Januar und Anfang Februar 
1992 im Küstengebiet der Provinz Huelva in Spanien haben unsere 
beiden Tôchter Pia sowie Sereina mit ihrem Mann Marcel auch nach 
Psychiden gesucht. Besonders erfolgreich waren sie entlang der Strasse 
bei El Rocio am Rande des grossen Schutzgebietes im Miindungsgebiet 
des Rio Guadalquivir. Die Brachstreifen beiderseits der Strasse sind 
teilweise mit Wald, teilweise mit hohem Gras und stellenweise mit Moos 
bewachsen (Abb. 1). Im Moos und speziell an den dazwischen wach- 


239 


Abb. 1. Lebensraum von P piae sp. n., hinter dem Zaun im Hintergrund beginnt das 
Schutzgebiet, das vom Fundgebiet durch eine 10-20 m breite „Feuerschneise“ getrennt 
ist. Diese Schneise wird immer wieder umgepflügt und ist völlig unbewachsenen. (Foto 
S. Parpan). 


senden Gräsern fanden sie eine grössere Zahl von Säcken, die mit Gras- 
stengeln längs bekleidet waren, wobei diese die eigentliche Sacklänge 
oft weit überragten. Die Mehrheit der Säcke waren 20-40 cm über dem 
Boden an den dürren Grasstengeln festgesponnen. 


Die mitgebrachten Säcke wurden in Schachteln ausgelegt und im Zim- 
mer gehalten. Zwischen dem 4.3. und 8.3.1992 schlüpften vier Männchen 
und ab dem 10.3. bis zum 15.3.1992 dann 36 Weibchen und eine un- 
verhältnismässig hohe Zahl von Parasitoiden. Durch die zeitliche Diffe- 
renz war es nicht möglich eine Copula zu erhalten und die erhoffte 
Nachzucht konnte nicht durchgeführt werden. Das Studium der Tiere 
ergab bald den Hinweis, dass es sich um eine neue Art, Unterart oder 
Form handeln musste. In der liebenswerten Weise war Herr Bourgogne 
in Paris bereit diese Tierchen anzusehen und zu vergleichen. Er kam 
auch zum Schluss, dass es eine der Prilocephala albida Esper, 1786 
nahe stehende, neue Art oder Unterart sein müsse. 


Am 10. Januar 1993 reisten die Tochter Pia, meine Frau Ruth und 
ich nochmals nach El Rocio um Raupen der fraglichen Art zu suchen. 


240 


Zu diesem Zeitpunkt waren die Raupen noch in der Bodenvegetation 
und sehr schwer zu finden. Mehrmals besuchten wir die Fundplätze 
und es gelang uns, mehrere Raupen und einige leere Säcke zu finden 
und zogen daraus nochmals einige Männchen, Weibchen und Schlupf- 
wespen, aber leider gelang es wieder nicht eine Nachzucht zu erhalten. 
Unsere Tochter Pia war am Auffinden und Sammeln dieser Art so sehr 
beteiligt, dass ich ihr diese neue Art widmen möchte, sie soll Ptilocephala 
piae sp. n. heissen. Diese neue Art ist in der Unterfamilie Oiketicinae, 
dem Tribus Oreopsychini und in der Gattung Ptilocephala nach der 
Art albida einzuordnen. 


Ptilocephala piae sp. n. 


Von den beiden Sammelreisen standen die folgenden Exemplare für die Ver- 
gleiche zur Verfügung : 


1992 : 4 Männchen, 36 Weibchen, alle ex. pupa 4-15. März, 3 Raupen und 
12 weitere Säcke, leg. Pia Hättenschwiler sowie Sereina und Marcel Parpan. 


1993 : 7 Männchen, 6 Weibchen, alle ex. larva 18. Februar — 7. März, 1 Raupe 
und 7 weitere Säcke gesammelt durch Tochter Pia, meine Frau Ruth und mir 
selber. Aus beiden Jahren zusätzlich noch die Säcke der parasitierten Raupen. 


Hoıortypus : @, El Rocio, Provinz Huelva, Spanien, ex. |. 22.2.1993. 
ALLOTYPUS : ®, El Rocio, Provinz Huelva, Spanien, ex. I. 10.-15.2.1992. 


Alle anderen Exemplare sind als Paratypus bezeichnet. Die Holotypus 
und Allotypus sind in meiner Sammlung. Von den Paratypen sind je 
ein Pärchen im Museo Nacional de Ciencias Naturales, Madrid und 
im Natural History Museum, London, die restlichen Exemplare be- 
finden sich in meiner Sammlung. 


MANNCHEN : Flügel-Spannweite 15-17 mm, im Mittel 16 mm. Alle Flü- 
gel glasklar mit weit auseinander stehenden feinen, kurzen Haaren, 
Adern dunkel, gut sichtbar, Ränder fast schwarz, am äusseren Rand 
aus kurzen, tief eingeschnittenen, meist 2-zackigen Schuppen bestehend. 
Vorderflügel breit, Apex gerundet mit 7, Hinterflügel mit 4 Adern aus 
der Discoidalzelle (Abb. 2), deutlich kürzer als Vorderflügel stark ge- 
rundet. Fühler doppelkammzähnig, nur gut einen Drittel der Flügel- 
länge erreichend, Kammzähne sehr lang, bewimpert mit vereinzelten 
weisslichen Haaren und Schuppen besetzt. Geisselglieder an der Basis 
kürzer als deren Durchmesser, gegen die Fühlerspitze sind sie 3-4 mal 
so lang wie der Durchmesser, 28-30 Glieder (Abb. 3). Ocellen fehlen, 
Labialpalpen kaum noch erkennbar, Gesicht und Körper lang, schwarz 
behaart. Augen klein, hochoval, Abstand etwa zwei mal Augenhöhe. 
Beine ohne Epiphysen und Sporne, schwarz, lang behaart, 5 Tarsen- 


241 


SN 
I 


Abb. 2. Flügelform und Geäder von P piae sp. n. 


Abb. 3. Fühler der Männchen von P piae sp. n. Nur wenige Kammzähne in ganzer 
Länge gezeichnet. 


glieder, das erste Glied viel länger als die anderen vier. Genital : Länge 
zu Breite = 3.2-3.8, Mittel 3.51. n — 5 und damit schmaler als bei ? 
albida, mit langem, feinem Saccus, er ist länger als der restliche Teil 
des Genitalapparates (Abb. 4). Die Sternite und Tergite aller Abdo- 
minalsegmente sind hart sklerotisiert, dunkelbraun, die Sternite sind 
in der Mitte durch einen breiten Spalt längs geteilt, was bei P albida 
oft weniger ausgeprägt ist. Bedornung auf der Intersegmentalhaut 7/8 
nur ventral. 


242 


Abb. 4. Männliche Genitalapparate. a — P piae sp. n. ; b — P albida Esper. 
c = Messpunkte für das Lange : Breite Verhältnis, andere Unterschiede sind zufällig. 


WEIBCHEN : Sehr stark reduziert, ungeflügelt, Beine noch als kurze 
Stummel erkennbar, Augen auf dunkle Flecken reduziert. Körper 5.5- 
7 mm lang, 2.5-3.5 mm Durchmesser, gegen hinten etwas dicker 
werdend, Kopf und Brustsegmente dorsal dunkelbraun, stark sklero- 
tisiert, Hinterkörper weisslich, etwas cr&mefarbig getönt, die Segmente 
tragen je einen Kranz von weisslicher, leicht bräunlich getönter After- 
wolle, der auf Segment 7 am stärksten ist und gegen den Kopf zu 
schwächer wird. Selten kann man alle sechs Kränze noch erkennen. 
Der Legeapparat besteht äusserlich aus einem kurzen, fleischigen Wulst. 


RAUPEN : 9-11 mm lang und 1.5-2 mm Durchmesser. Körper gelb- 
bräunlich mit vielen Platten von etwas dunklerer, bräunlicher Farbe. 
Kopf und die drei Brustglieder hart sklerotisiert mit dunklen Flecken, 
die zusammen 6-8 Längsreihen oder -Streifen bilden. Die letzten zwei 
Segmente ebenfalls mit harter, dunkler Rückenplatte. 


Die Säcke beider Geschlechter (Abb. 5) sind mit Grashalmen längs 
belegt, beim Männchen 11-14 mm lang, 3.5-4.5 mm Durchmesser, oft 
mit vereinzelten Halmen, die bis 23-25 mm lang sein können, End- 
röhre kurz, grau. Weibchen Säcke 11-13 mm lang und 3.5-5 mm Durch- 


messer mit vielen überragenden Halmen von einer Gesamtlänge bis 
zu 20-25 mm. 


243 


mm 
25 


0 


Abb. 5. Säcke von P piae sp. n. a — Männchen ; b — Weibchen ; c, d — verschiedene 
Erscheinungsformen der Säcke von P albida Esper. 


Puppen : Kopf-Brustplatte beider Geschlechter mit vier Borstenpaaren, 
die Platte der weiblichen Puppen ist sehr stark rückgebildet, die ein- 
zelnen Teile sind kaum noch erkennbar. Die Rückenplatten der Hinter- 
leibssegmente mit einer nach hinten gerichteten Dornenreihe, auf der 
Intersegmentalhaut eine Reihe, deren Dornen nach vorn gerichtet sind. 


ÖKOLOoGIE : Die ersten Männchen schlüpften am 18. Februar, die letz- 
ten am 7. März ab etwa 7 Uhr am Morgen, was in dieser Jahreszeit 
noch in der Dunkelheit ist, die letzten gegen 9 Uhr. Nach 15-20 Minuten 
sind sie schon flugbereit. Die Weibchen schlüpfen fast gleichzeitig, sie 
verlassen aber die Puppe nicht, sie sprengen nur die Rückenplatte der 
Brustsegmente der Puppe und trennen die Kopf-Brustplatte ab. Das 
Pheromon der paarungsbereiten Weibchen scheint durch den Sack zu 
dringen und stimuliert die wartenden Männchen, die dann ohne auf 
Gefahren zu achten in stürmischem Flug die Weibchen aufsuchen. Die 
Eier werden, wohl wie bei P albida beobachtet, in die Puppenhülle ab- 
gelegt wobei die Afterwolle des Weibchens abgerieben wird und zwi- 
schen den Eiern als Polsterung oder Isolation dient. Nach dem Ablegen 
der Eier ist das Weibchen auf ein Häufchen Haut reduziert und stirbt 
noch in der Puppenhülle. Die Paarung wurde nicht beobachtet, dürfte 
jedoch gleich ablaufen wie bei den anderen nahestehenden Arten in 
der Gattung, also im Sack in der Puppenhülle. Das Männchen kann 
dazu den Körper sehr weit ausdehnen und so durch die Sacköffnung, 
die die Raupe speziell dazu vorbereitet hat, in die Puppenhülle ein- 
dringen und die Geschlechtsöffnung des Weibchens erreichen. 


244 


Die Raupen leben von Gräsern, verschiedenen Kräutern und Moosen. 
Um den Sack der wachsenden Raupe anzupassen muss dieser der Länge 
nach aufgebissen und ein längerer Halm neu eingesetzt werden. Die 
Erweiterung geschieht also in Querrichtung. Die Grashalme werden auf 
die Länge des Sackes oder länger zurechtgebissen und erst dann einge- 
arbeitet. Die Entwicklung dauert ein Jahr, ob eine Diapause oder Ruhe- 
zeit eingehalten wird konnte nicht beobachtet werden, es muss aber 
angenommen werden, dass in den Spätsommer- oder Herbstmonaten 
eine Ruhe stattfindet. 


Die Raupen werden sehr stark von Parasitoiden befallen. Von den etwa 
100 Säcken, alle waren angesponnen, die 1992 gefunden wurden, waren 
nur 40 geschlüpft, der Rest war parasitiert durch verschiedene Wespen, 
darunter die zwei Arten mit ungefliigelten Weibchen Gelis fortificator 
Aubert, 1980 und Gelis meridionator Aubert, 1960 die durch Herrn 
Mag. Martin Schwarz, Zool. Inst. der Univ. Salzburg freundlicher- 
weise bestimmt wurden. Es ist allerdings môglich, dass die parasitier- 
ten Raupen sich anders verhalten und sie dadurch leichter gefunden 
wurden. 


Die Verpuppung der Männchen erfolgt in oder in der Nahe der Boden- 
vegetation, die Weibchen steigen 20-40 cm an den dürren Grashalmen 
empor und spinnen den Sack dort erhoht fest. dadurch kann der Lock- 
duft sich besser ausbreiten. Gegen Ende Januar oder in den ersten 
Februartagen werden die Säcke angesponnen, die Puppenruhe dauert 
etwa drei Wochen. 


Die neue Art wurde bisher nur entlang der Strasse in der Gegend von 
El Rocio in der spanischen Provinz Huelva gefunden. Der Lebensraum 
ist meistens trocken und stark besonnt. Die Vegetation besteht aus 
den üblichen niederen Pflanzen, besonders aber aus hartem, hohem 
Gras, das ab und zu von steinigen, moosbewachsenen Stellen unter- 
brochen ist. Bäume, meist Föhren, sind nur vereinzelt vorhanden. Das 
Fundgebiet liegt nur wenige Meter über dem Meeresspiegel. 


Diskussion 


Die neue Art steht der sehr weit verbreiteten Prilocephala albida nahe. 
Diese besiedelt ähnliche Lebensräume, wurde aber in der Umgebung 
von El Rocio, soweit mir bekannt, nicht gefunden. Es sind eine Reihe 
von Unterschieden vorhanden (Tab. 1), die auffälligsten sind sicher die 
Säcke und die Flugzeit. P albida fliegt im Mai und Juni und baut 
Säcke, die wirr mit Moos oder verschiedenen Blättern und Pflanzen- 
fragmenten belegt sind. P piae fliegt im Februar bis Anfang März und 


245 


baut Säcke die mit parallelen Grashalmen längs belegt sind (Abb. 5). 
Gewisse Unterschiede sind auch in den männlichen Genitalien zu finden. 
So ist das Verhältnis von Lange: Breite der männlichen Genitalien 
unterschiedlich. Dieses ist bei P piae 3.2-3.8, Mittel 3.51, bei P albida 
2.7-3.5, Mittel 3.07. 


Tab. 1. Gegenüberstellung einiger Merkmale der beiden Arten P piae sp. n. und P. al- 


bida Esper. 


Männliches Genital 
Verhältnis Lange : Breite 
Mittelwert 
Anzahl Präparate 


Flugzeit 
Sackform 


Anordnung der 
„Sackbekleidung“ 


Sack „Bekleidung“ 


Sack Durchmesse über 
„Bekleidung“ gemessen 


Endröhre der &-Säcke 


Ptilocephala piae 


3,2-3,8 
si 
5 


Februar-März 

länglich 

längs 

Grashalme, gut anliegend, 
parallel 

3,5-4,5 mm 


0,5-1,5 mm lang 


Ptilocephala albida 


2,7-3,5 
3,07 
14 


Mai-Juni 

rundlich 

Witr 

Pflanzenfragmente oder 
Moos, seitlich abstehend 
5-10 mm 


2-3 mm lang 


Verdankungen 


Ich möchte Herrn Jean Bourgogne ganz herzlich danken für seine Vergleiche 
mit den Belegen von P albida im Museum Paris und für die Übergabe all 
seiner Erfahrungen und Notizen über die in Frage stehende Gattung, ebenso 
geht mein Dank an meinen Freund Prof. Dr. Willi Sauter fiir die Beratung 
und das Prüfen des Manuskriptes und Herrn Mag. Martin Schwarz für das 
Bestimmen der Parasitoiden. Ein ganz besonderer Dank geht an Pia sowie 
Sereina und Marcel Parpan und meiner Frau Ruth fiir das fleissige und 
erfolgreiche Sammeln der interessanten Tierchen. 


Literatur 


BOURGOGNE, J., 1967. Materiaux pour une révision du genre Oreopsyche. 
Alexanor V : 1-40. 


246 


Nota lepid. 18 (3/4) : 247-265 ; 13.V.1996 ISSN 0342-7536 


Vergleichende Analyse der Zönosen tagaktiver 
Schmetterlinge im Sengsengebirge (Oberösterreich) 
(Lepidoptera) (*) 


Erwin HAUSER 


Forschungsgemeinschaft Wilhelminenberg, Otto Koenig-Institut Staning, Dorf a.d. Enns 69a, A-4431 
Haidershofen, Osterreich 


Summary 


Habitats and communities of butterflies and moths of the mountain-range 
“Sengsengebirge” (Upper-Austria) were studied in 1992 and 1993 by daytime 
counting in the area of the mountain “Hoher Nock”. The cluster-analysis 
of the communities produced four groups, which mainly depended on altitude 
and the location on the south or north side of the mountain-range, but not 
in all cases on the type of vegetation. A number of different communities 
and several endangered species have been observed in the investigated area. 
“Especially valuable” were south-facing grassland on rocky ground, woodland 
with sunny clearings, alpine meadows high up on the “Hoher Nock” and open 
vegetation of herbaceous plants on the northern side of the mountain-range. 
The extensively managed pasture “Misteleben” has been classified as “valuable”. 
Only three of the twenty high altitude species known from Upper Austria 
have been recorded in the Sengsengebirge (Boloria pales, Psodos alpinata and 
Psodos quadrifaria). This small number is assumed to be a result of warm 
inter- or postglacial periods. 


Zusammenfassung 


Lebensräume und -gemeinschaften tagaktiver Groß-Schmetterlinge in der 
montanen bis alpinen Stufe des Sengsengebirges sind in den Jahren 1992 und 
1993 beschrieben, verglichen und aus der Sicht des Naturschutzes bewertet 
worden. Eine Clusteranalyse der Falterzönosen auf der Basis korrigierter 
Jaccard-Indices ergab vier große Gruppen, die hauptsächlich durch ihre 
Höhenstufe sowie der Lage auf der Süd- oder der Nordflanke des Gebirges, 
nicht unbedingt aber durch den Vegetationstyp bestimmt waren. Für das unter- 
suchte Gebiet um den Hohen Nock konnte eine Vielfalt an Falterzönosen 
mit etlichen gefährdeten Arten belegt werden. „Besonders wertvoll“ waren süd- 
exponierte, flachgründige montane bis subalpine Gras- und Felsfluren, Wälder 


(*) Im Auftrag der Nationalparkplanung Kalkalpen, Leonstein (O.O). Geféidert durch 
das österreichische Bundesministerium für Umwelt, Jugend und Familie. 


247 


mit hohem Anteil an sonnenexponierten Lichtungen, die Matten der Gipfel- 
region und Hochstaudenfluren an der Nordflanke. Die untermontane und 
extensiv genutzte ,,Mistelebenweide“ wurde bezüglich ihrer Zönose tagaktiver 
Schmetterlinge als „wertvoll“ eingestuft. Von den etwa 20 in Oberösterreich 
vorkommenden tagaktiven Groß-Schmetterlingen, die ausschließlich die höhere 
Gebirgsstufe besiedeln (subalpin, alpin ; nur ausnahmsweise obermontan), sind 
nur drei Arten vertreten : Boloria pales, Psodos alpinata und Psodos quadri- 
faria. Der geringe Anteil an tagaktiven Hochgebirgsarten im Sengsengebirge 
ist vermutlich auf zwischen- oder nacheiszeitliche Wärmeperioden zurückzu- 
führen. 


Resume 


Les biotopes et les biocénoses des Macrolépidoptères à vol diurne des étages 
montagnard à alpin dans le massif de Sengsen (Haute-Autriche) ont été décrits 
en 1992 et 1993, comparés et qualifiés sur le plan de la protection de la nature. 
Une analyse de groupe (cluster) des bioc&noses de lépidoptères sur la base 
des indices Jaccard corriges a mis en Evidence quatre grands groupes deter- 
minés essentiellement par leur étage (altitude), ainsi que par leur place sur 
les flancs sud ou nord de la montagne, mais pas de façon absolue par le 
type de végétation. Pour la zone étudiée autour du Hoher Nock, on a pu 
mettre en évidence un grand nombre de biocénoses de lépidoptères avec beau- 
coup d’espèces menacées. Les zones herbeuses et rocheuses montagnardes a 
subalpines à fond plat exposées au sud ont été classées «particulièrement 
précieuses», ainsi que les forêts avec beaucoup de clairières ensoleillées, les 
pelouses de la région sommitale et les mégaphorbiées sur le flanc nord. Le 
pâturage «Misteleben» bas-montagnard en exploitation extensive a été classé 
«Précieux» en ce qui concerne sa biocénose de papillons volant de jour. Parmi 
les quelque 20 espèces de Macrolépidoptères diurnes de Haute-Autriche qui 
vivent exclusivement dans la partie supérieure de la montagne (étages subalpin, 
alpin : exceptionnellement montagnard supérieur), on ne trouve dans la litté- 
rature et selon les propres observations de l’auteur que trois espèces : Boloria 
pales, Psodos alpinata et Psodos quadrifaria. Le petit nombre d’espéces de 
haute montagne volant de jour dans le massif de Sengsen pourrait être un 
effet des périodes chaudes inter- et post-glaciaires. 


Einleitung 


Das Sengsengebirge ist Bestandteil der oberösterreichischen Kalkalpen 
und liegt zwischen dem hochalpinen Toten Gebirge im Westen und 
dem mittelgebirgsartigen Reichraminger Hintergebirge im Osten. Nörd- 
lich und südlich des Sengsengebirges erstrecken sich regionale Becken- 
landschaften (Mollner und Windischgarstener Becken). Die Achse der 
Gebirgskette liegt etwa in ostwestlicher Richtung, sodaß an der Nord- 
und der Südflanke vor allem durch differente Sonnenexposition und 


248 


Niederschlagsverhältnisse (Nordstaulagen) unterschiedliche Vegetations- 
typen auftreten. Das Sengsengebirge reicht mit seinen Gipfeln kaum 
über die subalpine Stufe hinaus, in den obersten Höhenstufen sind 
kurzrasige Matten mit Latschenbeständen (Legfôhren, Pinus mugo) ge- 
mischt. Seine hôchste Erhebung ist der Hohe Nock mit 1960 m See- 
hohe. 


In den Jahren 1992 und 1993 sind Untersuchungen an Schmetterlingen 
durchgefiihrt worden. Alle Falterdaten und die Ergebnisse der Nacht- 
falterzählungen legen bereits in publizierter Form vor (Hauser, 1995). 
Ziel war nicht nur eine faunistische Erhebung, sondern vor allem Be- 
schreibung und Vergleich der montanen bis alpinen Lebensräume tag- 
aktiver Schmetterlinge (Höhenstufen nach Adler er al., 1994). Als Unter- 
suchungsgebiet wurde ein Areal südlich und nördlich des Hohen Nock- 
Gipfels ausgewählt (Abb. 1). Zusätzlich sollten die Artengemeinschaften 
der an die höchstgelegenen Lebensräume angepaßten Tagschmetterlinge 
zwischen dem Sengsengebirge und den übrigen oberösterreichischen 
Kalkgebirgen verglichen werden. 


Im Hinblick auf die praxisbezogenen Planungsarbeiten für den National- 
park Kalkalpen wurde auf eine naturschutzrelevante Beurteilung be- 
sonderer Wert gelegt. Das Sengsengebirge ist nach dem oberösterreichi- 
schen Naturschutzgesetz als Naturschutzgebiet ausgewiesen, das Gebiet 
soll in den geplanten Nationalpark Kalkalpen eingegliedert werden. 


Methoden 


Die Arten sind im Freiland nach Higgins & Riley (1971) bestimmt 
worden (Netzfang oder Beobachtung mit dem Fernglas). Bei scheuen 
oder flüchtenden Tieren konnte manchmal nur die Gattung determiniert 
werden. Die Männchen der Weißlinge Pieris napi und Pieris bryoniae 
sind nicht sicher zu unterscheiden, daher wurden nur Nachweise von 
Weibchen den entsprechenden Arten zugeordnet. Untersuchungsgegen- 
stand waren alle tagaktiven Groß-Schmetterlinge, also jene Falter, die 
bei einer Begehung am Tag zu erwarten sind. Die Nomenklatur folgt 
Forster & Wohlfahrt (1960 bis 1981). 


Die Freilandarbeiten fanden am späten Vormittag bis zum frühen 
Nachmittag stets bei „Schönwetter“ statt (1992 war ein Jahr mit extrem 
heißem Sommer). Hohe Temperaturen, geringe Bewölkung und Wind- 
geschwindigkeiten waren bei der Arbeit in den oberen Höhenstufen 
von besonderer Bedeutung, weil bekannt ist, daß die dort verbreiteten 
Tagfalter ihre Aktivität mit zunehmenden Wind oder starker Bewölkung 
einstellen. Die Hauptarbeiten wurden im Jahr 1992 vom Mai bis Sep- 


249 


Feichtau 
Misteleben 


ne 


Niklbach 


0 1000 m 
ee EEE | 


[___] Grasfluren 
Felsfluren 
Hochstaudenfluren 
Latschenbestande 
ll] Nadelholzdominierte Walder 
BEE Laubholzdominierte Wälder 


Hinterer Rettenbach 


Abb. 1: Lage der Untersuchungsflächen im Gebiet um den Hohen Nock (620 bis 
1960 m.ü.M.). 


250 


tember durchgeführt (eine Zählung pro Monat, im Juli zwei). Zusätz- 
liche Begehungen im Sommer 1993 dienten lediglich dem Nachweis von 
Arten, die zwei Jahre zur Entwicklung benôtigen. 


Standortstruktur und Vegetation der 27 ,,Habitate“ (Abb. 1) sind aus- 
führlich bei Hauser (1995) wiedergegeben. Flächen ähnlicher Vegetation 
und Lage (Meereshöhe, Hangexposition) wurden in Anlehnung an 
Lepidopterologen-Arbeitsgruppe (1991) und Blab & Kudrna (1982) zu 
15 ,,Habitattypen“ zusammengefaßt und erst in dieser Form für den 
Vergleich ihrer Faltergesellschaften (-zônosen) verwendet. Die Habitat- 
typen fassen ein bis maximal vier Habitate zusammen. 


Die Falterzönosen der „Habitatypen“ wurden mit Hilfe einer Cluster- 
analyse auf der Basis korrigierter Jaccard-Zahlen (EJa) gruppiert und 
gemäß ihrer Ähnlichkeit in einem hierarchisch organisierten Dendro- 
gramm zusammengestellt. 


Die Artenidentität zweier Zönosen stellt den Anteil gemeinsamer Arten 
in Prozent dar (Jaccard-Zahl, Ja) (Mühlenberg, 1989 ; Balogh, 1958). 
Aufschlußreich ist auch die maximal erreichbare Artenidentität (Ja .x)» 
ein Maß für die Verschiedenheit der Artenzahlen, das bei gleicher 
Artenzahl in den Habitaten den Wert 100% erreicht. Zusätzlich wurde 
die Zahl E,, berechnet, die die Jaccard-Zahl durch die maximal er- 
reichbare Artenidentität relativiert (E,, = Ja/Ja,,,,). Er, gibt folglich 
an, ob sich die Artenspektren zweier Zönosen ähneln (E,, nahe | bei 
gleichzeitig hohem Ja), ob eine Zönose eine ,,verarmte“ Ausprägung 
der anderen ist (E,, nahe 1 bei gleichzeitig niedrigen Ja) oder ob sich 
die Verschiedenheit auf exklusive Arten in beiden Zönosen gründet 
(E,, nahe Null). Die Clusteranalyse wurde mittels der „single linkage“- 
Methode auf Basis einer ungewichteten Distanzmatrix (1- E,,) durch- 
geführt. Dieses Verfahren reagiert empfindlich auf Ausreißer und eignet 
sich gut zum Erkennen natürlicher Gruppen (Mühlenberg 1989 ; Back- 
haus et al. 1990). In der Clusteranalyse wurden alle Arten berücksichtigt, 
sowohl euzöne als auch zönosefremde Arten (s. unten). Dies deshalb, 
weil die Falter — auch wenn sie sich nicht im Habitat vermehren — 
ein ausgewähltes Angebot an Strukturen und Blüten nutzen sowie das 
charakteristische Topoklima bestimmter Habitattypen bevorzugen. So- 
gar die Wanderfalter und die gemeinhin als Ubiquisten geltenden Arten 
sind in diesem Sinn für eine Untersuchung von Falterzönosen bedeut- 
sam, was sich vor allem im Vergleich zwischen Offenland und Wald 
äußerte. 


Als „euzön“ wird eine Art dann bezeichnet, wenn sie eine deutliche 
Präferenz für einen bestimmten Habitattyp aufweist und darin als 
zönoseeigen gelten kann (d.h. sie vermehrt sich im Habitattyp und hält 


a! 


hier ihren Bestand auch ohne Zuzug von außen aufrecht) (Schwerdt- 
feger 1975 ; Erhardt 1985). Eine Schmetterlingsart kann in einem Habi- 
tattyp euzön, in einem anderen zönosefremd sein. Boloria pales und 
Psodos alpinata waren z.B. im Bereich der alpinen und subalpinen 
Matten euzön, auf den angrenzenden Fels- und Schuttflächen hingegen 
zönosefremd. Ähnlich war es beim Apollofalter, der für montane Fels- 
fluren als euzön gelten kann und als zönosefremdes Element in den 
angrenzenden Wälder auf Lichtungen angetroffen wird. 


Die Gefährdungskategorien für Oberösterreich lauten nach der Roten 
Liste (Bundesministerium für Umwelt, Jugend und Familie, 1994) : 
Kat. 0 : ausgestorben, ausgerottet oder verschollen ; Kat. 1 : vom Aus- 
sterben bedroht ; Kat. 2: stark gefährdet ; Kat. 3 : gefährdet ; Kat. 4: 
potentiell gefährdet ; Kat. 7: nicht bodenständige Weitwanderer mit 
starkem Rückgang ; ? : ungenügend erforscht ; + : nicht gefährdet. 


Ergebnisse 


Die Habitattypen 


Der Anschaulichkeit halber sind die 15 Habitattypen nach ihren Vege- 
tationsformen geordnet (Hochstaudenfluren : Hl bis H2 ; Grasfluren : 


40- 


CO] Summe 
EE euzön 
gefährdet 


30) 


20 


vll 


WANN RIRES I Ze BIRB2BS 


Anzahl der Arten 


Hochstau- Grasfluren Felsfluren Latschen Wald 
denfluren 


Abb. 2: Vergleich der Artenzahlen tagaktiver Schmetterlinge (Summe, euzöne Arten 
und in Oberösterreich gefährdete Arten) zwischen den Habitattypen. 


252 


WI bis W3; Felsfluren : Fl bis F3 ; Latschenbestände : L1 bis LA; 
Walder: Bl bis B3) (Tab. 1). Sie unterscheiden sich voneinander 
erheblich in der Artenzahl sowie der Menge an euzônen und gefährdeten 
Arten (Tab. 2, Abb. 2). 


Tabelle | 


Habitattypen im Untersuchungsgebiet. 
Gefährdungskategorie für Oberösterreich in Klammern. Vergleiche Abb. 1. 


Hochstaudenflur H1 : Ruderalflora/ Waldschlag/ Forststraße (obermontan, mäßıg son- 
nenexponiert, oberhalb Blumaualm) 

Charakterisierung : wiesenartiger Randstreifen einer Forststraße (Berg-Reitgras), 
rundum ein verbuschter Waldschlag ; Straßenböschung felsig, 1170 m.ü.M 

Habitate : Nr. 100 

Gefährdete Arten : Erebia oeme (4) 

Euzöne Art : Erebia pronoe : bei felsiger Straßenböschung 

Artenzahl : 20 (1 euzön, 1 gefährdet) 


Hochstaudenflur H2 : Hochstaudenfluren der Nordflanke (obermontan, Herzerlsee) 
Charakterisierung : bodenfeuchter, meist steiler Osthang, der groBteils mit Hochstau- 
den bewachsen ist (Senecio fuchsii, Veratrum album, Trollius europaeus, Aconitum 
napellus, Urtica dioica, Geranium sylvaticum) ; eingestreut sind spärlich bewachsene 
Schuttkegel mit Latschen und randlichen Grasfluren ; im untersten Bereich Matten 
von Luzula ; selten von Rindern beweidet ; Umland Fichten/ Lärchen/ Rotbuchenwald ; 
1250-1450 m.ü.M. 

Habitate : Nr. 102a, 102b 

Gefährdete Arten : Clossiana thore (2) ; Aricia artaxerxes allous (3) 

Artenzahl : 34 (7 euzön, 2 gefährdet) 


Grasflur W1 : untermontane Almweide (mäßig sonnenexponiert, Misteleben) 
Charakterisierung : als extensive Rinderweide genutzte Alm, zentral zwei alte Roßkasta- 
nienbäume ; auf der südexponierten Teilfläche ein geringflächiger Quellsumpf ; Umland : 
rotbuchendominierter Laubmischwald (einzelne Fichten) ; 710-750 m.ü.M. 

Habitate : Nr. 121 

Gefährdete Arten : keine 

Artenzahl : 32 (3 euzön) 


Grasflur W2 : Gipfelplateau Hohe Nock (alpine/subalpine Matten) 

Charakterisierung : dolinenreiche, im gesamten schwach nach Nordwesten geneigte 
Fläche ; der größte Teil von lockerem Latschenbestand bedeckt, dazwischen kurzrasige 
oder hochrasige (in Gräben) Matten ; im oberen Bereich keine Latschen, nur kurzrasige 
Matten : Dryas octopetala, Gentianella aspera, Helianthemum, Carex firma ; spärlich 
bewachsene Felsflur im Bereich der Dolinen : Saxifraga aizoides ; unbewirtschaftet ; 
1850-1960 m.ü.M. 

Habitate : Nr. 107 

Gefährdete Arten : Psodos alpinata (2) 

Artenzahl : 9 (2 euzön, 1 gefährdet) 


Grasflur W3 : montane Grasflur auf flachgründigem Boden (Südflanke, z.T. verbuscht) 
Charakterisierung : südostexponierter Pfeifengrasbestand, zum Teil mit Sorbus aria 
und Amelanchier ovalis verbuscht ; Umland : Rotbuchen (-Fichten)-Wald ; unbewirt- 
schaftet ; 750-890 m.ü.M. 

Haupthabitate : Nr. 1, 2a 

Gefährdete Arten : Erebia oeme (4) ; Strymonidia spini (4) 

Artenzahl : 20 (1 euzön, 2 gefährdet) 


DS 


Felsflur F1 : südexponierte montane Magerrasen (Felsflur) 

Charakterisierung : steile, nach Südosten geneigte Felshänge mit lückriger Trocken- 
vegetation (Globularia, Teucrium, Erica, Molinia, Buphthalmum, Vincetoxicum, Thy- 
mus), geringgradig verbuscht (Sorbus aria, Amelanchier ovalis) ; unbewirtschaftet ; 890- 
1030 m.ü.M. 

Habitate : Nr. 2b, 3a, 3b 

Gefahrdete Arten : Erebia oeme (4) ; Strymonidia spini (4) 

Artenzahl : 38 (9 euzön, 2 gefährdet) 


Felsflur F2 : subalpine/ alpine Felsfluren (südexponiert, Südflanke) 

Charakterisierung : steile südexponierte Felsflur und Schutt (spärliche Grasflur, Scabiosa 
lucida, Carduus defloratus) ; unbewirtschaftet ; 1600-1960 m.ü.M. 

Habitate : Nr. 13a, 14c 

Gefährdete Arten : Erebia oeme (4) ; Psodos alpinata (2) 

Artenzahl : 13 (3 euzôn, 2 gefahrdet) 


Felsflur F3 : subalpine Felsfluren (Nordflanke) 

Charakterisierung : spärlich bewachsene Schutthalden, steil nach Nord geneigt ; einzelne 
Latschen, Papaver alpinus, Pedicularis, Linaria alpina ; in Teilbereichen (Nr. 105) kurz- 
rasige Matten mit Zwergsträuchern (Rhododendron hirsutum, Scabiosa lucida, Parnas- 
sia palustris) ; unbewirtschaftet ; 1570-1830 m.ü.M. 

Habitate : Nr. 104, 105, 106 

Gefährdete Arten : Psodos alpinata (2) 

Artenzahl : 4 (1 euzön, 1 gefährdet) 

Die Habitate sind bezüglich der Schmetterlingsfauna extrem arten- und individuenarm. 
Psodos alpinata nur in den Habitaten mit Matten. 


Latschenbestand L1 : montane südexponierte Latschenflächen (Teilhabitate : Mager- 
rasen, mesophile Wiesenflecken) 

Charakterisierung : dichter Latschenbestand mit Lichtungen aus steinigem, xerother- 
mophilem Magerrasen (Globularia) ; Umland : lockerer Fichten-Lärchen-Wald ; 1130- 
1170 m.ü.M. 

Habitate : Nr. 5 

Gefährdete Arten : keine 

Artenzahl : 14 (1 euzön) 


Latschenbestand L2 : montane/untersubalpine Latschenfläche (Südflanke ; Teilhabitate : 
Zwergsträucher) 

Charakterisierung : dichter südexponierter Latschenbestand ; Primula elatior, Viola 
biflora, Daphne mezereum, Rhododendron hirsutum ; 1450-1520 m.ü.M. 

Habitate : Nr. 10 

Gefährdete Arten : keine 

Artenzahl : 12 (keine euzönen Arten) 

Boloria pales wahrscheinlich zönosefremd. 


Latschenbestand L3 : obersubalpines Latschenfeld (südexponiert ; Teilbiotope : oft 
großflächige hochgrasige Matten) 

Charakterisierung : abwechselnd dichte Latschenbestände und hochgrasige Matten, z.T. 
steil nach Süden geneigt ; in Teilbereichen mit kleinflächigen Felsfluren ; Scabiosa 
lucida, Gentianella aspera, Carduus defloratus ; 1780-1920 m.ü.M. 

Habitate : Nr. 12 

Gefährdete Arten : Clossiana titania cypris (4) ; Psodos alpinata (2.2) 

Artenzahl : 10 (3 euzön, 2 gefährdet) 


Latschenbestand L4 : nordexponierter subalpiner Latschenbestand (Teilhabitate : Nr. 103 
fast ausschließlich Latschen, Nr. 11 mit Zwergsträuchern) 

Habitate : Nr. 103, 11 

Gefahrdete Arten : keine 

Artenzahl : 11 (keine euzônen Arten) 

Boloria pales wahrscheinlich zönosefremd. 


254 


Wald B1 : montan/subalpiner lichter Lärchen-Fichten-Wald, Südflanke 
Charakterisierung : lockerer, südexpomierter Lärchen-Fichten-Wald (Nr. 9c), häufige 
Waldlichtungen mit z.T. anstehendem Fels (Nr. 9a) mit Helleborus niger, Adenostyles, 
Erica, Trollius ; am westlichsten Punkt des Weges eine wenige Quadratmeter große 
nasse Stelle bei 1360 m.ü.M. (zu Nr. 9a); bei ca. 1400 m eine kurzrasige Lichtung 
(Nr. 9b) mit geringer Neigung nach Süden ; 1300-1450 m.ü.M. Wegen der engen Ver- 
zahnung der Teilflächen erscheint die Behandlung als einen Habitattyp gerechtfertigt. 
Habitate : Nr. 9 mit den Teilflächen a, b und c 

Gefährdete Arten : Aricia artaxerxes allous (3) ; Erebia oeme (4) ; Melasina lugubris (1) 
Euzöne Arten der Habitatteile : Aricia artaxerxes allous : 9b ; Boloria pales : 9b (in 
9a zönosefremd ?) ; Lysandra coridon : 9a ; Melasina lugubris : 9a 

Artenzahl : 29 (4 euzön, 3 gefährdet) 


Wald B2 : fichtendominierte, montane, lichte Wälder (Südflanke) 

Charakterisierung : lockere, nach Süden oder Südosten geneigte Fichtenwälder ; Lich- 
tungen mit einem Mosaik aus Hochstauden und Zwergsträuchern (Aconitum napellus, 
Helleborus niger, Vincetoxicum hirundinaria, Adenostyles, Buphthalmum, Pteridium, 
Erica, Vaccinium myrtillus, im Wald Helleborus niger) ; 900-1300 m.ü.M. 

Habitate : Nr. 2c, 4, 7, 8 

Gefährdete Arten : Erebia oeme (4) ; Lopinga achine (4) 

Euzöne Arten der Habitatteile : Erebia ligea : v.a. im Waldesinneren ; Lysandra coridon : 
xerotherme Stellen 

Artenzahl : 28 (2 euzön, 2 gefährdet) 


Wald B3 : montane geschlossene Rotbuchenbestände (Süd- und Nordflanke) 
Charakterisierung : Rotbuchenwälder mit geringem Fichtenanteil auf der Nordflanke 
(Nr. 101, nach Nordost geneigt) oder Südflanke (südexponiert) ; Unterwuchs : gering 
(Südflanke ; Zyklamen) oder teilweise flächendeckend mit Vaccinium myrtillus (Nr. 101, 
zusätzlich Athyrium, Adenostyles, Luzula) ; 620-1250 m.ü.M. 

Habitate : Nr. 0, 6, 101 

Gefährdete Arten : Lopinga achine (4) 

Artenzahl : 8 (keine euzönen Arten, | gefährdet) 

Die obermontanen Rotbuchenwälder sind extrem arten- und individuenarm (Nr. 6 und 
101). 


Zwei Arten mit offenbar zweijähriger Entwicklung kamen im Gebiet 
vor. Der Hochgebirgsspanner Psodos alpinata war nur im Jahr 1992 
ım Gipfelbereich häufig, er fehlte im Folgejahr. Nachweise des Mohren- 
falters Erebia euryale isarica gab es in beiden Jahren an der Nord- 
flanke des Hohen Nocks im Umkreis der Misteleben, der Feichtau und 
des Herzerlsees. In den Habitaten der Südflanke fehlte diese Art 1992 
völlig, trat aber 1993 in einer Höhenlage von etwa 1000 bis 1400 m 
häufig auf. Forster & Wohlfahrt (1976) geben für Erebia euryale „eine 
vermutlich 2-jährige Entwicklung“ an, andere Quellen berichten vom 
Auftreten der Art je nach Gebiet jedes zweite Jahr bzw. alljährlich 
(Lepidopterologen-Arbeitsgruppe, 1991). Möglicherweise ist die Art 
zweijährig und kommt an manchen Orten, wie der Nordflanke des 
Hohen Nocks, in zwei Linien vor, von denen sich die eine in ungeraden, 
die andere in geraden Jahren zu Faltern entwickelt. Die äußerlich sehr 
ähnliche Erebia ligea wurde in den Habitaten des Südabhanges in 
beiden Jahren beobachtet. 


255 


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Abb.3 : Dendrogramm der Gesellschaften tagaktiver Schmetterlinge. 


Struktur der Zünosen tagaktiver Schmetterlinge 


Auf hohem Trenn-Niveau ergaben sich 4 deutliche Klassen (Cluster) von 
Faltergesellschaften (Abb. 3) : 


Die subalpinen/alpinen, südexponierten Felsfluren (F2) bildeten 
wegen ihrer eigentiimlichen Faltervergesellschaftung eine eigene 
Klasse. Diese war durch das gemeinsame Vorkommen von Arten 
charakterisiert, die ihre hauptsächliche Höhenverbreitung entweder 
über oder unter der Baumgrenze haben (Boloria pales, Psodos 
alpinata, Psodos quadrifaria als Hochgebirgsarten sowie Erebia 
oeme, Erebia pronoe, Lysandra coridon). Der Rest betraf Offen- 
lands-Ubiquisten und eine stenotope Edelfalter-Art (Euphydryas 
cynthia). Letztere wurde nur in diesem Habitattyp beobachtet. Am 
größten war die Übereinstimmung im Einzelvergleich noch mit den 
topografisch nahen, lockeren Wäldern bzw. Latschenflächen (Bl, 
L1), besonders fern war die Artzusammensetzung im Vergleich zu 
den einzelnen Habitattypen der Nordflanke (Tab. 3). 

Die artenreichen Faltergesellschaften der Pfeifengrasflur (W3), der 
Felsfluren (F1), des montanen Latschenbestandes (L1), des Fichten- 
waldes (B2), des lockeren Lärchen-Fichten-Waldes (Bl) und der 
untersubalpinen Latschenfläche (L2) hatten die ausschließliche Ver- 
breitung auf der Südflanke des Hohen Nocks bis zur Baumgrenze 
in ca. 1450 m als Gemeinsamkeit und bildeten eine eigene Klasse. 


259 


Tabelle 3 


Diagramm der Ja- und E;,-Werte beim paarweisen Vergleich der Habitattypen. 
Werte größer gleich Ja = 67% bzw. E;, = 0,67 sind markiert. 


a [WE [W2 [WS [RI EP LE LL LC BBA BIT 
21 | 33 | 12 | 29 | 32 | 22 | 19° | 36 | 23 | IS MON | 40) aie 
La 15 130 [29 Léo [sa loaf st [as [st [3s [5e [se | [ls 
38 | 14 | 23 1.27. | 12 | 12-24 | 18 | 13 7172677352 SU 
Lo [sr Lie [31 Loi fmf so [st Las [malas [as [as | 
12172|.297. 21025 7 12.1045 1 8 23 | 27 | 28 | 21 Ws 
a Las [30 Li raf ss [37 as [67 [50 [52 (esol 
8 7 3130 9 18 | 640) 362416 9 7 |W2 
is [as [st | Lac 126 ful. | Ian [on J 
AS 1014 422.33 11- 1.402250 W: 
BHAHHHHETENN 
16 3021225 17. | 432150 F1 
APS. 
20 LT ade lS dd 20% 27001821 F2 
SMAAAAE 
19 | 13. | 15 | 23 | 10 J) LORS 
53 (se far [st [se [se [20 | 
37 4 14 | 43 |50 | 22 BER 
os [ar Kansas | 
16..1.28 Sia 5 2 
19 [30 [9060] os | 
241 22 Aes L3 
26 [es Las Lo || 
21 18 | 12 IL4 
AH 
58 | 12 |B1 
oo Las | 
2 B2 
of 


Die Habitattypen liegen in vorwiegend offenem Gelände (Grasfluren, 
Hochstaudenfluren in Waldlichtungen) und auf skelettreichem Boden 
in sonnenexponierter Lage. Innerhalb der Klasse ergab sich eine 
Zweiteilung : die topographisch benachbarten Habitatgruppen W3 
und Fl setzten sich gegen die übrigen deutlich ab. Ebenfalls fiel 
auf, daß LI und B2 mit E,, = 1 (bei gleichzeitig niedrigem Ja-Wert) 
ident waren. Der Zönose des Latschenbestandes LI kann damit 
als ,,verarmte“ Gesellschaft des umgebenden lockeren Fichtenbe- 
standes B2 gelten. 


EJa 


260 


— Die Zônosen der Gipfelregion des Hohen Nocks (W2) und der 
obersubalpine, lockere Latschenbestand (L3, südexponiert) bildeten 
die dritte Klasse und umfassen die Faltergesellschaften der alpinen 
bzw. obersubalpinen Matten. 

— Die Zönosen der Nordflanke des Hohen Nocks bildeten die vierte 
Klasse (nur B3 ist aus Habitaten der Süd- und der Nordflanke 
zusammengesetzt), wobei die Gesellschaft der Mistelebenweide (W1) 
mit jener der dichten Rotbuchenbestände (B3) eng miteinander 
verwandt und die iibrigen voneinander recht isoliert waren. 


Die Latsche ist weder für die Raupen noch für die Falter der tagaktiven 
Schmetterlinge nutzbar, die in den Beständen dieser Art vorkommenden 
Faltergesellschaften sind daher auf kleine, von Latschen freie Stellen 
angewiesen. Die Faltergesellschaften der untersuchten Latschenbestände 
wiesen deshalb auch wenig Gemeinsamkeiten auf, fielen aber durch 
eine starke Ähnlichkeit zu den benachbarten Habitaten auf (L3/W2, 
L1/B2, L2/B1) (Tab. 3). Zusammenfassend waren die Faltergesell- 
schaften von der Höhenstufe und der Lage entweder auf der Nord- oder 
der Südflanke des Gebirges abhängig, nicht unbedingt aber vom Vege- 
tationstyp. 


Naturschutzorientierte Bewertung der Habitattypen 


Als „wertvoll“ wurden Habitattypen dann bezeichnet, wenn sie euzöne Arten 
beherbergten und eine große Artenzahl aufwiesen. „Besonders wertvoll“ 
war eine Standortgruppe dann, wenn sie von mehreren euzönen sowie 
in Oberösterreich gefährdeten tagaktiven Schmetterlingsarten bewohnt 
wurde. Die Artenzahl mußte ın diesem Fall nicht unbedingt hoch sein 
(Abb. 2). 


Ein Drittel der Habitattypen wiesen hohe Artenzahlen (über 25) auf, 
allen voran montane südexponierte Felsfluren/ Magerrasen (F1), ober- 
montane Hochstaudenfluren der Nordflanke (H2 ; Clossiana thore - 
Vorkommen) und die untermontane Misteleben-Weide (WI). Lockere 
Wälder mit großem Lichtungsanteil beherbergten ebenfalls viele tagak- 
tive Schmetterlingsarten (Bl, B2), was durch eine enge Verzahnung 
unterschiedlicher Habitatteile zu erklären ist (Abb. 2). Das gehäufte 
Auftreten von euzönen und gefährdeten Arten war meist mit der hohen 
Artenzahl korreliert. Die als extensive Rinderweide genutzte Misteleben- 
Alm wurde zu einem großen Teil von Schmetterlingsarten besiedelt, 
die ihren Verbreitungsschwerpunkt im Talbereich haben und als nicht 
gefährdet gelten ; sie wird der großen Artenzahl wegen als „wertvoll“ 
angesehen. Die übrigen der genannten Flächen sınd nach den oben 


261 


angeführten Kriterien als „besonders wertvoll“ für tagaktive Schmetter- 
linge einzustufen. 


Trotz geringer Artenzahl sind jene Habitattypen als „besonders wertvoll“ 
zu beurteilen, in denen mehrere euzöne und gefährdete Arten auftraten. 
Dazu gehören die Gipfelwiese (W2), die obersubalpinen lockeren 
Latschenbestände der Südflanke (L3) und die subalpine Felsflur des 
Knödelkares (F2, Südflanke). In diesen topographisch eng beisammen 
liegenden Flächen befanden sich die Hauptvorkommen der Hochge- 
birgsarten wie Boloria pales und Psodos alpinata. Zusätzlich sind 
Euphydryas cynthia und Clossiana titania cypris nachgewiesen. 


Wenige Arten mit keiner oder lediglich einer euzönen bzw. gefährdeten 
Art bezeichneten die subalpinen Felsfluren der Nordflanke (F3), die 
Latschenflächen (mit Ausnahme von L3) sowie die geschlossenen 
Rotbuchenwälder (B3). Die beiden als gefährdet geltenden Schmetter- 
lingsarten waren nicht auf diese Lebensräume beschränkt, sondern 
kamen auch in anderen vor (Psodos alpinata vor allem ın der Gipfel- 
region auf Matten, Lopinga achine auch im montanen Fichtenwald). 


Neben Kriterien wie Artenzahl und Anzahl gefährdeter und euzöner 
Arten in den einzelnen Habitattypen ist für eine naturschutzorientierte 
Bewertung des Untersuchungsgebietes auch die Vielfalt der darin 
vorkommenden Schmetterlingsgesellschaften von Bedeutung. 


Die insgesamt 105 E,, — Werte der paarweisen Vergleiche der Zönosen 
verteilten sich folgendermaßen : E,, bis 0,33 : 38 Werte ; E,, von 0,34 
bis 0,66 : 51 Werte ; E,, über 0,67 : 16 Werte (Tab. 3). Aus der relativ 
geringen Zahl hoher E,, — Werte kann auf eine große Vielfalt der 
Schmetterlingsgesellschaften im Transekt geschlossen werden. Wertvolle 
Standorte sind daher bezüglich ihrer Artenstruktur weitgehend nicht 
äquivalent und folglich in ihrer Gesamtheit schützenswert. 


Offene, blütenreiche Flächen (Grasfluren, Waldlichtungen, Hochstauden- 
fluren, Weiden) auf der Nord — sowie der Südflanke des Hohen Nocks 
sind für die tagaktive Schmetterlingsfauna von zentraler Bedeutung. 
Besonders die anthropogen weitgehend unbeeinflußten Standorte kön- 
nen als Rückzugsgebiete für gefährdete Falterarten gelten. 


Tagaktive Schmetterlinge der oberen Höhenstufen 


In den Alpen kommen eine ganze Reihe von tagaktiven Groß-Schmet- 
terlingen vor, die ihren Verbreitungsschwerpunkt in der subalpinen und 
alpinen Höhenstufe aufweisen. Sie gehören zu verschiedenen Familien 
der Echten Tagfalter und Dickkopffalter (Diurnea) (Pieridae, Nympha- 


262 


lidae, Satyridae, Lycaenidae, Hesperiidae), dazu kommen tagfliegende 
Arten der ,,Nachtfalter“ (Heterocera) aus den Familien Zygaenidae, 
Arctiidae, Psychidae, Hepialidae, Noctuidae und Geometridae (Forster 
& Wohlfahrt, 1960-1981). 


Für die hôchsten Kalkgebirgsstôcke Oberôsterreichs (Dachstein-Massiv, 
Totes Gebirge, Warscheneck, Haller Mauern) sind folgende tagaktive 
Hochalpenarten gemeldet (Kusdas & Reichl, 1973-1978 ; Huemer & 
Tarmann, 1993 ; Reichl, 1992, 1994) : 


TAGFALTER : 

Grünlicher Heufalter (Colias phicomone, Pieridae ; zusätzlich in der 
obermontanen Stufe), die Mohrenfalterarten Erebia epiphron, Erebia 
eriphyle, Erebia parthe, Erebia glacialis (= pluto) und Erebia gorge 
(Satyridae), der Hochalpen-Perlmutterfalter (Boloria pales, Nympha- 
lidae), der Helle Alpenbläuling (Albulina orbitulus, Lycaenidae) sowie 
die Dickkopffalter Pyrgus alveus alticola, Pyrgus andromedae und 
Pyrgus cacaliae. Zusätzlich liegen noch unsichere Angaben für wenige 
Tagfalter-Arten vor (z.B. Erebia melampus). 


TAGAKTIVE HETEROCEREN : 

Scioptera schiffermilleri (Psychidae), Epichnopterix montanella (Psy- 
chidae), Hepialus ganna (Hepialidae), Caloplusia hochenwarthi (Noc- 
tuidae), Anarta melanopa rupestralis (Noctuidae), Orphne tenebraria 
(Geometridae), Psodos coracina (Geometridae), Psodos noricana (Geo- 
metridae), Psodos canaliculata (Geometridae), Psodos alpinata (Geo- 
metridae), Psodos quadrifaria (Geometridae ; auch im obermontanen 
Bereich). 


Im Sengsengebirge fehlen diese Arten fast vollstandig. Eigene Erhe- 
bungen in den Jahren 1992 und 1993 in der Gipfelregion des Hohen 
Nocks und des Schneeberges erbrachten Nachweise lediglich für den 
Hochalpen-Perlmutterfalter (Boloria pales) und die tagaktiven Spanner- 
Arten Psodos alpinata und Psodos quadrifaria. Auch die tiergeogra- 
phische Datenbank Osterreichs, ZOODAT (Stand 14.5.1992), enthält 
keine weiteren hochalpinen Arten fiir diese Region. 


Für die meisten dieser Hochgebirgsarten waren auch im Sengsengebirge 
offenbar geeignete Lebensräume vorhanden, sodaß für das Fehlen 
dieser Schmetterlinge andere Ursachen zu suchen sind. So könnten sich 
die erwähnten Arten während zwischen- oder nacheiszeitlicher Wärme- 
perioden z.B. im Dachsteinmassiv bzw. im Toten Gebirge auf die höch- 
sten Gipfelbereiche zurückgezogen haben, während sie ım relativ nie- 
drigen Sengsengebirge ausstarben. Auf ähnliche weise deutete bereits 
Holdhaus (1954 : 282-285) die Artenarmut der Karpaten an „exklusiv 


263 


alpinen“ Käfern im Vergleich zu den Ostalpen. Extinktion dieser Arten 
während der Eiszeit und — aus welchen Gründen auch immer — Aus- 
bleiben der Wiederbesiedelung ist als Erklärung jedenfalls nicht stich- 
haltig, weil das Sengsengebirge während der Eiszeiten zum großen Teil 
gletscher- und firnfrei war (Franz, 1954). 


Bei manchen Arten ist das Bild komplizierter : Der Griinliche Heufalter 
(Colias phicomone, Pieridae) ist in den oberösterreichischen Kalkalpen 
und deren Vorbergen, z.B. am Schoberstein, ab der montanen Stufe 
verbreitet und fehlt nur am Traunstein und im Sengsengebirge (Kusdas 
& Reichl, 1973). Möglicherweise überlebte auch diese Art die Wärme- 
zeiten nur auf höheren Gipfeln, könnte sich aber dank einer (hier vor- 
ausgesetzten und nicht belegten) Wandertätigkeit in geeigneten Lebens- 
räumen wieder angesiedelt haben. Das Fehlen im Sengsengebirge und 
am Traunstein wäre demnach als Ausbreitungslücke zu deuten. 


Danksagung 


Für die effektive und liebenswürdige Unterstützung mit Schmetterlingsdaten 
aus der ZOODAT (Tiergeografische Datenbank Österreichs) danke ich Herrn 
Univ.-Prof. Dr. Ernst R. Reichl, Linz. 


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MUHLENBERG, M., 1989. Freilandökologie. 2. Auflage. UTB 595. 431 pp. 
Quelle und Meyer, Heidelberg, Wiesbaden. 

REIcuL, E. R. (1992, 1994). Verbreitungsatlas der Tierwelt Österreichs. Band 1, 
Tagfalter (1992) ; Band 2, Spinner und Schwärmer (1994). Forschungs- 
institut fiir Umweltinformatik, Linz. 

SCHWERDTFEGER, F., 1975. Ökologie der Tiere. Band III : Synökologie. 451 pp. 
Parey, Hamburg. 

WIMMER J., 1991. II. Beitrag zur Kenntnis der Macrolepidopterenfauna des 
Reichraminger Hintergebirges in Oberösterreich. Steyr. Ent. 25 : 1-41. 


265 


Nota lepid. 18 (3/4) : 266 ; 13.V.1996 ISSN 0342-7536 


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266 


Nota lepid. 18 (3/4) : 267-280 ; 13.V.1996 ISSN 0342-7536 


Butterfly diversity and rarity within selected habitats 
of western Andalusia (Spain) 
(Lepidoptera : Papilionoidea and Hesperioidea) 


J. M. MouiNA* & J. M. PALMaA** 


* CIDA “Las Torres-Tomejil”, Aptdo. Oficial, E- 41200 Alcala del Rio (Sevilla), Spain 
** c/ Candelilla, 5, 1 izqda, E-41006 Sevilla, Spain 


Summary 


A study was made of the butterfly fauna associated with six habitat types 
in eight localities of western Andalusia (Spain). Four assessment methods 
identified kermes oak wood and evergreen oak forest as the habitats supporting 
the greatest diversity and rarity of butterflies and skippers both at local and 
regional scale. Results suggest an important influence of woody plants, by 
means of microclimate effects, on butterfly communities. A comparison of the 
assessment methods, reveals that the Shannon-Weaver index (H’) is a poor 
discriminant of diversity in this study, and that richness and rarity are sufficient 
and better measures for site conservation assessment. 


Résumé 


Les auteurs ont étudié la faune des lépidoptères diurnes sur six types de bio- 
topes dans huit localités d’Andalousie occidentale (Espagne). Quatre méthodes 
d'évaluation ont permis d’identifier les bois de chênes kermès et la forêt de 
chênes à feuilles persistantes comme les biotopes abritant la plus grande 
diversité de Diurnes et d’Hespérides, ainsi que les plus rares, tant sur le plan 
local que régional. Ces résultats font penser que les lieux boisés exercent une 
forte influence sur les communautés de Diurnes du fait de leurs effets sur 
le microclimat. En comparant les méthodes d’évaluation, on découvre que 
l’Index Shannon-Weaver (H’) a peu de valeur pour distinguer la biodiversité 
dans la présente étude, et que la richesse de la faune et la rareté de ses espéces 
sont suffisantes et plus efficaces pour apprécier l’importance de la conservation 
des sites. 


Introduction 
Habitat loss is considered to be by far the most significant threat to 


insect conservation, with agricultural management as the major con- 
tributor. As agriculture continues to expand, natural lands become 


267 


more and more fragmented, increasing the need to integrate regional 
land use and conservation strategies (Van Hook, 1994). 


Insects, and especially butterflies, are receiving much attention because 
of their potential as indicator species of landscape structure and eco- 
logical changes (Ferrin & Martin-Cano, 1994 ; Kitahara & Fuji, 1994 ; 
Daily & Ehrlich, 1995). Surveys and monitoring have been successfully 
used for habitat assessment, having direct management implications 
for land acquisition and reserve planning (Cheverton & Thomas, 1985 ; 
Munguira & Thomas, 1992 ; Wood & Samways, 1992 ; Pollard & Yates, 
PCR) 


This paper deals with the diversity and rarity of the butterfly fauna 
in several habitats within the Mediterranean ecosystem of western 
Andalusia, an area which has been little investigated entomologically. 


Materials and methods 


Study area, sites and sampling 


The study was carried out during five months, between 5 March and 
21 July, 1994, with the exception of the second half of April and first 
half of May, due to bad weather. Eight sites were surveyed in the 
Penibetic mountains, approximately 100 km south-east of Seville 
(western Andalusia, Spain), in a mosaic landscape of agriculture, settle- 
ments and seminatural areas (Fig. 1). This part of our province has 
a typically Mediterranean climate that changes from the dry to the wet 
type with increasing altitude, here from 100 to 1100 m. The substrate 
is mainly calcareous (see also Molina & Palma, 1992 ; 1994). 


According to Rivas-Martinez (1987), potential vegetation belongs to the 
evergreen oak forest (Quercus rotundifolia Lam.). There is no natural 
forest today, and the actual vegetation comprises a mosaic of scrub 
types, successional stages, and climax remnants together with crops, 
mainly olive groves, cereal and sunflower fields. 


Butterflies and skippers were sampled by conducting fixed transect 
walks, mostly 200 to 400 m in length, recording the number of 
individuals of each species seen within 2.5 m either side of and 5 m 
in front of the recorder (Pollard, 1977 ; Descimon & Napolitano, 1991 ; 
Pollard and Yates, 1993). Population density for each transect is 
calculated as the number of individuals seen within the 5 m square 
per hundred meters of transect walk (an area of 500 m?) (Cheverton & 
Thomas, 1985), and the values obtained used in calculations of diversity. 
A total number of 48 samples evenly distributed among all sites were 
studied. 


268 


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269 


Local transects were divided into sections which coincide with changes 
in the nature of the vegetation (Pollard, 1977). Six distinct vegetation 
types, as an approximation to habitats, were subjectively distinguished, 
mainly according to the structure of dominant plant Pe (see 
Table 1) : 


(1) Evergreen oak forest is dominated by Quercus rotundifolia (Lam.) growing 
in association with Olea europaea L. and Ceratonia siliqua L.; plants such 
as Pistacea lentiscus L., Daphne gnidium L. or Smilax aspera L. are also 
characteristic. They appear mainly on limestone substrate. Clear-cut under- 
growth is usual as a result of forestry and grazing (‘dehesa’ landscape). 
Intensively managed evergreen oak forests usually have the shrub layer 
dominated by Cistus spp. 


(II) Kermes oak wood occurs naturally in the Penibetic mountains on steeper, 
stony slopes, impoverished soils and sunny sites. Quercus coccifera L. is 
dominant together with Asparagus albus L., Rhamnus oleoides L., and Cra- 
taegus monogyna Jacq. ; Aristolochia baetica L. is also characteristic. 


Open scrub occurs mainly as a result of grazing or as a consequence of 
abandoned cropping. Sites have characteristically dry and bare ground from 
late spring to autumn; there are some shrub species, climax remnants or 
early successional ones. 


(III) Thyme grassland occurs on sunny and calcareous sites as the penultimate 
or last retrograde successional stage in the natural vegetation series. Species 
of Fabaceae, Asteraceae and Lamiaceae, especially Thymus spp. are dominant. 
There are few grasses and herbs (Rivas-Goday & Rivas-Martinez, 1969). 


(IV) Broom scrub is dominated by Retama sphaerocarpa (L.) Boiss. in 
association with grasses and some herbs of small size which form a short 
turf. 


(V) Grasslands are structurally simplified habitats, the result of the complete 
elimination of woody plants, usually by fire or extensive grazing. Grasses and 
thistles are characteristic in the late spring and summer, giving a similar 
physiognomy to that of broom scrub. 


(VI) We have also considered road verges as a distinct habitat. Herbs and 
weeds here are most diverse, resulting from the introduction of many annual 
and biannual nitrophilous species. Examples of the climactic plant species 
are usually also present. 


Data analysis 


Four measures for the quantity and/or quality of Rhopalocera species 
were used to compare habitats: (a) species richness, i.e. the total 
number of species present, irrespective of their rarity ; (b) the Shannon- 
Weaver diversity index (H’) (Southwood, 1978); (c) the proportion 


270 


Habitat type | No. 


Evergreen 
oak forest 


Thyme 
grassland 


Table | 


Some characteristic habitats in the study area. 


Formation type 
(Southwood, 
1978) 


Woodland 
Trees dominant 


Scrub 
dominant plants 
not exceeding 
shrub layer 
height, usually 
not over 7.6 m 


Field 

dominant plants 
coincide with 
field layer, 
usually not more 
than 2 m high 


Open-ground 
dominant plants 
not usually more 
than 15 cm in 
height 


Field/ open- 
ground 


Local characteristics 


Hunting and 
forestry. 

Grazing, mainly 
cows. Every three 


years crops of cereal 


or sunflower 


Hunting. Occasional 


grazing. No more 


than 3-4 m in height 


None. Near olive 
groves. Vegetation 


seldom reaches more 


than | m in height. 
Bare ground 


Grazing for 5-8 
month/year, few 
cattle, mainly goats 


Grazing with few 
cattle, mainly goats 
and sheeps 


None. Occasional 
grazing. 
Nitrophilous plant 
communities. 


Approximate plant 
sociological type 


Paeonio-Quercetum 
rotundifoliae 
Riv.Mtnez., 1964 
transitional to Smilaci- 
Quercetum 
rotundifoliae Barber, 
Quezel & Riv.Mtnez., 
1981 


Asparago-Rhamnetum 
oleoidis Riv.God., 1959 
em Riv.Mtnez., 1975 


Teucrio- Corydothime- 
tum baeticum 
Riv.God. & 
Riv.Mtnez., 1969 


Saturejo- Corydothi- 
mion Riv.God. & 
Riv.Mtnez., 1968. 
Similar to Genisteto- 
Retametum 


Thero- Brachypodion 
(Br.-Bl., 1925) 
Riv.Mtnez., 1977 
Astragalo- Poeion 
bulbosae Riv.God. & 
Ladero, 1970 


Brometalia rubenti- 
tectori Riv.Mtnez. & 
Izco, 1977. 

Scolymo- 
Onopordetalia nervosi 
Br.-Bl. & O.Bolés. 
1957 corr Riv.Mtnez., 
1975 


Dr 


of species that are locally rare (LRS) ; and (d) the species rarity factor 
(SRE), an index ignoring common species and weighted towards those 
rarest in Seville province as a whole. 


The latter two measures were calculated as follows : (c) species of 
restricted distribution in the study area, locally rare species (LRS), were 
defined as those recorded to date in fewer than 50 % of the total number 
of UTM squares (10 km?) prospected (Molina & Palma, 1994, and 
some unpublished data). The proportion of locally rare species was 
expressed as the ratio of the number of these species to the total number 
of species on a site. 


The relative rarity of the species selected in (c) was quantified by 
assigning each a score reflecting its distribution for UTM squares in 
Seville province following the octave method proposed by Dony & 
Denholm, 1985) : 


No. of UTM (100 km?) squares 
with species present 1 2-3 4-7 8-15 16-31 >31 


Score 6 à 4 3 2 1 


Some unpublished data on geographical ranges taken from Molina & 
Palma (1995) were used as a source of distribution data. Summation 
of the scores for each locally rare species (c) gave (d) the SRF, the 
fourth measure used to assess habitats (see also Dony & Denholm, 
1985). 


Correlation among the assessment measures was carried out using the 
Spearman rank correlation coefficient (rs) (Southwood, 1978). 


Results 


A grand total of 1403 individuals of 47 species were registered during 
this study. Details of recorded species are included in Molina & Palma 
(1994 ; 1995 ; Table 2). The total number of species in the habitats 
ranged from 20 to 33, with a mean of 26 species. Tables 3 and 4 sum- 
marise the results obtained with the four assessment methods applied 
at each locality and habitat type. 


The greatest number of species and diversity were found in evergreen 
oak forest and in kermes oak wood. Minimum species richness and 
diversity were obtained both locally and as mean values in broom scrub, 
grassland and along road verges. 


272 


Table 2 


List of species under study. Names follow mainly those contained in Higgins (1975), 
and Gomez-Bustillo & Arroyo-Varela (1981, 1984) with some modifications 
in the arrangement of families (Ackery, 1984). 


Hesperiidae Pieridae 

Spialia sertorius (Hoffmansegg, 1804) Aporia crataegi (Linnaeus, 1758) 

Syrichtus proto (Ochsenheimer, 1808) Pieris brassicae (Linnaeus, 1758) 

Carcharodus alceae (Esper, 1780) Pieris rapae (Linnaeus, 1758) 

Carcharodus boeticus (Rambur, 1839) Pontia daplidice (Linnaeus, 1758) 

Thymelicus acteon (Rottemburg, 1775) Euchloe ausonia (Hübner, 1804) 

Thymelicus flavus (Brünnich, 1763) Euchloe tagis (Hübner, 1804) 

Lycaenidae Euchloe belemia (Esper, 1799) 

Lycaena phlaeas (Linnaeus, 1761) Anthocharis belia (Linnaeus, 1767) 

Quercusia quercus (Linnaeus, 1758) Colias crocea (Geoffroy in Fourcroy, 

Satyrium esculi (Hübner, 1804) 1785) 

Satyrium spini (Denis & Schiffermiiller, | Gonepteryx cleopatra (Linnaeus, 1767) 
1775) Papilionidae 

Tomares ballus (Fabricius, 1787) Papilio machaon Linnaeus, 1758 

Cupido lorquinii (Herrich-Schäffer, 1847) Iphiclides podalirius (Linnaeus, 1758) 

Pseudophilotes abencerragus (Pierret, 1837)  Zerynthia rumina (Linnaeus, 1758) 

Glaucopsyche melanops (Boisduval, 1828) 

Aricia cramera (Eschscholtz, 1821) 

Lysandra albicans (Herrich-Schäffer, 1851) 

Lysandra bellargus (Rottemburg, 1775) 

Polyommatus icarus (Rottemburg, 1775) 

Nymphalidae 

Vanessa atalanta (Fabricius, 1807) 

Vanessa cardui (Linnaeus, 1758) 

Melitaea aetherie (Hübner, 1826) 

Euphydryas aurinia (Rottemburg, 1775) 

Hipparchia statilinus (Hufnagel, 1766) 

Hipparchia fidia (Linnaeus, 1767) 

Melanargia occitanica (Esper, 1793) 

Melanargia ines (Hoffmansegg, 1804) 

Maniola jurtina (Linnaeus, 1758) 

Hyponephele lupinus (Costa, 1836) 

Pyronia bathseba (Fabricius, 1793) 

Pyronia cecilia (Vallantin, 1894) 

Coenonympha pamphilus (Linnaeus, 1758) 

Coenonympha dorus (Esper, 1782) 

Pararge aegeria (Linnaeus, 1758) 

Lasiommata megera (Linnaeus, 1767) 


293 


Table 3 


Species richness (S), Shannon-Weaver diversity (H”), locally rare species (LRS) 
and (proportion), and species rarity factor (SRF) at habitats and sampling sites. 
Spearman rank correlation values (r,). Critical value n = 19, p < 0.05, 

2 tails = 0.454) 3 Ts(S/H') = 0.796 3 Ts (S/LRS) = 0.581 5 Ts (S/ SRF) = 0.567 5 


Ts(H'/LRS) — 0.476 ; I, (H’/SRE) — 0.435 ; Ts(LRS/SRE) — 0.987). 


Site number and name | S | H’ | LRS |SRF\ Habitat type 
Sierra Vaquera 5 (0.24) | 20 | Evergreen oak forest. 


11 | 1.61 | 3 (0.27) | 14 | Broom scrub. 
Pinus spp. afforestation 
(2-3 years old). 


2 | Mte. Becerrero 4 (0.24) | 17 | Kermes oak wood. Some 


Quercus spp. trees. 
.00) 


4 | Mte. Aguila 


Grassland. Abandoned crop. 
Road verge. 


. 


© 
—> 
ir 
© 
© 
= 


Grassland. Grazing. 


Road verge. 


NTN] 
&} N | oO 
OI 
N 
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Kermes oak wood. 


n| & 
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DE 

N}N 

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Thyme grassland. 


Road verge. 


| Evergreen oak forest. 
Kermes oak wood.  : 


wos Grassland. Grazing. 


12 
mea Cistus spp. undergrowth. 


| 
1.72 | 1 (0.14) Broom scrub. À lot of bare 


La Gomera 
ground. 
7 | Fte. La Roya 22052 |S ORS) Evergreen oak forest. Clear- 
cut undergrowth. 
18 | 2.31 | 4 (0.22) Thyme grassland. 


Barrancos Blancos | 21 | 2.63 | 3 (0.14) Broom scrub. Some climactic 
plant species remnants. 


11] 2.12 [0 (0.00) | 0 [Roadverge DD 


Fifteen regional rare species have been found in the sampled habitats, 
20% of those occurring in Seville province. Six are very rare, occupying 
3 or less UTM 100 km? squares, and 21 are locally rare within the 
-study area (Molina & Palma, 1995). 


The mean number of LRS was 6, among 1/3 and 1/4 of the species 
in each habitat (Tables 3,4). Kermes oak wood has the greatest mean 
of LRS. In the evergreen oak forest the proportion of LRS has a 
mean value of 1/5. The species rarity factor (SRF) was also greater 


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274 


Table 4 


Mean values and (coefficient of variation) of abundance (Ab), species richness (S), 
Shannon-Weaver diversity index (H’), locally rare species (LRS), 
and species rarity factor (SRF) in the six habitat types. 
n = sites visited. 


Habitat n Ab S H’ LRS SRF 
Evergreen oak forest | 4 224.5 18.50 238 325) 10275 
0759) (14.54) (10.35) (38.72) (50.29) 

Kermes oak wood 3 169.3 19.00 DS 4.67 ACT 
(39.25) (22.74) (12.74) (24.74) (22.88) 

Thyme grassland 2 216.5 17.50 225 4.50 17.50 
(17:78) (2.85) (4.09) (IS) (12210) 

Broom scrub 3 178.3 13.00 1899 2235 9.00 
(52.00) (45.23) (28.18) (49.49) (89,59) 

Grassland 3 185.3 12.30 2.08 0.00 0.00 
(37.68) (21.30) (4.87) (0.00) (0.00) 

Road verge 4 183.5 11.50 2.18 1.00 3.05 

i (32.48) (18.86) (11.21) (81.65) (70.13) 


both locally and as a mean value in kermes oak wood and in evergreen 
oak forest (Tables 3, 4). 


During this study none of the rare species occurred in all habitats 
(Table 5), though they tended to be biased to evergreen oak forest 
and woods. Nymphalidae and Lycaenidae are the families with the 
greatest number of locally rare species. The latter includes the more 
habitat-restricted species and has the greater mean rarity score (see 
Table 5). 


Some successional stages contain very rare species, which increase the 
values of both LRS and SRF (Table 4). This result is in agreement 
with that obtained by Thomas & Mallorie (1985) for the Moroccan 
butterflies. 


Discussion 


The richness and diversity pattern obtained within the study area is 
similar to that described from other agricultural areas in the Iberian 
Peninsula, although there are some changes in species composition and 
smaller H’ diversity values (Sanchez & Viejo, 1986 ; Viejo, 1986 ; Baz, 
1986). A possible explanation is the expansion of cereal crops and 
olive groves as monocultures in the region. On a regional scale, a mosaic 
of various biotope patches provides a high degree of habitat hetero- 
geneity that may enhance species richness and rare species survival 


275 


Table 5 


Distribution of locally rare species among habitat types. 
I= Evergreen oak forest, // = Kermes oak wood, /// = Thyme grassland, 
IV = Broom scrub, V = grassland, and V/= road verge. 


Presence (+) in the habitats Occurrence in — 
Seville province 
Species I UL PTE DAZ V VI | No. of Rarity 
UTM score 
squares 
+ 
iy Ar 


Carcharodus. alceae 
+ 
+ 
+ 
Lea 
+ 
ane 


Carcharodus boeticus 

Quercusia quercus 
Satyrium spini 

* one sole individual, near a well-conserved evergreen oak forest. 

P = present, but not recorded during the sampling period included in this study, 

and not considered in calculations. 


+ 


Tomares ballus 
Cupido lorquinii 
Glaucopsyche 
melanops 

Lysandra albicans 
Lysandra bellargus 
Vanessa atalanta 
Melitaea aetherie 
Euphydryas aurinia 
Hipparchia statilinus 
Hipparchia fidia 
Melanargia occitanica 
Hyponephele lupinus 
Coenonympha dorus 
Pararge aegeria 
Aporia crataegi 
Anthocharis belia 
Iphiclides podalirius 


+++++ 


— 
+ 
+++ 


++ ++ 
= 
+ 


+ 


— 
D PU DPD Un © R Un ND WN NOB BB BR © 


+r + +++ 
+ 
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= 


(Duelli er al., 1990), whereas homogeneity within our study area should 
produce quite the reverse. 


Local butterfly richness and diversity have been correlated with plant 
species richness and habitat complexity by several authors (Erhardt, 
1985a,b ; Thomas & Mallorie, 1985 ; Baz, op. cit. ; Spitzer et al., 1987 ; 
Leps & Spitzer, 1990). In this study evergreen oak forest and kermes 
oak wood have the greatest values for both richness and diversity ; 
this is perhaps to be expected, since these habitats have suffered least 
human disturbance, thereby conserving many of the plant species of 
the potential climax vegetation of the region (Rivas-Martinez, 1987). 


Within habitats, diversity of woody plants has been considered as an 
indicator of the diversity of food resources for insects (Lawton, 1983) ; 


276 


but also trees, shrubs and climbers are most important components 
of habitat complexity and heterogeneity, providing varying types of 
microhabitats which would be differentially suitable for species in space 
and time (Thomas, 1984 ; Spitzer et al., 1987 ; Munguira & Thomas, 
1992 ; Wood & Samways, 1992). 


In our Mediterranean ecosystem, climate makes many sites unsuitable 
for butterflies because of the annual late spring and summer drought 
from mid-June until autumnal rains. During this time, open habitats, 
such as low scrub, grassland, and bare ground are the most unsuitable, 
whereas shaded habitats act as refuges for butterfly species (Viejo ef 
al., 1992). Woody plants provide shade, protection from desiccation 
by wind, lower midday temperatures, and other microhabitat modi- 
fications, including floristic composition (Montoya-Oliver, 1982 ; Mon- 
toya-Oliver & Mesôn-Garcia, 1982). 


Many open habitats have been mainly caused, and are used by cattle. 
Grazing by sheep and goats should reduce diversity and floristic 
composition of plant communities, especially shrubs (Van Hook, 1994), 
particularly affecting the rarest butterflies in the study area. 


Field data support the important influence of woody plants on com- 
munities (compare Tables 1, 2 and 3), but as no objective measure- 
ments were made of the vegetation diversity or structure in the habitats, 
this needs further research. However, this should explain both mean 
abundance and local rarity values obtained for kermes oak woods, 
since these habitats have the best floristic and structural conservation. 
This is mainly owing to their occupying marginal land of low agri- 
cultural value and their management as hunting reserves. Positive effects 
of shrubs on butterfly communities have been recently reported by 
Arechavaleta et al. (1994) and Ferrin & Martin-Cano (1994). 


The Penibetic Mountains give shelter to many regionally uncommon 
or rare species of butterflies (Molina & Palma, 1995). In this study 
these species are generally associated with the better conserved sites 
and vegetation types. The presence in some successional habitats of 
very rare species. (e.g. Lysandra albicans (Herrich-Schäffer, 1851) and 
Lysandra bellargus (Rottemburg, 1775) in thyme grassland, or Meli- 
taea aetherie (Hiibner, 1826) in broom scrub) may be explained by 
the existence of specific ecological requirements for the larval stages 
(Thomas, 1984). 


Rarity patterns and index values obtained also suggest actual differences 
among habitats, even when these are very close to each other, and 
support a sedentary character for rare butterflies (Erhardt, 1985a,b). 


Da 


High values of species diversity are often an important, if not a unique, 
criterion when selecting sites for conservation (Margules & Usher, 
1981). In this study, diversity values were very similar among habitats 
and did not represent a good discriminant measure. Only the H’ index 
was well associated with species richness, whereas there was little or 
no correlation with rarity values (Table 3). Relative similarity in diver- 
sity values among habitats may be due to influx of species from adjacent 
habitats during late spring and summer, which could vary from year 
to year, depending on the weather. 


Conclusions 


All assessment methods using butterfly communities as indicators, 
identified kermes oak wood (©. coccifera) as the least disturbed and 
most interesting habitat in the study area for nature conservation. 


There is indirect evidence that within habitats the presence of woody 
plants act as microclimate modifiers during the hottest period of the 
flight season. Together with the intensity of human influence upon 
vegetation, which decreases diversity, these seem to be the determining 
factors of butterfly species richness, diversity and rarity in the study 
area. 


Butterfly conservation policy must focus on kermes oak woods if 
characteristic Mediterranean communities are to be preserved, whereas 
attempts to conserve individual species must involve a study of their 
specific habitat requirements within the succession. When assessing 
sites, species richness, together with an evaluation of the faunal quality, 
would be a better discriminant than diversity index values. 


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CHEVERTON, M. R. & THomas, C. D., 1985. Land use and conservation 
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Descımon, H. & NAPorITANO, M., 1990 (1991). L’étude quantitative des 
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280 


Nota lepid. 18 (3/4) : 281-288 ; 13.V.1996 ISSN 0342-7536 


Larval food shortage and adult dispersal 
in Callimorpha dominula (L.) 
(Lepidoptera : Arctiidae) 


Denis F. OWEN 


School of Biological and Molecular Sciences, Oxford Brookes University, Headington, Oxford OX3 OBP 


Summary 


In Oxfordshire, England, the moth Callimorpha dominula (L.) occurs in well- 
established colonies in damp and marshy habitats. Specimens from the within 
colonies are smaller than those from outside, evidently because larvae just 
out from hibernation feed on an extremely limited food source. It is suggested 
that moths constantly leave the main colonies and found small and perhaps 
temporary breeding populations elsewhere, but whether they leave as a result 
of starvation in the larval stage is a matter for conjecture. 


Résumé 


En Angleterre, dans l’Oxfordshire, l’Arctüde Callimorpha dominula (L.) se 
présente en colonies bien établies dans certains biotopes humides et marécageux. 
Les exemplaires provenant de ces colonies sont plus petits que ceux de 
l'extérieur, évidemment parce que leurs chenilles sont juste sorties de l’hivernage 
ne disposent pour se nourrir que de ressources très limitées. On peut supposer 
que certains imagos quittent régulièrement les colonies principales et fondent 
de petites populations, peut-être temporaire qui se nourrissent ailleurs, mais on 
ne peut que des conjectures quant à savoir si ces départs sont dus au manque 
de nourriture pour le stade larvaire. 


In Oxfordshire, England, the scarlet tiger moth, Callimorpha dominula 
(Linnaeus, 1758), forms discrete colonies in damp marshy places, 
especially in the vicinity of the River Thames which flows through 
the county. In some years a colony may comprise tens of thousands 
of adult moths, while in others there may be no more than a few 
hundred. The population size of one colony, at Cothill, has been 
monitored since 1939, chiefly to detect changes in frequency of a mutant 
allele (medionigra) that affects wing coloration. This is a rare example 
of long-term ecological research. Indeed, the first 32 years of work 
at Cothill stimulated Ford (1975) to claim that, “No natural population 


281 


of animals in the world has been so fully quantified as that of the 
Scarlet Tiger moth, Panaxia dominula”. 


The moths (and larvae) are repeatedly (and perhaps increasingly) found 
outside the main colonies, either as strays or as temporarily established 
populations of small size and low density. These occurrences never 
seem to lead to the formation of strong colonies like the one at Cothill. 


C. dominula has a single generation a year, flying (usually in sunshine, 
but also at night) in July. The eggs are scattered over vegetation and 
hence young larvae (not female moths) select potential foodplants. The 
larvae feed in August-October, then hibernate in the third instar beneath 
leaf litter, sometimes feeding a little on warm winter days, especially 
in February and March when a few new plant shoots may become 
available. Feeding recommences in April and pupation occurs in June. 
The two main foodplants are Symphytum officinale (Boraginaceae) 
and Eupatorium cannabinum (Compositae), but in years of high 
population size up to 43 species of plants belonging to 28 families 
may be utilised (Owen, 1994). This expansion of hostplant range in 
years of high population is associated with the depletion of Symphytum 
leaves earlier in the season. The high diversity and disparity of food- 
plants occurs in May, when the vegetation in the colonies is well- 
developed ; in April, when larvae emerge from hibernation, there are 
few plants available, except Symphytum and the rosettes of Cirsium 
(Compositae). 


Adult size 


Several workers have commented on the relatively small size of moths 
in the Cothill colony. Moths from North Hinksey, a nearby high density 
colony are also small. Accordingly, I measured the wing lengths of 
moths obtained from Cothill and Hinksey. I also measured the wing 
lengths of moths obtained from various sites outside the main colonies. 
I refer to these moths as the “outsiders”. The measurements were made 
with a vernier calliper (to the nearest 0.1 mm). A wing was measured 
from its apex to the middle of the thorax, a measurement that 
accurately reflects body size. The results are given in Table 1 as N (the 
sample size), x (mean wing length) and SD (the standard deviation 
of the mean). In Table 1 the relatively small sample size for the outsiders 
is because of difficulty in obtaining specimens from the sites where 
they occur. All moths were wild-caught, none reared or bred. 


Males and females are of similar size and do not differ significantly : 
Cothill, = 1.703, d.f. = 38, North Hinksey, t = 1.365, d.f. = 38, and 


282 


Table | 


Wing length (in mm) of Callimorpha dominula in 
three samples from Oxfordshire. 


Cothill males 
females 


North Hinksey males 
females 

outsiders males 
females 


outsiders, { = 0.661, d.f. = 10, and so the sexes can be added together 
for the inter-site comparisons. Comparing Cothill to outsiders, t = 6.924, 
d.f.=50, p< 0.001, and North Hinksey to outsiders, ¢ = 7.987, 
d.f. = 50, p < 0.001). Both comparisons give statistical confirmation 
to the small size of Cothill and North Hinksey moths and the relatively 
larger size of the outsiders. 


The origin of the outsiders 


Fig. | shows the distribution of C. dominula in Oxfordshire. The species 
is exceptionally well-recorded, chiefly because of its attractive coloration 
and mainly diurnal flight activity. The distribution shown includes well- 
established colonies (three are marked on the map) and occasional 
sightings which may or may not indicate the existence of a colony: 
many recorders do not differentiate between stray individuals and 
colonies. 


North Hinksey (A in Fig. 1) was established artificially in 1951 by 
the introduction of 4000 fertile eggs which, assuming 200 eggs from 
each female, came from about 20 pairs of moths. The circumstances 
of its establishment are given in Sheppard & Cook (1962), and there 
is little doubt that the site was chosen because it looked suitable, i.e. 
like Cothill. The point of the North Hinksey introduction was to 
monitor the fate of the medionigra allele which has been introduced 
at a known frequency. An essential requirement for such monitoring 
is that colonies under investigation should be ecologically and geo- 
graphically isolated from one another to an extent that alleles are not 
exchanged. Virtually all the early publications on C. dominula claim 
that colonies are isolated and that allele exchange is unlikely. Evidently 
E.B. Ford and his colleagues did not encounter outsiders as they are 
not mentioned in the numerous publications on the population genetics 
of C. dominula (review in Jones, 1989). Indeed, Ford (1975), stressing 


283 


gr met | 


Fig. 1. Map of Oxfordshire showing records of C. dominula in 2 X 2 km squares 
in the period 1960-1993, and the location of the North Hinksey (A), Cothill (B) and 
Sheepstead Hurst (C) colonies. The distribution of the moth is strongly associated 
with the River Thames, as shown. The large squares are 10 X 10 km. The map is derived 
from information held by the Oxfordshire Biological Records Centre, Standlake, 
Oxfordshire. 


284 


the isolation of the Sheepstead Hurst colony (C in Fig. 1) writes, “The 
whole area is completely surrounded by agricultural land and a neigh- 
bouring aerodrome, and the insect, though sometimes present in great 
numbers, seems to be closely restricted to this habitat and completely 
isolated [my italics] from any of its other colonies, the nearest of which 
is that at Cothill”. 


On 2 May 1993, I walked from Sheepstead Hurst to Cothill, following 
the small stream (Sandford Brook) that links the two sites. Along the 
banks of the stream there were Symphytum plants and C. dominula 
larvae were found feeding on many of them, as shown in Fig. 2. Away 
from the stream the land is agricultural (exactly as described in Ford 
(1975)) and there are no Symphytum or other plants suitable as food 
for C. dominula. It is difficult to imagine that the situation was sub- 
stantially different 30 years earlier : the pattern of land use has probably 
changed, but the essential structure of the countryside would have been 
much the same then as now. Thus it is likely that these two “isolated” 
colonies have been and still are linked together by a corridor of Sym- 
phytum. Moths found in the vicinity of black spots shown in Fig. 2 
would count as outsiders. Hence one source of outsiders 1s that female 
moths follow Symphytum corridors and deposit eggs. This could lead 
to the establishment of small (perhaps temporary) populations away 
from the main colonies. 


Both moths and larvae are repeatedly found in the densely populated 
and built-up environment of the City of Oxford. They are reported 
from gardens and from weedy patches where the land has recently 
been disturbed. It is difficult to believe that these individuals are the 
offspring of females that have followed a Symphytum corridor from 
one of the established colonies. One possibility is that contrary to 
expectation, the moth disperses for long distances over unsuitable 
habitat, and that it does this quite frequently, a possibility never enter- 
tained by the many previous C. dominula workers. Another is that 
the diffuse City of Oxford population is derived from accidental escapes 
or deliberate releases from captive stocks held by the Zoology Depart- 
ment, University of Oxford, in the 1950s and 1960s. Whatever the 
explanation, the apparently hostile environment of the City of Oxford 
contrasts markedly with the damp, marshy habitat occupied by the 
main colonies. 


Larval starvation and dispersal 


As Ford (1975) claimed, C. dominula 1s indeed a well-studied species, 
but there are questions about its ecology that remain unanswered. For 


285 


S 
N 
oe * 


Fig. 2. The distribution of larvae of C. dominula (solid dots) in the area between 
colonies (triangles) at Cothill and Sheepstead Hurst. 


286 


example, why are the main colonies located in wet, marshy places, 
and yet low density breeding can occur almost anywhere ? And why 
are adult moths from the main colonies significantly smaller than the 
outsiders ? 


At Cothill, the site occupied by the colony is dominated by the reed, 
Phragmites, while at North Hinksey a tall horsetail, Equisetum telma- 
teia provides the main ground cover. At North Hinksey in particular, 
the moth is confined to only a relatively small area of the apparently 
suitable habitat, and it is difficult to see why this should be so. At 
both sites the early spring larvae usually eat all the available Symphytum 
shoots as they appear and are clearly deprived of food. Symphytum 
starts to grow in February, considerably earlier than virtually all other 
possible foodplants, and it is not until later in the spring (mid-April 
in most years) that additional foodplants become freely available. The 
inevitable conclusion is that third instar larvae just out of hibernation are 
in danger of starving, especially in years of exceptionally high population 
density. This conclusion is not new: more than 30 years ago Cook 
(1962) suggested that at Cothill larvae might sometimes starve. 


Cook (1961) measured wing lengths of C. dominula from Cothill and 
compared them with similar measurements of specimens from a colony 
in south-west England at Axmouth, Devon. His Cothill measurements 
are virtually identical to those of mine given in Table 1, differing by 
only 0.2 mm (males, N = 27) and 0.1 mm (females, N = 25). Interesting- 
ly, moths from Axmouth were considerably larger than those from 
Cothill, but are virtually identical to my Oxfordshire outsider measure- 
ments, differing by 0.1mm for males (N=68) and for females 
(N = 42). Cook (1961) concluded that Cothill is at the edge of the 
moth’s geographical range and that the size difference between Cothill 
and Axmouth is determined primarily by environmental dissimilarities 
between the colonies which, by implication, could be larval food 
shortage at Cothill in early spring, but not in the milder area of Ax- 
mouth. 


This explanation does not, however, account for the large size of the 
Oxfordshire outsiders, all obtained within 20 km of Cothill (and North 
Hinksey). These moths are derived from larvae that are unlikely to 
have experienced food shortage when they start to feed immediately 
after hibernation. The Symphytum plants utilised by outsider larvae 
are surrounded by other vegetation suitable as larval foodplants, 
including Lamium spp. (Labiatae) and Urtica dioica (Urticaceae). It 
seems therefore that adult size is not so much a matter of geography 


287 


but rather is determined by inter-site variation in the availability of 
larval food in early spring. 


As already indicated, the outsiders probably originate from the main 
colonies, and that areas of countryside surrounding these colonies may 
periodically support small breeding populations which may or may 
not persist over a few years or more. It is not known what triggers 
moths to leave the colonies and breed elsewhere, but the possibility 
remains that it is associated with larval food shortage early in an 
individual’s life. But the most outstanding problem in C. dominula’s 
ecology is why the main colonies occur in such small and specialised 
areas, and why even within these areas the moths rarely expand into 
all the apparently suitable habitat, even though they can breed elsewhere 
in a variety of totally different habitats. 


Acknowledgements 


I thank David A.S. Smith for useful discussion and advice and Derek Whiteley 
for drawing Figs 1 and 2. My research on C. dominula is supported by a 
grant from the Cockayne Trust administered by the Natural History Museum, 
London. 


References 


Cook, L. M., 1961. Influence of larval environment on adult size and fecundity 
in the moth Panaxia dominula L. Nature 192 : 282. 

Cook, L. M., 1962. Some observations on the condition of over-wintering 
larvae of the scarlet tiger moth, Panaxia dominula (Linnaeus). Ento- 
mologist 95 : 47-50. 

Forp, E. B., 1975. Ecological genetics. 4th ed. Methuen, London. 

Jones, D. A., 1989. 50 years of studying the scarlet tiger moth. Trends Ecol. 
Evol. 4 : 298-301. 

Owen, D. F., 1994. Increase in larval foodplant diversity during a population 
explosion of the moth, Panaxia dominula (L.) (Lepidoptera : Arctiidae). 
Nota lepid. 16 : 267-273. 

SHEPPARD, P. M. & Cook, L. M., 1962. The manifold effect of the medionigra 
gene of the moth, Panaxia dominula, and the maintenance of a poly- 
morphism. Heredity 17 : 415-426. 


288 


Nota lepid. 18 (3/4) : 289-296 ; 13.V.1996 ISSN 0342-7536 


Zur Identität von Epermenia falciformis 
(Haworth, 1828) (Lepidoptera : Epermeniidae) 


Axel SCHOLZ 


Heerstrasse 10b, D-89269 Vöhringen-Illerberg, Deutschland 


Summary 


Epermenia falciformis (Haw.) is treated as a species separate from E. illigerella 
(Hübn.), with which it has previously been synonymised. The adults of E. 
falciformis and the closely related E. illigerella and E. petrusella (Heyl.) are 
illustrated and compared. The male and female genitalia of E. falciformis 
are figured and the biology and distribution is outlined. In order to stabilise 
the nomenclature a neotype of E. illigerella is designated. 


Zusammenfassung 


Epermenia falciformis (Haw.), eine bislang mit E. illigerella (Hübn.) synony- 
misierte Art, wird als eigenständiges Taxon erkannt. Die Falter von E. falci- 
formis und den nah verwandten E. illigerella und E. petrusella (Heyl.) werden 
abgebildet und verglichen. Die männlichen und weiblichen Genitalien von E. 
falciformis werden dargestellt und die Biologie und Verbreitung behandelt. 
Im Interesse der nomenklatorischen Stabilität wird ein Neotypus von E. 
illigerella festgelegt. 


Resume 


Epermenia falciformis (Haw.), jusqu’à présent considérée comme synonyme 
de E. illigerella (Hüb.) est reconnue comme taxon autonome. L'auteur présente 
des figures de E. falciformis et des espèces proches E. illigerella et petrusella 
(Heyl.) et les compare. Illustrations des genitalia mâles et femelles de E. falci- 
formis ; exposé de sa biologie et de sa répartition. Pour stabiliser la nomen- 
clature, l’auteur fixe un néotype de E. illigerella. 


Einleitung 
Schon seit Jahren steckten in der Sammlung des Autors zwischen einer 
Serie gezüchteter Tiere von Epermenia illigerella (Hübner, [1813]) einige 


gefangene, der genannten Art habituell sehr ähnliche Exemplare. Die 
vorgenommenen Genitaluntersuchungen ergaben tiberraschenderweise, 


289 


daß die fragliche Art in den ¢ Genitalien (!) Epermenia petrusella (Hey- 
laerts, 1883) nahesteht. Um eine Konspezifität auszuschließen wurde 
Vergleichsmaterial von petrusella mit dem Ergebnis untersucht, daß eine 
weitere, zwischen illigerella und petrusella stehende Art existiert. 


Nach Durchsicht der einschlägigen Literatur Konnte schließlich eine voll- 
ständige Übereinstimmung der Genitalien der fraglichen Art mit den bei 
Pierce & Metcalfe (1935 : Taf. 51) für illigerella abgebildeten Genitalien 
festgestellt werden. Die Vermutung, daß illigerella sensu Pierce & Met- 
calfe mit der aus England beschriebenen, als Synonym zu ülligerella 
auct. angesehenen Epermenia falciformis (Haworth, 1828) identisch sein 
könnte, bestätigte sich durch das Auffinden des Haworth’schen Typus. 


Folglich wird Epermenia falciformis (Haworth, 1828) als „bona species“ 
in die Literatur wieder eingeführt, und eine Abgrenzung gegen die 
nahestehenden Arten Epermenia illigerella (Hübner, [1813]) und Eper- 
menia petrusella (Heylaerts, 1883) vorgenommen. 


Abkürzungen 


BMNH Natural History Museum, London 


DEI Deutsches Entomologisches Institut 

TLME Tiroler Landesmuseum Ferdinandeum, Innsbruck 
YM York Museum, England 

ZSM Zoologische Staatssammlung München 
Bemerkung 


Hübners als „Originalbeschreibung“ geltende Abbildung von C. [alo- 
tripis| illigerella ([1813]: Taf. 48, Fig. 333) läßt sich nicht eindeutig 
interpretieren und könnte auch als falciformis gedeutet werden (beide 
Taxa kommen bei Augsburg, woher Hübners Original möglicherweise 
stammte, vor). 


In seinem Supplement und Revision zu Hübners Werk gibt Herrich- 
Schäffer (1854 : 207-208) einen kurzen Kommentar zu Hübners Ab- 
bildung und eine Diagnose von illigerella (?). Zur Biologie bemerkt er: 
„... die Raupen nach Stainton im Mai zwischen den Blättern von Aego- 
podium podagraria“. Stainton (1854 : 234) seinerseits schreibt : ,,... the 


(1) Die 6 und ® Genitalien von E. illigerella (Hübn.) und E. petrusella (Heyl.) sind 
bei Gaedike (1966 : 664-665) abgebildet. 
(2) „982. Illigerella H. 333. Kenntlich, die Spitze der Vorderflügel nicht sichelförmig 


290 


larva feeding (according to Fischer) on Aegopodium podagraria in 
May“. Demnach lagen Herrich-Schäffer selbst keine gezüchteten Exem- 
plare vor, und es ist nicht zu ersehen, ob Herrich-Schäffer bei seiner 
Beschreibung die „echte“ illigerella oder falciformis vor sich hatte. 


Da Hiibners Material verschollen ist, wird im Interesse der nomenkla- 
torischen Stabilität und um künftige Fehldeutungen auszuschließen, 
gemäß ICZN (Artikel 75 : (a)(c)) ein Neotypus von Epermenia illigerella 
(Hübner, [1813]) festgelegt. Ausgewählt wird ein & mit folgender Etiket- 
tierung (deponiert in der Zoologischen Staatssammlung München) : 


„Neotypus &“ [gedrucktes rotes Etikett], ,,Epermenia (Calotripis) illi- 
gerella (Hübner, [1813]) Taf. 48, Fig. 333., desig. A. Scholz 1994“ [ge- 
drucktes, rotes Etikett], „Deutschland, Bayern, Augsburg-Lechauen, 
4.V1.1982, ex larva Aegopodium podagraria, A. Scholz“ [gedrucktes 
Etikett] (ZSM). 


Epermenia falciformis (Haworth, 1828) 


Recurvaria falciformis Haworth, 1828. Lep. Brit. 4 : 555. 


Fehlinterpretationen : (3) 
Epermenia illigerella (Hübner) ; Pierce & Metcalfe, 1935 : 83, Taf. 51 (¢9 
Genit.). 
Epermenia illigerella (Hübner) (part) ; Gaedike, 1966 : 664-665. 
Epermenia petrusella (Heylaerts) ; Prose, 1992 : 242. 
Epermenia petrusella (Heylaerts) ; Larsen, 1993 : 135-136, Fig. 5. 


Locus TYPICUS : England 

HoıoTypus @ : ,,falciformis“ [handgeschriebenes Etikett von Haworth] ; 
„Holotype“ [rundes Etikett]; „Recurvaria falciformis Haworth, teste 
K. Sattler, 1994“ [Etikett von Sattler] (YM). 


genug, die zwei Schuppenzähnchen am Innenrande nicht angegeben ; sie gehen aber 
sehr leicht verloren. 

Lutea, ferrugineo-nebulosa, alis anter. puncto disci ad 1/3 & 2/3, cilus ter fusco divisis. 
Ledergelb, Vorderflügel stark rostgelb wolkig ; in der Mittellängslinie bei 1/3 und 2/3 
je ein schwarzer Punkt, von letzterem ein brauner Schrägwisch zum Afterwinkel, an 
welchem die Franzen von drei eisengrauen Linien getheilt sind“. 

(3) Auf eine vollständige Behandlung aller Epermenia falciformis (Haworth, 1828) 
betreffenden Literaturzitate wird zugunsten der nachgeprüften und gesicherten Angaben 
verzichtet. 

Alle Literaturangaben über ein Vorkommen von Epermenia illigerella (Hübner, [1813]) 
in England beziehen sich hôchstwahrscheinlich auf Epermenia falciformis (Haworth, 
1828) (Sattler, in litt.). 


21 


Diagnose 


IMAGo (Abb. 2, 4): Spannweite 14-l6mm. Fühler grau, lehmgelb ge- 
zähnt, Wurzelglied lehmgelb, Palpen lehmgelb, Endglied schwarzbraun 
beschuppt ; Kopf, Thorax und Tegulae lehmgelb, Stirn blaBgelb ; Beine 
schwarz beschuppt, Innenseite blaßgelb ; Abdomen dunkelgrau. Vor- 
derflügel : Grundfarbe blaßgelb, lehmgelb durchmischt ; Wurzelfeld 
blaßgelb, am Costalrand lehmgelb und schwarz beschuppt ; Dorsalrand 
mit zwei schwarzen Schuppenzähnen, der erste größere bei 1/3, der 
zweite bei 1/2 des Dorsalrandes ; knapp vor dem ersten Schuppenzahn 
zieht eine schwarze, distal lehmgelb begrenzte Binde schräg zu einem 
schwarz beschuppten, meist den Costalrand erreichenden trapezoiden 
Fleck ; an der dorsodistalen Ecke dieses Flecks ein schwarzer, hell um- 
grenzter Discoidalpunkt, von dem eine gebogene schwarze Binde schräg 
zum Tornus bis in die Fransen und eine weitere schwarzbraune Binde 
zum Costalrand vor den Apex gehen ; der Costalrand vor dem Apex 
und der Apex selbst sind schwarz beschuppt, die Außenrandfransen 
unterhalb des Apex weißgelb, die schwarze subapicale Fransenlinie an 
dieser Stelle konkav verlaufend, so daß der Eindruck einer „sichel- 
förmigen“ Flügelspitze entsteht. Hinterflügel : einfarbig grau ; Fransen 
grau, basal heller, Apex schwarzbraun gefranst. 


E. falciformis ähnelt sehr illigerella (Abb. 1), bei falciformis sind die 
Schrägbinde vor dem ersten Schuppenzahn und der basale Teil des 
Schattenflecks immer schwarz begrenzt, bei i/ligerella höchstens dunkel- 
braun begrenzt ; der Apex und die subapicale Fransenlinie sind bei 
falciformis breit und tiefschwarz, bei illigerella schmäler und grau- 
schwarz. Im Gesamthabitus erscheint falciformis durch den Kontrast 
der bleichgelben Grundfarbe zur lehmgelben und schwarzen Beschup- 
pung lebhafter gezeichnet als i/ligerella. E. petrusella (Abb. 3) ist von 
falciformis und illigerella sofort am geraden Verlauf der subapicalen 
Fransenlinie und den einfarbigen Außenrandfransen (kein „sichelför- 
miger“ Apex) zu trennen. 


& GENITALIEN (Abb. 5, 6) : Uncus lang, distal zugespitzt, basal konkav 
ausgeschnitten ; Tegumen breit, umgekehrt herzförmig ; Ampulle distal 
stark gebogen, zugespitzt ; Cucullus lang und schmal, Ampullenspitze 
überragend ; Sacculus basal breit, distal verschmälert, caudal Fortsatz 
schmal und spitz aufgebogen ; Juxta u-förmig, dorsolateral sehr stark 
verbreitert, Anellus etwa quadratisch, caudal beiderseits stark sklero- 
tisiert ; Aedoeagus etwa so lang wie die Valve, leicht gebogen, mit einem 
stark gefurchten, basal zugespitzten, distal stärker sklerotisierten Cor- 
nutus. 


297 


Abb. 1-3. Epermenia spp. 1 — E. illigerella (Hübn.) & ; 2 — E. falciformis (Haw.) 6 ; 
3 — E. petrusella (Heyl.) 9. 


Abb. 4-7. Epermenia falciformis (Haw.). 4 — Imago, Flügelpaar vergrößert ; 5 — 6 
Genital ; 6 — idem, Aedoeagus ; 7 — Q Genital (Spitze des Signums nachgezeichnet !). 


E. falciformis ist von illigerella leicht durch den schmalen, distal zu- 
gespitzten Uncus zu unterscheiden, bei illigerella ist der Uncus löffel- 
artig verbreitert ; meist reicht ein Abpinseln des Abdomens aus. Die 
Genitalien von falciformis und petrusella differieren wenig, sichere Unter- 
schiede liegen im Bau des Uncus, der bei falciformis etwa gleichbreit, 
bei petrusella länger und medial verschmälert ist ; der Sacculus is bei 
falciformis basal breit mit schmalem, spitz aufgebogenem Caudalfort- 
satz, bei petrusella basal schmal mit kurzem, stumpfen Caudalfortsatz, 
der Cornutus im Aedoeagus bei falciformis breiter und stark gefurcht, 
bei petrusella schmäler mit einer angedeuteten Furche. 


® GENITALIEN (Abb. 7): Papillae anales länglich oval, membranös ; 
Apophyses posteriores etwas länger als Apophyses anteriores ; Apo- 
physes anteriores gegabelt, innere Schenkel den Caudalrand des VIII. 


\ 


294 


Tergits erreichend ; Caudalrand des VII. Sternits konkav, sklerotisiert ; 
Ostium bursae sklerotisiert, breit ringförmig, mit caudaler Ausbuch- 
tung ; Ductus bursae gleichbreit wie das Ostium, zur Bursa etwas ver- 
breitert und gefurcht ; Bursa oval, Signum eine ovale, an den Rändern 
unregelmäßige Platte mit einem spitzen, dornförmigen Fortsatz. 


E. falciformis ist sowohl von illigerella als auch von petrusella durch 
das breit ringformige Ostium, den breiten Ductus bursae sowie das 
anders gestaltete Signum zu unterscheiden. 


Biologie 


Ei, Raupe und Puppe unbeschrieben. Die Falter wurden in England 
(Devon) von Heckford mehrfach aus Angelica sylvestris (Fam. Um- 
belliferae) gezüchtet (Sattler, in litt.). Generationsfolge : 2 Generationen, 
erste von Mai bis Juli, zweite von August bis September. Habitate : 
Feuchte Gebiete im Flachland (Moore, FluBauen). 


Verbreitung 


Schweiz : Graubünden : Landquart (Sattler, in litt.). Österreich : Vorarl- 
berg : Feldkirch-Bangs (Huemer, in litt.). Deutschland : Bayern : Isar- 
gebiet, München ; Württemberg : Allgäu, Oberschwaben ; Thüringen ; 
Nordrhein-Westfalen : Moseltal, Düsseldorf. Holland : Eindhoven. Bel- 
gien: Prov. Luxembourg: Heinstert. England: Südl. Grafschaften : 
Cornwall, Devon, Hampshire, Berkshire, Surrey, Oxfordshire, Cam- 
bridgeshire, Norfolk. Wales : Pembrokeshire (Sattler, in litt.). 


Danksagung 


An erster Stelle möchte ich mich sehr herzlich bei Herrn Dr. R. Gaedike 
(DEI) für seine Hilfe und Unterstützung in allen Fragen, Ausleihe von Material 
und Literaturbeschaffung bedanken. Herr Dr. K. Sattler (BMNH), der das 
Problem in England untersuchte, stellte freundlicherweise seine gesamten 
Ergebnisse zur Verfügung. Mr. K.R. Tuck (BMNH) und Mr. M. Denton (YM) 
ermöglichten durch ihre Hilfe das Auffinden des Haworth’schen Typus. Mit 
ausgeliehenem Material bzw. Informationen unterstützten mich die Herren 
W. Biesenbaum (Velbert-Langenberg), Dr. T. Grünewald (Landshut), S. Koster 
(Callantsoog), W. Pröse (Hof) und Dr. P. Huemer (TLMF). Ganz besonderen 
Dank schulde ich Herrn R. Sutter (Bitterfeld) und A. Lingenhöle (Rißegg) 
für Fotoarbeiten. 


295 


Literatur 


GAEDIKE, R., 1966a. Ergebnisse der Albanien Expedition 1961 des Deutschen 
Entomologischen Institutes. 53. Beitrag. (Lepidoptera : Epermeniidae). 
Beitr. Ent. 16 : 461-466. 

GAEDIKE, R., 1966b. Die Genitalien der europäischen Epermeniidae (Lepi- 
doptera : Epermeniidae). Beitr. Ent. 16 : 633-692. 

GAEDIKE, R., 1975. Beitrag zur Kenntnis der Mikrolepidopterenfauna der 
Balkanhalbinsel (Epermeniidae, Acrolepiidae, Douglasiidae). Beitr. Ent. 
25 : 221-226. 

GAEDIKE, R., 1993a. Nomenklatorische Bemerkungen zu den von H. T. 
Stainton 1851 beschriebenen Epermeniidae und Tineidae (Lepidoptera). 
Nota lepid. 15 (3/4) : 228-232. 

GAEDIKE, R., 1993b. Zur Kenntnis der Epermeniidae der Ostpaläarktis (Lepi- 
doptera). Nota lepid. 16 (2) : 91-104. 

Haworth, A. H., 1803-[28]. Lepidoptera Britannica XXXVI, 610 S. Londini. 

HERRICH-SCHAFFER, G. A. W., 1853-55. Systematische Bearbeitung der 
Schmetterlinge von Europa 5: 1-394; 1847-1854, ibidem 5 : Taf. 1-124, 
1-7, 1. Regensburg. 

HüÜBnER, J., 1796-[1836]. Sammlung europäischer Schmetterlinge 8: 78 S. 
(1796), 71 Taf. (1796-[1836]). Augsburg. 

HÜBNER, J., 1816-[26]. Verzeichnis bekannter Schmetterlinge. 431 S. Augsburg. 

International Commision on Zoological Nomenclature, 1985. International 
Code of Zoological Nomenclature. Third edition adopted by the XX 
General Assembly of the International Union of Biological Sciences. 
xx + 338 S. International Trust for Zoological Nomenclature, London. 

LARSEN, K., 1993. Some species of Microlepidoptera new to the Belgian fauna 
(Lepidoptera : Tortricidae & Epermenüdae). Phegea 21 (4) : 131-136. 

Pierce, F. N. & METCALFE, J. W., 1935. The genitalia of the tineid families 
of the Lepidoptera of the British Islands. xxıı + 116 S., 68 Taf. Oundle, 
Northants. 

Prose, H., 1992. Rote Liste gefährdeter Kleinschmetterlinge Bayerns. 
SchrReihe Bayer. Landesamt Umweltsch. 111 : 237-255. 

STAINTON, H. T., 1854. Insecta Britannica. Lepidoptera : Tineina. VIII, 313 S. 
10 Taf. London. 


296 


Nota lepid. 18 (3/4) : 297-303 ; 13.V.1996 ISSN 0342-7536 


Eupithecia veratraria Herrich-Schäfter, 1850 
new to Scandinavia, with the redescription 
of E. veratraria ssp. arctica Viidalepp, 1974 
(Lepidoptera, Geometridae) 


Toomas TAMMARU*, Tarmo VIRTANEN*, Kai RUOHOMAKI* & Lauri 
KAILA** 


* Laboratory of Ecological Zoology, Department of Biology, and Kevo Subarctic Research 
Institute, University of Turku, FIN-20500 Turku, Finland 
** Zoological Museum, PB 17, FIN-00014 University of Helsinki, Finland 


Summary 


The occurrence of Eupithecia veratraria H.-S. in Fennoscandia is confirmed. 
Three populations of this species, one of them abundant, were found in Tana 
and Berlevag communes, northernmost Norway. The larvae feed on Veratrum 
album L., the only known host plant of the species. Compared to central 
European specimens, those from Norway are characterised by an indistinct 
wing pattern and smaller size. The Norwegian population is considered to 
belong to ssp. arctica Viidalepp, described from the Polar Urals. E. v. arctica 
is redescribed on the basis of new material from the type locality. 


Zusammenfassung 


Das Vorkommen von Zupithecia veratraria H.-S. in Fennoskandien konnte 
bestätigt werden. Drei Populationen dieser Art, eine davon zahlreich, wurden 
in den Gemeinden Tana und Berlevag im nördlichsten Norwegen gefunden. 
Die Raupen leben an Veratrum album L., der einzigen bekannten Futterpflanze 
dieser Art. Im Vergleich mit zentraleuropäischen Tieren sind die Falter aus 
Norwegen kleiner und haben eine unscharfe Zeichnung. Die norwegische 
Population wird als ssp. arctica Viidalepp angesehen, eine aus dem nördlichen 
Ural beschriebene Unterart. E. v. arctica wird auf Grund neuer Materialien 
aus dem Typenfundort nochmals beschrieben. 


Resume 


Les auteurs confirment la présence d’Eupithecia veratraria H.-S. en Fenno- 
scandie. Ils ont trouvé trois populations de cette espèce, l’une d’entre elles 
abondante, dans les communes de Tana et Berlevag, dans l’extrême nord de 
la Norvège. Les chenilles se nourrissent de Veratrum album, seule plante- 
hôte connue de cette espèce. Comparativement aux exemplaires d’Europe 


297 


centrale, les veratraria de Norvège se caractérisent par un dessin flou et une 
taille plus petite. On considère la population de Norvège comme appartenant 
à la ssp. arctica Vüdalepp d’€crite de la partie polaire des Monts Oural. Re- 
description de la ssp. arctica sur la base du nouveau materiel de la localité- 


type. 
Introduction 


Eupithecia veratraria Herrich-Schäffer, 1850, is a transpalaearctic 
species which displays an alpine distribution throughout the mountains 
of central Europe. Its occurrence in northern Europe westwards from 
Russia has been disputed for a long time. E. veratraria was reported 
from northern Norway by earlier sources (Schneider, 1893 ; Haanshus, 
1933; Juul, 1948). Later, these records were considered doubtful 
(Knaben, 1976) and recent faunistic reviews (Skou, 1984 ; Mikkola et al. 
1989) did not consider the species as belonging to the fauna of the 
Fennoscandian countries. E. veratraria has repeatedly been mentioned 
also from the Baltic countries (e.g. Sulcs & Viidalepp, 1972). However, 
these reports have turned out to be based on misidentifications (Sulcs 
et al., 1981). 


Closest to Fennoscandia, E. veratraria is known to occur in the Ural 
Mountains (Vidalepp, 1974 and pers. comm.). In addition, one 
confirmed specimen was collected at Ponoi in eastern Kola peninsula 
(Fig. 1, D) in 1899 (Kozlov & Jalava, 1994). Veratrum album L. (Lilia- 
ceae), the only known host plant of E. veratraria, occurs continuously 
from the Polar Urals westwards to Arctic Norway (Hultén & Fries, 
1986). The record from Kola indicates that E. veratraria probably 
follows the range of its host plant throughout north-eastern Europe. 
Its occurrence in northernmost Norway could therefore be expected. 


New records from Norway (Fig. 1) 


In August 1993 three larvae of this species were found in an 
inflorescence of V. album close to Vestertana (70°26’N, 27°50’E) in 
northernmost Norway. Further records were made in September 1993, 
when hundreds of larvae of E. veratraria were collected at Leirpoll- 
skogen (70°27N, 28°40’E). Some larvae were also found at Store 
Molvik (70°47’N, 28°45’E). On 20th July 1994, 27 adults (8 64, 1999) 
were collected at Leirpollskogen where also naturally laid eggs were 
found. On 22nd July, 11 more specimens (6 @@, 599) were collected 
at the same locality and 3 (1 4, 299) specimens in the vicinity of 
Vestertana (J. Pöyry leg.). 


298 


Fig. 1. Records of Eupithecia veratraria (dots) and the main distribution of its host plant 
Veratrum album (shaded) in northernmost Europe. Localities mentioned in the text : 
A — Vestertana, B — Leirpollskogen, C — Store Molvik, D — Ponoi. 


At Leirpollskogen and Vestertana, the species was found in luxuriant 
mountain birch (Betula pubescens tortuosa (Ledeb.) Nyman) forests 
and on half-open pastures where V. album grows along streams ; Store 
Molvik is situated north of the timberline, V. album grows on river 
banks at a coastal settlement there. All localities are situated at low 
altitudes (0 - 50 m above sea level). E. veratraria was not found at higher 
altitudes in spite of the abundant occurrence of the host plant there. 


Main subspecies of E. veratraria, and the subspecific status 
of the Norwegian population 


Since the eastern subspecies of Æ. veratraria are little known, it is 
pertinent to give a short review of the subspecific division of this species. 


The nominate subspecies E. veratraria veratraria Herrich-Schäffer, 1850 
from mountains of central Europe is characterised by a uniform ground 


299 


colour and a distinct wing pattern consisting of a black discal spot, 
black vein streaks and scattered white markings (Fig. 2). 


All southern Siberian populations from the Altai mountains to the 
Pacific are considered to belong to E. veratraria homophaea Dyakonov, 
1926 (Mironov, 1991). The moths are smaller and have a more irregular 
fasciate wing pattern than the nominate subspecies, and the black vein 
streaks, typical for E. v. veratraria, are absent. The discal spot may 
be weak or absent. Terminal and costal parts of the forewing are usually 
darkened, the latter sometimes reddish brown. 


E. veratraria arctica Viidalepp, 1974 was described from northern parts 
of the Ural Mts. as a small subspecies with indistinct wing pattern 
(Viidalepp, 1974). Since the original description was based on only 
2 specimens, and the range of variation was unknown, we give a re- 
description of it on the basis of new material from the type locality. 


Eupithecia veratraria ssp. arctica Viidalepp, 1974 


REDESCRIPTION : Wingspan 18 - 22 mm (males), two females 17 and 
24 mm. Forewing leaden grey with weak yellowish tinge. Wing pattern 


Fig. 2. Eupithecia veratraria. Upper row: ssp. arctica, Tana, Norway, 46699; middle 
row : ssp. arctica, Polar Urals, 669Q; lower row : ssp. homophaea, Primorye region, 
OQ ; ssp. veratraria, central Europe, é@. 


300 


variable, markings highly diffuse with little contrast except small discal 
spot. Distally often unicolorous, darker grey, subterminal fascia weak 
or absent. Costal area usually darkened, but never brownish. Cilia 
weakly spotted. Hindwing as forewing, discal spot sometimes absent. 


The moths lack the sharply traced dentate fasciae typical for E. v. 
homophaea as well as the spotty pattern of the nominate ssp. No 
differences in the structure of either male (including examination of 
everted vesica) or female genitalia were found between FE. v. arctica 
and other subspecies. 


The wild-caught specimens from Norway closely resemble the moths 
from the type locality of E. v. arctica in their external appearance 
(Fig. 2). However, the Norwegian specimens seem to be more variable, 
including specimens with a clearly fasciate wing pattern. Characteristic 
traits of the Norwegian population are unicolorous cilia and no 
darkening of the costal region of the forewings. No differences in genital 
structure were found. Many reared Norwegian individuals, but none 
wild-caught from the same locality, resembled the ssp. veratraria by 
having a uniorm ground colour and well-developed subterminal fasciae. 
The genetic background of some of the subspecific differences can there- 
fore be questioned. 


As the differences between the Ural and Norwegian populations are, 
in Our Opinion, minor, we consider the Norwegian population as. be- 
longing to E. veratraria arctica. The view is supported by the continuous 
distribution of Veratrum album, and therefore most probably also of 
E. veratraria, from the Urals to Arctic Norway. 


MATERIAL EXAMINED 


ssp. veratraria — Central Europe : 9 specimens labelled “Dietze” ; 2 4@ label- 
led “Wien, Isaak” (ZBI) ; 1 & labelled “Bohemia” and “Staudinger” (ZMH) ; 


ssp. homophaea — Russia : Altai Mts., 12 specimens (ZMH) ; Primorye region, 
14 ; Khabarovsk region, 7 ; Kemerovo region, | (ZBI) ; 


ssp. arctica — Russia: Polar Urals, Krasnyi Kamen (66°53’N 65° 10’E) 
16.07.1969. (in the original description erroneously 16.05.), 1 4, holotype, 
(ZBI) ; ibidem, 15.-16.07.1994, 2 33, J. Kullberg, J. Jalava and S. Koponen 
leg. ; Ob delta, 8 km N Labytnangi (66° 42’N 66°35’E), 27 44, 2 ©, J. Kullberg, 
J. Jalava and S. Koponen leg. (ZMH); Kola peninsula, Ponoi, 1899, 1 spe- 
cimen (ZMH); Norway: Berlevag commune, autumn 1993, 1 © ex larva; 
Tana commune, autumn 1993, 37 specimens ex larva ; ibidem, 41 specimens, 
20.-22.07.1994 (ZBI, ZMH, authors’ and J. Pöyry’s private collections). 


301 


ABBREVIATIONS 


ZBI Institute of Zoology and Botany of the Estonian Academy of Sciences, 
Tartu, Estonia | 

ZMH Zoological Museum, Finnish Museum of Natural History, Helsinki, 
Finland. 


Description of the preimaginal stages and notes on ecology © 


At Leirpollskogen, eggs of E. veratraria were found singly on stamens 
of V album. They were about 0.5 mm long and 0.3 mm wide, and 
varied from white to orange in colour (probably, due to different age). 
In some cases, there were up to three eggs per flower of V. album. 


In late August — early September 1993 most of the larvae collected 
were full-grown and pupated within a few days in the laboratory. They 
were dark brown to black in colour, without any pattern, and up to 
12mm long. The larvae match perfectly with the description of E. 
veratraria larvae from Central Europe (Dietze, 1913 ; Weigt, 1990), 
both in appearance and feeding habits. About 10% of the larvae were 
parasitised by Campoletis rectangulata (Aubert, 1977) (Hymenoptera, 
Ichneumonidae). This species was hitherto known only from the Alps 
(R. Jussila, pers. comm.). 


Larvae pupated in soil within a thin but tough cocoon. The dark yellow 
pupae were about 7-8 mm long. After the first hibernation, less than 
10% of pupae gave adults and more eclosed after the second winter. 
This suggests a strong inclination of Norwegian E. veratraria pupae 
for multiple hibernations, as is characteristic for central European 
populations (Dietze, 1913). We would expect a strong natural selection 
for such an emergence pattern since V. album does not bear flowers 
every year. 


Most adults collected 20th-22nd July 1994 were in worn condition and 
the sex ratio was female-biased. We conclude that the peak flight was 
one or two weeks earlier, E. veratraria is thus a species of subarctic 
midsummer. The adults were observed in flight during the evening 
(about 18.30 - 19.30 local time) but not at noon, and they were only 
found in the close vicinity of their host plant. They were observed 
feeding on flowers of V. album. 


Acknowledgements 


We are most thankful to J. Poyry for putting his data at our disposal as well 
as to M. Kozlov, J. Pöyry and J. Viidalepp for comments on the manuscript. 


302 


L. Aarvik and C. Eliasson helped to find the Norwegian references. The 
parasitoid could not have been determined without the kind help of R. Jussila. 


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Staats-Universität zu Tartu (Lepidoptera, Geometridae). TRÜ Toimetised 
327 : 86-93. (in Russian, with German summary). 

WEIGT, H.-J., 1990. Die Blütenspanner Mitteleuropas. (Lepidoptera, Geome- 
tridae : Eupithecuni). Teil 3 : Eupithecia sinuosaria bis pernotata. Dort- 
mund. Beitr. Landeskde 24 : 5-100. 


303 


Nota lepid. 18 (3/4) : 304-305 ; 13.V.1996 ISSN 0342-7536 


Short communication — Kurze Mitteilung — En bref 


Rubiginous larvae of Hyles hippophaes (Esper, 1793), an autosomal 
recessive variety (Lepidoptera : Sphingidae) 


E. A. LOELIGER 


Hofdijck 48, NL-2341 ND Oegstgeest, The Netherlands 


Aberrant colouration of the caterpillar of Hyles hippophaes (Esper, 1793) has 
so far only been reported by Pittaway (1993), who mentions that “in a very 
rare colour form, all green colouration is replaced by pinkish-brown”. Due 
to the lack of further information, the following observations seem to be worth 
reporting. 


In 1982, a rubiginous-coloured young L; caterpillar of H. hippophaes, was 
observed openly sunning itself on a twig of Hippophae rhamnoides, near the 
Swiss village of Susten, in the Rhône Valley (Fig. la). The caterpillar was 
highly conspicuous against the olive-green coloured plant. The caterpillar and 
pupa developed normally. A normal male emerged in 1983 and mated with 
a female originating from an olive-green larva which had been found in the 
same region. The resulting F, larvae were all olive-green, the pupae well- 
shaped. From the subitaneous pupae a first F, was obtained in the same 
year, consisting of 30 L;, of which 6 were rubiginous. A second F, was not 
achieved until 1985, because many of the F, pupae remained in diapause 
for more than one year. Rearing this time outdoors on Eleagnus angustifolia, 
produced almost three times as many L; were obtained, 28.5% (23/80) being 
of the rubiginous variety (all instars, but most obvious in L;), a percentage 
which is consistent with the hypothesis of a hereditary, recessively transmitted 
colour variety. The crucial F3 was finally raised in 1986, both parents coming 
from rubiginous larvae. One hundred percent of the caterpillars (resulting in 
24 male and 22 female pupae) were rubiginous, paler when reared on Eleagnus 
angustifolia (Fig. 1b) and darker on Dutch Hippophae rhamnoides (Fig. 1c), 
confirming the aforementioned hypothesis of an autosomal recessive trait. 


Since the lower surface of the leaves of many Hippophae rhamnoides plants 
is also rubiginous, we believe that the gene responsible for this larval variety 
of hippophaes is adaptive rather than being a mutation caused by, for instance, 
toxic pollution of the atmosphere by the chemical industry in the region. 


References 


Pitraway, A. R., 1993. The Hawkmoths of the Western Palaearctic. 240 pp. Harley 
Books, Colchester. 


304 


Fig. 1. Larvae of H. hippophaes. a — Propositus larva on Hippophae rhamnoides, 
Susten, VS, Switzerland, 1982 ; b — Rubiginous F, larva on Eleagnus angustifolia ; 
c — Larva of the F3 generation on Dutch Hippophae rhamnoides. Colouration 
approximating that of the dark rubiginous lower surface of the leaves. 


Nota lepid. 18 (3/4) : 306-307 ; 13.V.1996 ISSN 0342-7536 


Short communication — Kurze Mitteilung — En bref 


Additional experiments to unravel the enigma of Hyles hybrid pauli 
Mory (Lepidoptera : Sphingidae) 


E. A. LOELIGER 


Hofdijck 48, NL-2341 ND Oegstgeest, The Netherlands 


The origin of Hyles hybrid pauli Mory (Mory, 1901) remained enigmatic even 
after extensive experiments with primary and secondary hybrids from A. 
euphorbiae euphorbiae Linn. and H. hippophaes hippophaes Esp. (Loeliger, 
1976). Recently, we tested the hypothesis, expressed on repeated occasions 
(Denso, 1909 ; John, 1932 ; Fischer, 1933), that Hyles hybrid pauli Mory might 
be a cross between a male Hyles hybrid fischeri John and a female H. hippo- 
phaes hippophaes Esp. 


H. hybr. fischeri, which is the cross of a male A. livornica livornica Esp. 
with a female H. hippophaes hippophaes Esp., was obtained in 1993. The 
parents of the livornica male had been collected in the spring in southern 
Turkey (courtesy Dr. ©. Kurz, Münster, D) and the parents of the hippophaes 
female were the offspring of the mating of a male of the caucasica variety 
found on E. angustifolia in Anatolia and a female of the “ordinary” hippophaes 
variety (courtesy F. Weber, Riehen, CH) (Loeliger, in prep.). 


The crucial cross between a fischeri male and hippophaes female, obtained 
in the early summer of 1994, resulted in no more than two eggs, one of which 
appeared to undergo embryogenesis (H. Harbich, Salz, D). However, the 
caterpillar did not hatch. Of the 22 eggs deposited by a female A. tithymali 
tithymali Bds (courtesy Dr. O. Kurz, Miinster, D) after mating with a fischeri 
male, none displayed signs of developing into a larva. In contrast, two of 
the 23 eggs resulting from the mating of two fischeri specimens seemed to 
be fertilized (observation H. Harbich, Salz, D) ; however, the caterpillars never 
hatched. Finally, the many hundreds of eggs deposited on vine leaves by two 
fischeri females after successful mating with hippophaes males were sterile. 


This infertility of H. hybr. fischeri is probably due to the tiny size of its eggs 
and to the extreme size difference to the eggs of H. hippophaes. The eggs of 
fischeri are distinctly smaller than those of /ivornica (roughly 1:4) and are only 
about one sixteenth the size of hippophaes eggs (pers. comm. Dr. F. Karrer, 
Zofingen, CH). 


In conclusion, evidence in favour of the above-mentioned hypothesis is meager, 
although not nil. Experiments should be repeated. We suspect, however, that 


306 


the H. hybr. pauli Mory specimen is an artefact consisting of parts of two 
sphingid specimens glued together, the abdomen (and antennae ?) belonging 
to a hippophaes female and the rest coming from a specimen containing 
tithymali and / or dahlii genes. This would also explain the conspicuous colour 
difference between the two parts, hippophaes being olive-green and the other 
species rubiginous brownish. 


The hypothesis that the father of A. hybr. pauli Mory was a stray H. dahlii 
Gey. (Kysela, 1908) is very unlikely. Still another hypothesis, i.e. that the 
very strange appearance is the result of the mergence of atavistic properties 
(Wladasch, 1939; 1943), is also improbable since we have never observed 
such conspicuous discrepancies in marking and colouration among countless 
hybrids of the Hy/es genus. 


Acknowledgements 


We would like to thank all the above mentioned colleagues for the excellent cooperation 
in our endeavour to unravel this everlasting enigma. 


References 


Denso, P., 1911. Celerio hybr. hippophaës & * euphorbiae Q und euphorbiae & x 
hippophaës 9. Ent. Z. 25 : 151-153. 

FISCHER, E., 1933. Ein sekundärer /ivornica-Hybrid und über Kennzeichen von Cel. 
hybr. pauli und gillmeri. Int. ent. Z. Guben 27 : 209-213. 

JoHn, K., 1932. Ein weiterer neuer Celerio-Hybrid. Ent. Z. 46 : 161-162. 

KYSELA, E., 1908. Beitrag zur Erkenntnis der Deilephila-Hybriden. Mitt. ent. Ver. 
Polyxena 2 : 63-82. 

LOELIGER, E. A., 1976. The enigma of Celerio hybr. Pauli Mory. Tijdschr. ent. 119 : 
217.219. 
LOELIGER, E. A., in prep. Breeding of an F2 of Hyles hybr. vespertilioides Boisduval 
after induction with ecdysone of metamorphosis in diapausal female pupae. 
Mory, E., 1901. Ueber einige neue schweizerische Bastarde des Sphingiden-Genus 
Deilephila und die Entdeckung abgeleiteter Hybriden in der Natur, sowie Be- 
schreibung einer neuen varietät von Deilephila vespertilio Esp. Mitt. schweiz. 
ent. Ges. 10 : 333-360. 

WLADASCH, E., 1939. Eine atavistische Rückschlagsform von Celerio euphorbiae im 
Vergleich mit C. dahlii, C. tithymali und Cel. hybr. pauli und den dunklen Formen 
von Celerio euphorbiae L. Ent. Rdsch. 56 : 224-227 ; 303-304 ; 331-332. 

WLADASCH, E., 1943. Ueber die vermutliche Abstammung des Celerio hybr. pauli 
Mory. Z. wien. ent. Ges. 28 : 25-43. 


307 


Nota lepid. 18 (3/4) : 308-309 ; 13.V.1996 ISSN 0342-7536 


Book reviews — Buchbesprechungen — Analyses 


Swallowtail Butterflies of the Americas. A Study in Biological Dyna- 
mics, Ecological Diversity, Biosystematics and Conservation. Hamilton 
A. TYLER, Keith S. Brown, Jr. & Kent H. Wizson. 376 Seiten, zahl- 
reiche Abbildungen und Farbtafeln. 22 X 28 cm. Scientific Publishers : 
Gainesville, Florida 32604, USA, 1994. ISBN : 0-945417-90-X (gebun- 
den), 0-945417-91-8 (kartonierte Studentenausgabe). Preis : $ 49.50 (ge- 
bunden), $ 24.50 (kartoniert). 


Dies ist ein echtes Arbeitsbuch! In den zwölf einleitenden Kapiteln werden 
(teils frei übersetzt) Schwalbenschwänze in Kultur und Natur ; Ökologie und 
Verhalten ; Populationsbiologie der Imagines ; Jugendstadien und Futterpflan- 
zen ; Chemie der Futterpflanzen ; Mimikry ; Genetik und Hybridisierung ; 
Schutz der Arten und ihrer Lebensräume ; Biogeographie ; Systematik, Evolu- 
tion und Phylogenie ; Merkmale zur Klassifizierung und Diversität und Bio- 
systematik abgehandelt. Daran schließt sich ein „systematischer Teil“ an, der 
alle 143 Arten der Region auf Farbtafeln und mit Verbreitungskarten vorstellt. 
Die Systematik wird erfrischend nüchtern und distanziert abgehandelt und 
auf den ihr zukommenden Rang verwiesen : notwendig, aber nicht Ziel. 


Im Vergleich zu den „Swallowtail Butterflies“ von Scriber et al. (s.u.) bemühen 
sich die Autoren nicht, einzelne wenige Aspekte möglichst abgerundet darzu- 
stellen. Vielmehr versuchen sie, auch an ungelöste Fragen heranzuführen und 
den Leser selbst zur Formulierung (und Beantwortung) von Fragen zu ani- 
mieren. Die parallele Studentenausgabe ist also kein Zufall. 


Es ist schwer, bei der gebotenen Fülle einzelne Aspekte herauszugreifen. Mir 
hat besonders gefallen, mit welchem Nachdruck die Jugendstadien gleichbe- 
rechtigt neben die Imagines gestellt werden. Immerhin werden die „frühen 
Stände“ von 105 Arten „komplett“ oder wenigstens teilweise abgebildet, etliche 
davon erstmalig. 


Die verschiedenen Informationspakete sind z.T. reichlich chaotisch verteilt. 
So tauchen z.B. die Tafeln 67, 70 und 72 mitten im Bestimmungsteil an Stellen 
auf, mit denen sie absolut nichts zu tun haben. Auf diese Weise entsteht ein 
skizzenhafter Gesamteindruck, der vielleicht sogar beabsichtigt ist ; zumindest 
zwingt er zum Nachschlagen. 


Schade allerdings, daß die Herausgeber offenbar keinen sonderlichen Wert auf 
die Qualität der Abbildungen gelegt haben, die daher extrem heterogen sind. 
Vor allem auf den ersten 48 (von insgesamt 100) Farbtafeln stören mich zudem 
das oft allzu winzige Format der Einzelfotos und ihre rücksichtslose „Zu- 
sammenstückelung“. Dies überrascht mich vor allem deshalb, weil der Bedeu- 
tung ästhetischer Aspekte beinahe das ganze erste Kapitel gewidmet ist. 


308 


Dies kann den Wert des Werkes insgesamt jedoch nicht schmälern. Es bietet 
seine enorme, dicht gepackte Informationsfülle leicht lesbar in jenem selbst- 
ironischen Stil, der dem europäischen Fachwissenschaftler so schwer aus der 
Feder kommt. Und das alles zu einem äußerst günstigen Preis — sofern man 
nicht die signierte Kunstlederausgabe bevorzugt (ISBN 0-945417-92-6, Preis 
$ 150.00). 


Alexander PELZER 


Swallowtail Butterflies : Their Ecology and Evolutionary Biology. J. 
Mark SCRIBER, Yoshitaka TsuBAK1I & Robert C. LEDERHOUSE (Hrsg.). 
459 Seiten, zahlreiche Abbildungen und 32 Farbtafeln. 22 X 28 cm, 
gebunden. Scientific Publishers : Gainesville, Florida 32604, USA, 1995. 
ISBN : 0-945417-89-6. Preis : $ 65.00. 


Das Buch basiert auf den Ergebnissen eines Kongresses in Yokohama (Japan). 
Die insgesamt 35 Einzelbeitrage sind auf die Sektionen Chemische Okologie 
und Verhaltensphysiologie, Lebensweisen und Populationsdynamik, Paarungs- 
biologie und Mimikry, Okologische Genetik und Evolution und Naturschutz 
und Erhaltung der Diversität aufgeteilt. 


Es verwundert nicht, daß sich inhaltliche Überschneidungen mit den Swallow- 
tail Butterflies of the Americas von TYLER et al. (s.o.) ergeben. Hier stehen 
jedoch die Phänomene und nicht die einzelnen Arten im Vordergrund, und 
neben den amerikanischen werden vor allem ostasiatische Arten behandelt. 
Europäische und afrikanische Arten sind hingegen deutlich geringer vertreten. 


Erklartes Ziel der Herausgeber ist eine monographische Tiefe bei den ange- 
schnittenen Themen, und dieses Ziel wird auch erreicht. Von Interesse ist das 
Werk damit nicht nur fiir Schwalbenschwanz-Enthusiasten, sondern ebenso 
als Einstiegs- oder Nachschlagewerk für alle, die sich mit ökologischen Frage- 
stellungen bei Insekten befassen. 


Ein Autor konnte offenbar der Versuchung nicht widerstehen, seinen eigenen 
Beitrag zu einem Fachgebiet überproportional darzustellen. Er zitiert sich auf 
1 Seiten fast 40 mal, wobei so gut wie alle Zitate „graue Literatur“, nämlich 
unveröffentlichte Berichte oder in Vorbereitung befindliche Arbeiten betreffen. 
Hier hätten die Herausgeber strenger sein können. 


Insgesamt jedoch ein sehr solider Band, der den aktuellen Wissensstand über 
viele Teilgebiete der Ökologie von Schwalbenschwänzen zusammenfassend 
darstellt. Bemerkenswert scheint mir die geringe Repräsentanz europäischer 
Wissenschaftler unter den Autoren. Liegt das wirklich nur daran, daß Europa 
so wenige Arten der Papilioniden „abbekommen“ hat ? 


Alexander PELZER 


P.S. : Als Europäer ist mir aufgefallen, daß Archon apollinus in beiden Büchern 
konsequent als A. apollinis erscheint. Auch Datenbanken haben ihre Tücken... 


309 


Nota lepid. 18 (3/4) : 310-312 ; 13.V.1996 ISSN 0342-7536 


Vol 18 — 1995 


Dates of publication — Publikationsdaten — Dates de publication 


18(1): 31.1.1996 pp. 1-92 
18 (2): 29.11.1996 pp. 93-180 
18 (3/4): 13.V.1996 pp. 181-312 


Contents — Inhalt — Sommaire 


BENYAMINI, D. : Pupal summer diapause in Chilean Pieris brassicae 
(:innaeusy 1759) (Pieridae)? re UIID 
CUPEDo, F. : Die morphologische Gliederung des Erebia melampus- 
Komplexes, nebst Beschreibung zweier neuer Unterarten: 
Erebia melampus semisudetica ssp.n. und Erebia sudetica 
belledonnae ssp.n. (Satyııdae) es 
DENNIS, R. L. H. : Oviposition in Zerynthia cretica (Rebel, 1904) : 
loading on leaves, shoots and plant patches (Papilionidae).... 

Dennis, R. L. H. — cf. SHREEVE, T. G. 

DRECHSEL, T. — cf. LOBEL, H. 

FAUCHEUX, M. J. : Sensilla on the ovipositor of the carpet moth, 
Trichophaga tapetzella 1; (Tineidae) 1... ei er 

FIBIGER, M., HACKER, H. & MoßBERG, A. : Notes on the Orthosia 
rorida (Frivaldsky, 1835) species group, with the description 
of a new species from Crete : Orthosia sellingi sp. n. (Noctui- 
dae, Hadeninae)®..........:: ec est oo geese ote osteo ee eee 

FIEDLER, K. — cf. SANETRA, M. 

GASKkIN, D. E.: Hesperioidea and Papilionoidea of the Ionian 
island of Kefalonia, Greece : Additional species and a review 
of faunal’ components"! oc tee See MEERE 

HACKER, H. — cf. FIBIGER, M. 

HATTENSCHWILER, P. : Dahlica wehrlii (Müller-Rutz, 1920) wieder 
gefunden. Beschreibung des Weibchens und Ergänzungen zur 
Kenntnis des Männchens und der Okologie (Psychidae) ...... 

HATTENSCHWILER, P.: Eine neue Montanima — Art aus dem 
Altai-Gebiet (Psychidae) nn... ence ee 

HATTENSCHWILER, P. : Eine neue, im Februar-März fliegende Pri/o- 
cephala — Art aus dem südlichen Spanien (Psychidae) ........ 

Hauser, E.: Vergleichende Analyse der Zönosen tagaktiver 
Schmetterlinge im Sengsengebirge (Oberösterreich). ............. 


310 


No. 


3/4 


3/4 


3/4 


3/4 


m 


184 


95 


193 


203 


218 


225 


233 


239 


247 


KaAILA, L. — cf. TAMMARU, T. 

Kozıov, M. V. : Subalpine and alpine assemblages of Lepidoptera 
in the surroundings of a powerful smelter on the Kola Penin- 
ELLE INT ROSE Ne ee a er er 

Kozıov, M. V. & Rosinson, G. S. : Identity and distribution of 
two dimorphic oriental fairy moths — Nemophora decisella 
(Walker, 1863) and Nemophora cantharites (Meyrick, 1928) 
TENTE ee ry el A ot ee 

LopeL, H. & DRECHSEL, T.: Zur Lebensweise der Raupe von 
Euchalcia emichi (Rogenhofer, 1873) in Zentralanatolien 
Mk) GNoctuidae, Plusinae) 70222 0.21. 

LOELIGER, E. A. : Rubiginous larvae of Hyles hippophaes (Esper, 
1793) an autosomal recessive variety (Sphingidae) ............... 

LOELIGER, E. A. : Additional experiments to unravel the enigma 
cles hybrid paul Mory (Sphingidae).........................- 

MAIER, C. & SHREEVE, T. G.: Endothermic heat production in 
PRES SPECIES OL NY Mp alid AC 0... 

MoBERG, A. — cf. FIBIGER, M. 

MoLiNA, J. M2. & PALMA, J. M.: Butterfly diversity and rarity 
within selected habitats of western Andalusia (Spain) (Papi- 
Iameideatand ELESMeMmOidea): 22.5 4.2 coo idee 

Owen, D. F.: Larval food shortage and adult dispersal in Calli- 
OC OM EN PME TS RER en. 

PASSERIN D’ENTREVES, P. : Révision des Scythrididae paléarctiques. 
VI. Les types de Scythrididae du Natural History Museum de 
Londres, du Muzeul de Historia Naturala «Grigore Antipa» 
de Bucarest et du Zoologisches Museum der Humboldt- 
Winnversitat de Berlin (Premiere partie)... 

RosInson, G. S. — cf. Kozıov, M. V. 

RUOHOMAÄRI, K. — cf. TAMMARU, T. 

SANETRA, M. & FIEDLER, K. : Behaviour and morphology of an 
aphytophagous lycaenid caterpillar: Cigaritis (Apharitis) 
EUROS Klug, 1834 (EV CACTI AE) nn. een 

SCHOLZ, A.: Zur Identität von Epermenia falciformis (Haworth, 
1.828) STONES RER Re 

SHREEVE, T. G., DENNIS, KR. L. H. & WırLıams, W. R.: Uni- 
formity of wing spotting of Maniola jurtina (L.) in relation 
to environmental heterogeneity (Satyrinae) ........................ 

SHREEVE, I. G. — cf. MAIER, C. 

TENNENT, J. : The distribution of the genus Tarucus Moore, [1881], 
in the Maghreb States of Morocco, Algeria and Tunisia, with 
notes on species identification (Lycaenidae) ......................... 


3/4 


17 


170 


304 


306 


127 


267 


281 


139 


57 


289 


eh 


161 


3 


TAMMARU, T., VIRTANEN, T., RUOHOMAKI, K. & Karna, L. : Eupi- 
thecia veratraria Herrich-Schäffer, 1850 new to Scandinavia, 
with the redescription of E. veratraria ssp. arctica Vüdalepp, 
1974 (Geometridac) 2.555 de Stas ee ee 3/4 
VIRTANEN, I. — cf. TAMMARU, T. 
WILLIAMS, W. R. — cf. SHREEVE, T. G. 


Obituary — Nekrolog — Nécrologie 


Dirk HAMBORG (1957-1995) Re RS PER 2 

Editorial... NARBE ELA ee ARE 3/4 
Book reviews — Buchbesprechungen — Analyses 

An Illustrated Key to European Sessidae ..................................... 3/4 

Die Schmetterlinge Baden-Württembergs. Band 3 & 4 .................. 2 

Die Zünslerfalter (Pyraloidea) Mitteleuropas ............................... l 

Swallowtail Butterflies of the Americas ........................................ 3/4 

Swallowtail Buttexflies. ..........200000.0...0 Re 3/4 

The Nepticulidae of eastern Europe and Asia ...................2...e0...... 2 

Notices... ee RE PAR ee l 

| 

2 

2 

3/4 

3/4 


New taxa described in Vol. 18 
Neue Taxa in Band 18 beschrieben 
Nouveaux taxa décrits dans le Vol. 18 


PSYCHIDAE 
Montanima aurea Hattenschwiler, 1996 .................................... 3/4 
Piilocephala piae Wattenschwilets, 1996 222.2... ee 3/4 
NOCTUIDAE 
Orthosia selling Tibiser 199... eee ee ee eee 3/4 
SATYRIDAE 
Erebia melampus semisudetica Cupedo, 1996 ............................... > 
Erebia sudetica belledonnae Cupedo, 196 7 nn en ee 2 


312 


297 


126 
183 


234 
241 


204 


116 
119 


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Nota lepidopterologica 


Vol. 19 No. 1/2 Basel, 21.X1.1996 ISSN 0342-7536 


Editor : Alain Olivier, Luitenant Lippenslaan 43 B14, B-2140 Antwerpen- 


Borgerhout, Belgium. 


Assistant Editors : Dr. Roger Dennis (Wilmslow, GB) 
PD Dr. Andreas Erhardt (Binningen, CH) 
Dr. Enrique Garcia-Barros (Madrid, E) 
Dr. Christoph Hauser (Stuttgart, D) 
Dr. Alexander Pelzer (Wennigsen, D) 
Dr. Erik J. van Nieukerken (Leiden, NL) 


Contents — Inhalt — Sommaire 


nn GP oat cake Se quete gant aunie das sehensssPadea 
HAUSMANN, A. : The morphology of the geometrid moths of the Levant 
and neighbouring countries. Part I : Orthostixinae and Geometrinae..... 
HAUSMANN, A. : Systematic list of the geometrid moths of the Levant 
and neighbouring countries. Part I : Orthostixinae and Geometrinae..... 
HATTENSCHWILER, P.: Sciopetris karsholti, eine neue Psychide aus 
mriesen(Eepidoptera!/Psychidae) Hi Hal 2 ne PHo nn. tunes 
LOELIGER, E. A. & KARRER, F.: On the induction of metamorphosis 
of Lepidoptera by means of ecdysone and 20-hydroxyecdysone ........ 
VANHOLDER, B. : The migration of Danaus plexippus (Linnaeus, 1758) 
during October 1995 in the UK (Lepidoptera: Nymphalidae, Da- 
NIEE) eos Mane! SUN Riad RS Sas Tee RS ANR a ESS 


Editorial 


With the present double issue, I start my first contribution as new 
Editor of Nota lepidopterologica. When it became known to me that 
my predecessor was about to resign, I proposed myself as a possible 
candidate for taking over his job and I became elected as Editor at the 
Xth European Congress of Lepidopterology Gale de la Sierra, 
Madrid, Spain, 3.-7.V.1996). 


In my own name as well as in name of all SEL-members, I wish to 
express my deep gratitude to Mr. Steven E. Whitebread for the excellent 
work he has offered us during about six years of editing. We can 
confirm that Steven certainly achieved his aim of improving the quality 
(content and style) of our journal. It is one of the most challenging 
aspects of my new job to maintain the same high standards he has 
attained. 


My greatest preoccupation so far has been to gather enough manu- 
scripts to be able to fill a complete volume and so to keep up with 
publication schedule by publishing the whole Vol. 19 still in 1996. In 
doing so, I have selected a series of manuscripts received from my 
predecessor as well as a few new ones, including one single article 
co-written by myself that, I hope, the readers will have the indulgence 
to appreciate. This took me a lot of time and energy as I had to do 
all the work on my very own (except for the bulk of the work on 
both Hausmann’s articles) and I hope the resulting present Volume 
will not be perceived as inferior to the preceding one. 


From Vol. 20 on, I will continue the editing in co-operation with a team 
of Assistant Editors. Of the people whom I contacted for this job so 
far, the following persons have accepted the offer : Dr. Roger Dennis, 
PD Dr. Andreas Erhardt, Dr. Enrique Garcia-Barros, Dr. Christoph 
Hauser, Dr. Alexander Pelzer and Dr. Erik J. van Nieukerken. 


In the future, I would like at least to try to achieve a regular publication 
schedule. 


We badly need more manuscripts. All authors who wish to publish in 
Nota are therefore kindly requested to send their manuscript right away 
to the Editor. I would like to ask expressly for contributions by micro- 
lepidopterists, as these are sadly lacking for the moment. 


Finally, I would like to emphasize that the future of both our Journal 
and our Society depend on the money we get through our membership 
fees. Those who have not paid their 1996 contribution yet are urged 
to do it now, without any further delay. 

Alain OLIVIER 


Nota lepid. 19 (1/2) : 3-90 ; 21.X1.1996 ISSN 0342-7536 


The morphology of the geometrid moths of the Levant 
and neighbouring countries 


Part I : Orthostixinae and Geometrinae 


Axel HAUSMANN 


Zoologische Staatssammlung, Münchhausenstr. 21, D-81247 München, Germany 


Summary 


This is the first of a series of articles on the Geometrid moths occurring in 
the Levant and neighbouring countries (Middle East). Morphological de- 
scriptions of 53 taxa at species or subspecies level are accompanied by 
illustrations of venation (for all genera) and of male and female genitalia 
(for all taxa). Differential diagnoses of genera and discussions on possible 
relationships are added. In a general revision of the systematic arrangement 
more than 100 genera are attributed to 19 Old World tribes or subfamilies. 
The validity of subfamily rank for Orthostixinae Meyrick, 1892 (Oenochro- 
minae auct. nec Guenée, 1857) is confirmed, including seven genera, and its 
differential features given. One new tribe is established : Microloxiini tribus 
n. with the type genus Microloxia Warren, 1893. Three new genera and one 
new subgenus are established : Microbaena gen. n. with type species Pho- 
rodesma pulchra Staudinger, 1897 ; Proteuchloris gen. n. with type species 
| Geometra (Phorodesma)| neriaria Herrich-Schaffer, 1852 ; Acidromodes gen. 
n. with type species Acidaliastis nilotica Wiltshire, 1985 ; Victorinella subgen. 
n. of Victoria Warren, 1897, with type species Victoria sematoperas Prout, 
1916. Three new species and three new subspecies are described : Thetidia 
persica sp. n. from N. Iran, Victoria wiltshirei sp. n. from the Sudan, Hemi- 
dromodes unicolorata sp. n. from Kenya, Pseudoterpna coronillaria halperini 
ssp. n. from Israel, Microbaena pulchra minor ssp. n. from the Sudan and 
Eucrostes indigenata lanjeronica ssp. n. from Spain. 


Further changes concern the following taxa : Archaeobalbini Viidalepp, 1981, 
emendation of Archeobalbini. Terpnini Inoue, 1961, homonym to Terpnini 
Kuznetsov, 1904 (= synonym to Geometrini Duponchel, [1845]). Terpnini 
sensu Inoue “synonym” (but unavailable) to Pseudoterpnini Warren, 1893. 
Neohipparchini Inoue, 1961, syn. n. to Geometrini Guenée, 1844. Ochrognesuni 
Inoue, 1961 syn. n. to Nemoriini Gumppenberg, 1887. Nemoriini Ferguson, 
1969, homonym to Nemoriini Gumppenberg, 1887. Apetovia Krulikovsky, 
1918, syn. n. to Heliothea Boisduval, 1840. Aglossochloris Prout, 1912, down- 
graded from genus to subgenus (of Thetidia Boisduval, 1840). Antonechloris 


3 


Raineri, 1994, downgraded from genus to subgenus (of Thetidia Boisduval, 
1840). Hissarica Viidalepp, 1988, downgraded from genus to subgenus (of 
Xenochlorodes Warren, 1897). Phaiogramma Gumppenberg, 1887, stat. n., 
removed from synonymy of Chlorissa Stephens, 1831, and raised to generic 
rank. Pseudoterpna coronillaria cinerascens (Zeller, 1847), stat. n. raised from 
synonymy to subspecies (Italy). Hemistola siciliana Prout, 1935, stat. n. raised 
from subspecies (of Hemistola chrysoprasaria (Esper, 1795)) to species (Sicily). 
Euchloris dissimilis Warren & Rothschild, 1905, syn. n. to Acidaliastis micra 
Hampson, 1896. Three species are transferred from Chlorissa to Diplodesma 
Warren, 1896 : Diplodesma approximans (Warren, 1897), comb. n., Diplodesma 
subrufibasis (Prout, 1930), comb. n., Diplodesma eborilitoris (Fletcher, 1958), 
comb. n., nec (Prout, 1930: unavailable). Provisionally six species are in- 
cluded in Phaiogramma (transferred from Chlorissa) : Phaiogramma faustinata 
(Millière, 1868), comb. n. (type species), Phaiogramma stibolepida (Butler, 
1879), comb. n., Phaiogramma discessa (Walker, 1861), comb. n., Phaiogramma 
patialensis (Rose & Devinder, 1985), comb. n., Phaiogramma pulmentaria 
(Guenée, 1857), comb. n., and Phaiogramma polemia (Prout, 1920), comb. n.. 
Lectotypes are designated for Orthostixis cribraria amanensis Wehrli, 1932, 
and Holoterpna pruinosata (Staudinger, 1898). 


Zusammenfassung 


Gegenstand der vorliegenden Verôffentlichung sind die Geometriden (Ortho- 
stixinae und Geometrinae) der Levante einschließlich der umliegenden Lander 
(Naher Osten). Die Morphologie wird fiir 53 Taxa der Artgruppe (Arten und 
Unterarten) beschrieben. Für alle behandelten Genera wird die Flügeläderung, 
für alle behandelten Taxa sowohl der männliche als auch der weibliche Genital- 
apparat abgebildet. Desweiteren werden für Triben und Gattungen Differen- 
tialdiagnosen erstellt sowie môgliche Verwandtschaftsbeziehungen diskutiert. 
Die Systematik der behandelten Geometriden-Unterfamilien wird einer gene- 
rellen Uberpriifung unterzogen, mehr als 100 Gattungen werden den 19 er- 
wähnten Triben bzw. Unterfamilien der ,,Alten Welt” zugeordnet. Für Ortho- 
stixinae Meyrick, 1892 (Oenochrominae auct. nec Guenée, 1857) wird der 
Rang einer Unterfamilie bestatigt, in die Konzeption der Differentialmerkmale 
werden 7 Gattungen einbezogen. Eine neue Tribus wird aufgestellt : Micro- 
loxiini tribus n. mit der Typus-Gattung Microloxia Warren, 1893. Drei neue 
Gattungen und eine neue Untergattung werden beschrieben : Microbaena gen. 
n. mit der Typusart Phorodesma pulchra Staudinger, 1897 ; Proteuchloris gen. 
n. mit der Typusart [| Geometra (Phorodesma)| neriaria Herrich-Schäffer, 1852 ; 
Acidromodes gen. n. mit der Typusart Acidaliastis nilotica Wiltshire, 1985 ; 
Victorinella subgen. n. von Victoria Warren, 1897, mit der Typusart Victoria 
sematoperas Prout, 1916. Drei neue Arten und drei neue Unterarten werden 
beschrieben : Thetidia persica sp. n. aus dem Iran, Victoria wiltshirei sp. n. 
aus dem Sudan, Hemidromodes unicolorata sp. n. aus Kenia, Pseudoterpna 
coronillaria halperini ssp. n. aus Israel, Microbaena pulchra minor ssp. 0. 
aus dem Sudan, und Eucrostes indigenata lanjeronica ssp. n. aus Spanien. 


4 


Weitere taxonomische Änderungen : Archaeobalbini Viidalepp, 1981, Emen- 
dation von Archeobalbini. Terpnini Inoue, 1961, jiingeres Homonym von 
Terpnini Kuznetsov, 1904 (~synonym zu Geometrini Duponchel, [1845]), und 
„synonym” (aber unverfügbar) zu Pseudoterpnini Warren, 1893. Neohipparchini 
Inoue, 1961, syn. n. von Geometrini Guenée, 1844. Ochrognesiini Inoue, 1961, 
syn. n. von Nemoriini Gumppenberg, 1887. Nemoriini Ferguson, 1969, homo- 
nym zu Nemoriini Gumppenberg, 1887. Apetovia Krulikovsky, 1918, syn. n. 
von Heliothea Boisduval, 1840. Aglossochloris Prout, 1912, herabgestuft von 
Gattungs- zu Untergattungs-Rang (von Thetidia Boisduval, 1840). Antonech- 
loris Raineri, 1994, herabgestuft von Gattungs- zu Untergattungsrang (von 
Thetidia Boisduval, 1840). Hissarica Vüdalepp, 1988, herabgestuft von Gat- 
tungs- zu Untergattungs-Rang (von Xenochlorodes Warren, 1897). Phaio- 
gramma Gumppenberg, 1887, stat. n., aus der Synonymie von Chlorissa 
Stephens, 1831, in den Gattungsrang erhoben. Pseudoterpna coronillaria 
cinerascens (Zeller, 1847), stat. n., aus der Synonymie in den Rang einer Unter- 
art erhoben (Italien). Hemistola siciliana Prout, 1935, stat. n., vom Unterarts- 
Rang (von Hemistola chrysoprasaria (Esper, 1795)) in den Rang einer Art 
erhoben (Sizilien). Euchloris dissimilis Warren & Rothschild, 1905, syn. n. 
von Acidaliastis micra Hampson, 1896. Drei Arten (ehemals Gattung Chlorissa 
Stephens, 1831) werden mit neuen Gattungen kombiniert : Diplodesma ap- 
proximans (Warren, 1897), comb. n., Diplodesma subrufibasis (Prout, 1930), 
comb. n., Diplodesma eborilitoris (Fletcher, 1958), comb. n., nec Prout, 1930 
(nicht verfügbar). Nach dem derzeitigen Stand umfaßt die Gattung Phaio- 
gramma sechs Arten (übertragen aus Chlorissa) : Phaiogramma faustinata 
(Milliere, 1868), comb. n. (Typusart), Phaiogramma stibolepida (Butler, 1879), 
comb. n., Phaiogramma discessa (Walker, 1861), comb. n., Phaiogramma 
patialensis (Rose & Devinder, 1985), comb. n., Phaiogramma pulmentaria 
(Guenée, 1857), comb. n., und Phaiogramma polemia (Prout, 1920), comb. n.. 
Festlegung von Lectotypen erfolgte bei Orthostixis cribraria amanensis Wehrli, 
1932, und Holoterpna pruinosata (Staudinger, 1898). 


Introduction 


This is the first paper in the first of three series of publications by 
the author on the Geometridae of the Middle East. The three series 
are ( 


1) Morphology of the species and subspecies occurring in the Levantine 
basin and neighbouring countries, with figures of male and female 
genitalia of all taxa, of venation of all genera, description of external 
structure of the imago and discussion of relationships at subspecies, 
species, genus and tribe level. 


2) Systematic list of these species with some important synonyms, 
citation of original descriptions, locus typicus and a short survey of 
the geographical distribution within the study area. 


3) Faunistic data, phenology and ecology of the species occurring in 
the state of Israel (administration frontiers of 1990). 


Only the morphology of adult stages will be considered in this paper. 
The study area includes S. Turkey south of the summits of the Taurus 
Mountains between Göksu and the River Tigris, Cyprus, Syria, 
Lebanon, Irag, Kuwait, N. Saudi Arabia (north of a line from Medina 
to Kuwait), Israel, Jordan, Egypt (the reasons for adopting these 
boundaries are given in the second series of publications). 


In the sections on “differential features” only some of the more im- 
portant features are described. Characters mentioned in the description 
of tribes are not repeated where they are also valid for genus or species. 
In some cases the key differential diagnostic characters of tribes do 
not clearly reflect the phylogenetic relationships, which need further 
studies to be elucidated. Diagnoses are valid for the study area, but 
the author tried to check the validity for taxa from other regions all 
over the world (mainly “Old World” regions). Similarly Old World 
tribes not represented in the study area are taken into consideration, 
which should assist the construction of a better classification of the 
family Geometridae. 


The systematic arrangement of the subfamily Geometrinae at tribal 
level largely follows the results of the extensive studies published by 
Inoue (1961) and Viidalepp (1981). The author however tried to re- 
examine the whole arrangement of tribes and genera on the basis of 
the results of the present morphological studies. Type species of more 
than 100 genera of Orthostixinae and Geometrinae have been examined 
and these genera attributed to the various tribes. Many tropical genera 
cannot be included in the tribes mentioned here. They need further 
studies with the aim of establishing new taxa at family group (tribal 
level). 


The following structural details are given for all species and subspecies 
(SQ) : venation, frenulum, frons, vertex, tongue, palpus, antenna, hind- 
tibia/tarsus, genitalia (incl. sternite 8 and tergite 8). Additional features 
are included if they show diagnostic characters. The terminology of the 
genitalia largely follows that of Ferguson (1985), Pitkin (1993) and Scoble 
(1994). Terminology of venation according to Pitkin (1993 : Fig. 68). 


Terms and abbreviations used in the descriptions of venation and 


oh) ——— 


external structure : “separate” = separate origin from cell of two veins ; 


6 


“connate” = veins arising from cell (upper or lower angle) at the same 
point ; “palpus” = labial palpus. Measurements of antennal branches 
or cilia refer to the longest, and comparison is made with the thickness 
of the filament at the point where this branch or cilium arises (usually 
at 1/3 to 1/2 length of flagellum) — 


Further abbreviations 


ZSM Zoologische Staatssammlung München 
BMNH The Natural History Museum, London 
TAU Tel Aviv University collection 


NMW Naturhistorisches Museum, Wien 
ZFMK Zoologisches Forschungsinstitut und Museum Alexander 
Koenig, Bonn 


SYSTEMATIC PART 
Subfamilia ARCHIEARINAE Fletcher, 1953 


DIFFERENTIAL FEATURES: Venation: M2 of hindwing rudimentary. 
“Larva with 16 Legs” (Prout, 1915 : 1). Accessory tympanum reduced 
(Compare Cook & Scoble, 1992 : 229). 


REMARKS : Not represented in the study area. 
Subfamilia ORTHOSTIXINAE Meyrick, 1892 


DIFFERENTIAL FEATURES: Venation: Forewing with accessory cell 
(double in Myinodes and Eumegethes) ; R1 and R2 fused or free, both 
arising below cell apex ; R3-R5 and M1 widely separate in all examined 
genera; Sc + RI and Rs of hindwing approximate, but not touching 
(slightly touching in Myinodes and Eumegethes), in some Orthostixis 
spp. with cross vein (R1) near wing base ; M2 tubular in all wings. 
Frenulum stout and long in @, replaced by tuft of stiff hair-scales in 
female. Frons usually strongly convex. @ antenna filiform, ciliate (in 
the study area). Ansa of tympanal organ broad at base, tapering to 
the end in all species examined (compare Cook & Scoble 1992 : 227). 
& genitalia : Costa of valva strongly sclerotised. Surface of valva often 
wrinkled. ® genitalia: Surface of posterior part of corpus bursae 
strongly wrinkled, anterior part membranous and with smooth surface ; 
tergite 8 laterally tapering. 


Type species of the following additional genera have been examined : 
Gypsochroa Hübner, [1825], Derambila Walker, [1863], Naxa Walker, 
1856, Ozola Walker, 1861. 


REMARKS: Validity of subfamily rank for “Orthostixinae” was pos- 
tulated by Bleszinski (1960 : 22) and Kovacs (1987 : 190), but without 
presenting a clear concept. The seven genera mentioned here are very 
probably monophyletic and would perhaps be better placed between 
Larentiinae and Ennominae. The African and Indoaustralian genera 
Derambila and Ozola seem to link (in external appearance, venation 
and genitalia) the three groups Myinodes/ Eumegethes, Orthostixis/ 
Naxa and Gypsochroa. Male genitalia of Gypsochroa reveal relation- 
ships with Myinodes/ Eumegethes ; the venation of Gypsochroa is 
almost identical to that of Derambila. Nevertheless Gypsochroa has 
until recently been treated by many authors as a genus of Larentiinae 
near Chesias Treitschke, 1825 (e.g. Leraut, 1980: 143). The genitalia 
resemble those of the latter genus only superficially. Ansa of tympanum 
is as described above for Orthostixinae. Compare also the remarks 
to the genus Orthostixis. 


MYINODES Meyrick, 1892 


DIFFERENTIAL FEATURES: Venation see Fig. |: Forewing: RI and 
R2 arising separately from cell, forming with Sc two accessory cells ; 
crossvein Ri between cell and Sc weak. Hindwing : Rs + M1 distinctly 
stalked. @ genitalia: Uncus tapering, without basal lobes (“soci”) ; 
costa of valva strongly sclerotised ; ventral part of valva medially with 
stout inwardly directed projection ; surface of valva slightly wrinkled 
in the apical part. See also Hausmann (1994b). © genitalia : Apophyses 
anteriores more than 3/4 length of apophyses posteriores ! 


REMARKS: @ Genitalia in the group Myinodes/ Eumegethes are quite 
uniform, furthermore indicating relationships to the genera Ozola and 
Gypsochroa (see above). ® genitalia of both Myinodes/ Eumegethes 
have comparatively short ductus bursae, signum bursae lacking. 


Myinodes shohami Hausmann, 1994 


For data on the morphology and differential features with regard to 
the other species of the genus see Hausmann (1994b). Holotype re- 
examined. Venation : Fig. 1. 


GENITALIA 6 : see Fig. 30 (N. Israel). 
GENITALIA Q : see Fig. 95 (N. Israel). 


8 


EUMEGETHES Staudinger, 1898 


DIFFERENTIAL FEATURES : Venation see Fig. 2 : Forewing like that of 
Myinodes. Hindwing : Rs + MI not stalked. Cilia of antenna longer 
than in Myinodes. & genitalia : Quite similar to genitalia of the genus 
Myinodes. However the surface of the valva is strongly wrinkled as 
in the species of the tribes Calothysanini and Cyclophorini (Sterrhinae). 
@ genitalia: Uncus tapering, basal lobes absent. Juxta with two 
posterior processes. Aedeagus with stout single cornutus. © genitalia : 
Apophyses anteriores about half length of apophyses posteriores. 
Shows only minor differences from the genitalia of Myinodes, perhaps 
also related to Derambila. 


Eumegethes tenuis Staudinger, 1898 


EXTERNAL FEATURES: Venation see Fig. 2 and generic description. 
Tongue developed (about 2.5mm). Palpus with dark brown scales, 
comparatively long (about 0.8 mm = twice the diameter of the eye). 
Frons whitish, strongly convex, tapering to one central projection. 
Vertex whitish. Antenna of & ciliate with two rows of long cilia (longest 
2-2.5 times width of flagellum), of ® ciliate, cilia as long as width of 
flagellum. Hindtibia of both sexes with two pairs of long spurs. Relative 
length hindtibia/hindtarsus 3.0/2.2 mm. 


GENITALIA @ : see Fig. 31 (C. Algeria) and generic description. 
GENITALIA Q : see Fig. 96 (C. Algeria) and generic description. 


REMARKS : No Egyptian material available. Eumegethes picta (Turati, 
1934), described from E. Libya, provisionally has to be considered a 
form of E. tenuis (Rungs, 1981 : 223). More detailed studies are required 
to resolve this question. 


ORTHOSTIXIS Hübner, [1823] 


DIFFERENTIAL FEATURES : Venation see Fig. 3: Forewing : The first 
crossvein between Sc and cell must be identified as “R2” because Sc 
usually turns off to costa (sometimes distinctly) before joining this 
crossvein ; connection of Sc and this R2-crossvein must be interpreted 
as diverging RI from Sc; Rl-crossvein from cell to Sc (as present 
in Myinodes/ Eumegethes) lost. Hindwing : both Rs/MI and M3/ 
CuAl separate. Frons comparatively flat, only in the ventral part 
strongly convex. Hindtibia with two short spurs in both sexes. & 
genitalia : Uncus broad, not tapering towards end ; small basal lobes 


9 


beside uncus (“soci”) present; costa of valva strongly sclerotised, 
spinulose ; juxta with long posterior projection ; aedeagus without stout 
cornuti. Q genitalia : Apophyses anteriores | / 3-1/2 ener of apophyses 
posteriores ; ductus bursae fairly long. 


REMARKS: The relationship between the genera Orthostixis and Naxa 
is very close, as their & and © genitalia reveal (e.g. posterior projection 
of juxta, long ductus bursae, shape of signum etc.). In @ genitalia 
“soci” are present in both Ozola and Orthostixis (probably synapo- 
morphic character). Female genitalia are also somewhat similar. 


Orthostixis cribraria cribraria (Hübner, [1799]) 


EXTERNAL FEATURES : Venation (Fig. 3) characterised by a short cross- 
vein (RI) between Sc and Rs in the hindwing. Tongue developed and 
long (5 mm). Palpus somewhat weak, with outstanding scales, length 
in male 1.1-1.5 times diameter of eye ; female 1.5-1.7 times diameter 
of eye, tip pale brown. Frons and vertex whitish. Antenna of @ dentate, 
length of cilia 3/4 width of flagellum ; antenna of Q slightly dentate, 
length of cilia 1/3 width of flagellum ; in both sexes with light brown 
scales on the upper side. Hindtibia in both sexes with one pair of short 
spurs. There is little structural variation within the area of distribution ; 
specimens from Italy have somewhat smaller palpus. 


GENITALIA @ : see Fig. 32 (Hungary). No differences between specimens 
examined from Hungary, Italy, N. and C. Turkey. 


GENITALIA 9: see Fig. 97 (N. Turkey). Little difference between 
specimens examined from Sicily and N. Turkey: In Sicily the ductus 
bursae is broader near the corpus bursae and less sclerotised. Anthrum 
narrower. Signum bursae is more slender and more pointed in Turkish 
specimens. 


REMARKS: The distribution of the nominate subspecies probably does 
not reach the study area. Further studies are required with material 
from the “contact-zone” with the following taxon around the Taurus 
mountains. 


Orthostixis cribraria amanensis Wehrli, 1932 


EXTERNAL FEATURES : See characters of the nominate subspecies. Wings 
of both types of amanensis rather short. At the ZSM there are small 
specimens (infrasubspecific forms) from N. Greece and Bulgaria. Wehrli 
(1932 : 3) gives a more basal position of the postmedian line ; however 


10 


this character is variable in the populations of S. Turkey and Lebanon. 
The main differential features of the subspecies lie in the male genitalia. 


GENITALIA @ : see Fig. 33 (Paralectotype from the Amanus mountains). 
The spinulose field at the apex of the valva is more extended distally 
than its equivalent in the nominate subspecies. No difference observed 
between the specimens examined from the Taurus and the Amanus 
mountains. Lebanese specimens have a still more extended spinulose 
field. 


GENITALIA © : unknown. 


REMARKS: Lectotype (4, Syria, Taurus, Marasch, VII.1931, leg. 
Pfeiffer, coll. ZFMK) and Para-Lectotype (4, [Amanus], coll. ZFMK) 
designated and examined. 


Orthostixis cinerea Rebel, 1916 


EXTERNAL FEATURES: Venation as in O. cribraria (compare Fig. 3). 
Length of tongue 5.5 (Q) to 6.5 (4) mm! Length of palpus as in O. 
cribraria, scales not strongly adherent as in O. calcularia, tip (3/4 of 
last segment) dark brown. Antenna of male dentate, length of cilia 
3/4 width of flagellum ; antenna of female slightly dentate, length of 
cilia 1/2 width of flagellum, with dark brown scales on the upper side. 
Hindtibia in both sexes with one pair of short spurs. 


GENITALIA @ : see Fig. 35 (Cyprus). 


GENITALIA Q : see Fig. 99 (Cyprus). Ductus bursae long. Signum bursae 
triangular. 


REMARKS: Holotype (@, coll NHMW) examined. External and 
genitalic structure reveal a closer relationship to O. cribraria than to 
O. calcularia. | 


Orthostixis calcularia Lederer, 1853 


EXTERNAL FEATURES : Venation similar to that of O. cribraria (compare 
Fig. 3), but Sc + RI and Rs of hindwing not connected by crossvein 
RI at base. Length of tongue 1.0 (©) - 1.5 (4) mm only! Length of 
palpus as in O. cribraria, but stouter, scales adherent, tip dark brown. 
Frons and vertex white. Antenna of male dentate, cilia as long as width 
of flagellum ; antenna of female slightly dentate, length of cilia 1/3 
width of flagellum, with dark brown scales on the upperside in both 
sexes. Hindtibia (4, 9) with one pair of short spurs. No structural 
variation within the area of distribution. 


11 


GENITALIA @ : see Fig. 34 (C. Turkey). 


GENITALIA Q : see Fig. 98 (C. Turkey). Apophyses broad. Ductus bursae 
short. Signum bursae small. 


Subfamilia ALSOPHILINAE Herbulot, 1962 


DIFFERENTIAL FEATURES: Venation: Forewing: R3-RS distinctly 
stalked ; R3-R5 and MI typically, but not always, separate, M2 
developed, tubular. Hindwing: Sc + RI and Rs anastomosing for a 
long distance as in the Larentiinae (except some East Asiatic species) ; 
Rs and MI typically shortly stalked (with a very few exceptions). 
Female brachypterous. 


REMARKS: Not represented in the study area. The exceptions in 
venation render the validity of subfamily rank doubtful. A/sophila is 
placed in “Oenochrominae” in Inoue (1977). 


“Subfamilia HELIOTHEINAE” Exposito, 1979 


DIFFERENTIAL FEATURES (compare Exposito, 1979): Venation : M2 
tubular in all wings. Hindwing : Sc + RI and Rs not anastomosing ; 
M2 arising from cell very near MI. Ansa of tympanal organ broad 
at its base, tapering towards end, not dilated in the central part. 
Genitalia & : not unlike genitalia of many Geometrinae species : uncus 
tapered ; socius present in Heliothea, reduced in Petovia; shape of 
aedeagus rather similar to the typical shape in many Geometrinae 
species. 


TYPE SPECIES of the following genera examined : Heliothea Boisduval, 
1840 (West Mediterranean), Apetovia Krulikovsky, 1918 (Central 
Asiatic) and Petovia Walker, 1854 (Ethiopian). 


REMARKS: Not represented in the study area. Validity of subfamily 
rank very doubtful. The possibility of inclusion in the subfamily Geo- 
rnetrinae as a separate tribe (as done in Vives Moreno, 1994: 371) 
should be considered. With regard to the tympanal structure I hesitate 
to change the status formally ; the above described feature is common 
in “Oenochrominae” (s.l.) and Orthostixinae and quite unusual in Geo- 
metrinae (compare Cook & Scoble, 1992). The three above mentioned 
taxa of genus group apparently monophyletic. ¢ genitalia of Heliothea 
and Apetovia extremely similar to each other. Therefore the latter is 
downgraded to synonymy: Apetovia Krulikovsky, 1918, syn. n. to 
Heliothea Boisduval, 1840. 


LZ 


Subfamilia GEOMETRINAE Guenée in Duponchel, 1844 [1845] 


DIFFERENTIAL FEATURES : Venation : Forewing : Rl and R2-R5 typical- 
ly, but not always, arising separately from cell and R2-R5 stalked. 
Hindwing : Rs appressed or fused to Sc + RI for short distance only 
(exceptions e.g. many Microloxuni genera and Xenochlorodes) ; M2 
fully developed in both wings and arising from above middle of cell. 
Ansa of tympanal organ narrow at the base, widening in the middle 
and then tapering to the end in nearly all species occurring in the 
study area ; compare Cook & Scoble (1992 : 228). Antenna of male 
usually bipectinate (exceptions in the study area: Aplasta, Chlorissa 
and Neromia). Wing colour mainly green. 


Tribus PSEUDOTERPNINI Warren, 1893 
(Terpnini Inoue, 1961) 


DIFFERENTIAL FEATURES : Venation : Forewing: RI arising from cell 
below cell apex ; M3 and CuAl unstalked. Hindwing: Rs + MI 
typically connate or very shortly stalked (exception : Aplasta). M3 and 
CuAl unstalked, often separate. Frenulum of @ typically present 
(exception : Holoterpna and Aplasta). Hindtibia of both sexes typically 
with four spurs (exception : e.g. Holoterpna). Q antenna filiform, with 
very short cilia. Abdominal crests developed (exception : Holoterpna 
and Aplasta). Tympanum as described for subfamily. @ genitalia: 
“Uncus” deeply forked (main synapomorphic character); “socius” 
absent or rudimentary (e.g. Pseudoterpna or some Dindica species ; 
homology to the socius of other Geometrinae not confirmed) ; basal 
coremata of valva present; costa of valva at base widely extended 
into inwardly directed sclerites ; sternite 8 and tergite 8 simple. © 
genitalia : Apophyses anteriores comparatively short ; lamella anteva- 
ginalis usually band-shaped. 


TYPE species of the following additional genera examined : Dindica 
Moore, 1888 ; Metallolophia Warren, 1895 ; Pachyodes Guenée, 1857 ; 
Mimandria Warren, 1895 ; Agathia Guenée, 1857 ; Dooabia Warren, 
1894. 


REMARKS: The subdivision into the subtribes “Terpnina Inoue, 1961” 
and “Pseudoterpnina Herbulot, 1963” (Vudalepp, 1981 : 95) is excluded 
from consideration in this paper (Terpnina and Terpnini are invalid 
with regard to Kuznetsov’s (1904) Terpnidae (= synonym to Geo- 
metrini), Pseudoterpnina has to be associated with “Warren, 1893” 
according to article 36 ICZN ; compare remarks in the systematical 


13 


part). Some features of Holoterpna and Aplasta are anomalous within 
Pseudoterpnini ; the systematical position of Aplasta is fairly isolated. 
According to genitalic morphology Dooabia is near Agathia. Bifurcation 
of uncus also occurs in Mimandria (M. insularis Swinhoe, 1904, 
examined), although Janse (1935 : 260) states “simple uncus”. 


PINGASA Moore, [1887] 


DIFFERENTIAL FEATURES : Venation (compare Fig. 4) : Forewing: Sc 
and RI free, R2-R5 stalked, R2-R5 and MI connate. Hindwing : 
Sc + RI and Rs appressed for short distance only ; both Rs/MI and 
M3/CuAl usually unstalked, sometimes on very short common stalk ; 
A2 present, rudimentary. Frenulum in both male and female present. 
Third segment of palpus elongate, “naked” (Janse, 1935 : 255), 1.e. 
slender, smooth, with short adherent scales. 4 and © hindtibia typically 
with four spurs. Wing pattern quite characteristic and rather uniform. 
Genitalia & : dorsal part of anellus (“superjuxta”; compare Inoue, 1961 : 
89) strongly sclerotised. 


Pingasa lahayei multispurcata Prout, 1913 


EXTERNAL FEATURES : Venation see Fig. 4: In one male from Saudi 
Arabia however R2-5 and MI very shortly stalked. Length of tongue 
6-7 mm. Palpus (@) 1.5 times diameter of eye, last segment with very 
short adherent scales, length of this segment ca. 0.4 mm ; total length 
in the female 2.5 times diameter of eye, length of last segment ca. 
1.1 mm. Frons only slightly convex, black, lower 1/3 creamy white. 
Male antenna bipectinate, length of longest branches ranging from twice 
to 2.5 times width of flagellum at same point. Hindtibia (3) with two 
pairs of spurs of unequal length. Only minor structural variation 
between the specimens examined from S. Iran, Saudi Arabia and 
Nigeria. 

GENITALIA G : see Fig. 36 (Saudi Arabia). Only minor variation be- 


tween the specimens examined from S. Iran, Saudi Arabia, Morocco 
and Nigeria. 


GENITALIA Q : see Fig. 100 (N. Nigeria). 


REMARKS : According to Wiltshire (1990 : 107) Pingasa multispurcata 
Prout, 1913, is probably a synonym of P /ahayei (Oberthür, 1887). How- 
ever wing colour is somewhat more brownish in the Asiatic populations. 
Absence of records from Egypt and Sudan suggests disjunct distribution 
area. 


14 


PSEUDOTERPNA Hubner, [1823] 


DIFFERENTIAL FEATURES : Venation (compare Fig. 5) : In the forewing 
no accessory cell. R2-R5 and MI shortly stalked. In the hindwing 
Sc + RI and Rs appressed for about 1/3 length of cell; Rs and MI 
unstalked ; M3 and CuAl unstalked ; A2 present, rudimentary. Frenu- 
lum present in male, absent in female (only some long hairs present). 
Hindtibia (39) with two pairs of spurs of unequal length in all species. 
Tarsus somewhat shortened. Genitalia & : Socius rudimentary, adherent 
to base of uncus ; “superjuxta” (compare Pingasa) absent ; harpe and 
gnathos spined. 


Pseudoterpna coronillaria axillaria Guenée, 1857 


EXTERNAL FEATURES: Venation : see generic description. Forewing : 
R2-R5 and MI unstalked. Tongue developed, length ca. 5 mm. Length 
of palpus 1.5 times diameter of eye (upper side black) in both sexes. 
Frons slightly convex, black. Antenna of male bipectinate, longest 
branches three times width of flagellum ; antenna of female filiform, 
ciliate, longest cilia 1/3 width of flagellum. Width of male hindtibia 
2-3 times width of tarsus. Length of hindtarsus 3/4 length of hindtibia. 
Some structural variation within the area of distribution of the species : 
In West Mediterranean populations including North Africa (= P c. 
coronillaria and P. c. algirica Wehrli, 1930) width of male tibia about 
2.5-4 times width of tarsus, tongue somewhat shorter (4-5 mm). Slight 
differences also in ratio tarsus/tibia : 1/2-2/3 in European subspecies. 


GENITALIA @: see Fig. 37 (C. Lebanon). Gnathos without the 
prominent terminal spines present in the West Mediterranean popu- 
lations. Shape of harpe somewhat variable, short and triangular in 
the specimens examined from the locus typicus. Distal cornutus long 
and slender as in the nominate subspecies, central cornutus com- 
paratively slender, its teeth short. 


GENITALIA © : see Fig. 101 (C. Lebanon). Shape of corpus bursae 
variable, but always more elongate than in the other subspecies. Rather 
similar to P. rectistrigaria (Fig. 104 ; see below). Apophyses posteriores 
slightly shorter than in European subspecies. 


Remarks: P c. coronillaria (Hübner, [1817]) ; as illustrated on pl. 
93, figs. 479-482 in Hübner is bluish-grey coloured, postmedian line 
strongly marked (Spain, France, Corsica ; locus typicus one of these 
countries). North African populations (P. c. algirica Wehrli, 1930) are 
characterised by a brownish ground colour, dark marginal area and 


15 


forewing apex slightly pointed. The typical P c. axillaria is small, 
brightly coloured, postmedian line very delicate, antemedian line almost 
wanting, forewing apex rounded. Culot (1919 : 7) mentions P. c. axil- 
laria from “Syria” flying at the same locality together with specimens 
that resemble the nominate subspecies. To date the author has not 
been able to make similar observations. 


Pseudoterpna coronillaria halperini ssp. n. 


HoıoTyPpE: @, NE. Israel, Qibbuz Senir, 5 km S. Banyas, 300 m, 
14.-16.1V.1987, leg. G. Müller, coll. TAU. 


PARATYPES : 19, NE. Israel, Banyas, 380 m, 22.1V.1988, leg. G. Miiller, 
coll. ZSM ; 16, id., 12.1V.1988 ; 24 id., coll. TAU ; 28, NE. Israel, 
Shetula, 4 km NE Fassuta, 680 m, 10.1V.1987, leg. G. Müller, coll. 
ZSM; 18, NE. Israel, Qibbuz Senir, 5 km S. Banyas, 300 m, 
11.IV.1988, leg. G. Müller, coll. ZSM ; 19, id., 12.1V.1988. 


EXTERNAL FEATURES : Venation see Fig. 5 : Forewing : R2-R5 and M1 
in the typical populations near Mt. Hermon usually stalked, other 
Israeli and Jordanian populations slightly more variable in this 
character. Larger and with darker wing colour than in typical P. c. 
axillaria from Beirut. Postmedian and antemedian lines much more 
distinctly marked. Other external features as described for P c. axillaria 
from Lebanon. 


GENITALIA @ : see Fig. 38 (NE. Israel). Similar to that of P c. axillaria, 
but harpe longer ; valva longer and narrower. No structural difference 
in genitalia of specimens examined from N. Jordan, and also 14 from 
N. Lebanon (!). 


GENITALIA © : see Fig. 102 (NE. Israel). Corpus bursae broader and 
more rounded than in P c. axillaria, but at the anterior (oral) end 
more elongate than in the European subspecies. Apophyses posteriores 
shorter. 


REMARKS: Distribution areas of P c. axillaria and P. c. halperini 
probably somewhat isolated from each other by the Bekaa Plain. 
According to the data published by Wiltshire (1939 : 43) with strikingly 
different phenology : “June-December” for P. c. axillaria ; end of March 
to beginning of May in P c. axillaria from Israel. The populations 
from NW. Israel — although identical in their external appearance 
with typical P c. halperini — are excluded from the type series, since 
they are slightly different in the male genitalia. An illustration of the 
imago of P. c. halperini will be given in a subsequent paper. 


16 


Both subspecies are probably well isolated from the nominate subspecies. 
Although P. coronillaria is said to occur in the Balkans, W. Turkey 
and Taurus, the author has never seen material from there (perhaps 
it only occurs as very local relict populations). The species is also absent 
from N. Italy (except Liguria). 


C. Italian populations characterised by short, broad and dentate 
cornuti, short harpe with long spines, gnathos without the long spines 
present in the specimens from Spain and N. Africa (see Figs. 39, 103), 
and the bright grey wing colour. External appearance of the populations 
from Sicily and S. Italy is transitional to the North African populations, 
genitalically however they correspond to the specimens from C. Italy. 
Provisionally I refer these populations of C. Italy, S. Italy and Sicily 
to Pseudoterpna coronillaria cinerascens (Zeller, 1847), stat. n., comb. 
n. (perhaps beginning speciation). The venation of P c. cinerascens 
is characterised by R2-R5 and MI of forewing often being shortly 
stalked, but they are never stalked in the nominate subspecies (Spain, 
France, Corsica). 


Pseudoterpna rectistrigaria Wiltshire, 1948 


EXTERNAL FEATURES : Venation : Hindwing: Both Rs/MI and M3/ 
CuAl separate ; Rs and CuAl arising subapically below angle of cell. 
Tongue developed, length about 4 mm. Length of palpus (Q) about 
1.5 times diameter of eye, only the upper side black. Frons convex, 
black. Antenna (©) filiform, with very short cilia (1/4 width of flagel- 
lum). Relative length hindtibia/ hindtarsus 3.7/3.0 mm (©). 


GENITALIA @: see Wiltshire (1948 : 80, Fig. B). Harpe very short, 
triangular. 


GENITALIA 9 : see Fig. 104 (Cyprus). Quite similar to that of P c. 
axillaria. Corpus bursae more slender. Apophyses posteriores slightly 
shorter than in European subspecies. 


REMARKS : Compare Hausmann (1994a). 
Holoterpna Püngeler, 1900 


DIFFERENTIAL FEATURES : Venation see Fig. 6: Forewing: R2-R5 
distinctly stalked, R2-R5 and MI connate (exceptionally very shortly 
stalked). Hindwing : Sc + RI and Rs appressed for a short distance 
only; Rs and MI (as in Aplasta) shortly stalked, somewhat variable ; 
M3 and CuAl not stalked, connate (or exceptionally separated by a 
short distance) ; A2 present, rudimentary. Frenulum in both & and 


17 


Q absent (as in Aplasta). Tongue lacking. & antenna shortly bipectinate, 
@ antenna filiform. & and © hindtibia with only one pair of terminal 
spurs. Hindtarsus not shortened. 5 genitalia: Uncus forked to 1/3- 
1/2 of its length only ; socius absent ; valva without spinulose harpe ; 
basal coremata of valva small. © genitalia: Lamella antevaginalis in 
female genitalia only slightly sclerotised. 


REMARKS: Type species of the genus is Holoterpna diagrapharia 
Piingeler, 1900. The synonymisation of Dyschloropsis Warren, 1895, 
with Holoterpna in Vojnits (1976 : 170) is erroneous, as the genitalia 
of both type species evidently show! Dyschloropsis with the type species 
Jodis impararia Guenée, 1857, is a valid genus, which has to be placed 
in the Thalerini. The African species “Holoterpna errata Prout, 1922” 
must be transferred to another genus. Some features of Holoterpna 
are reminiscent of Hemistolini (frenulum, venation, papillae anales in 
© genitalia). 


Holoterpna pruinosata (Staudinger, 1898) 


EXTERNAL FEATURES : See generic description. Palpus white, small, in 
both sexes ca. 1.3 times diameter of eye ; last segment very slender, 
with short adherent scales, somewhat resembling that of Pingasa, but 
not elongate. Frons rather flat, brown. Vertex white. Antenna of @ 
shortly bipectinate, length of longest branches 1.5 times width of 
flagellum ; antenna of female filiform, almost without cilia. Hindtibia 
of both sexes with two terminal spurs of nearly equal length. Length 
of tarsus somewhat exceeding length of tibia (distinguishing character 
from Pseudoterpna). 


GENITALIA @: see Figs. 40, 41 (N. and C. Israel). Specimens from 
C. Israel with valva slightly more tapered and narrower than in the 
populations from N. Israel. 


GENITALIA Q : see Fig. 105 (N. Israel). Apophyses posteriores 3-4 times 
length of apophyses anteriores. Surface between papillae anales cha- 
racteristically folded. 


REMARKS: Lectotype designated (4, Israel, Jerusalem, 1895, leg. 
Paulus, coll. ZMB). Two specimens examined from Trieste (N. Italy) 
structurally without differences from the populations from N. Israel. 


APLASTA Hubner, [1823] 


DIFFERENTIAL FEATURES: Venation see Fig. 7: Forewing venation 
rather variable: Sc always fused with RI, length of fusion very 


18 


variable ; Rl exceptionally re-anastomosing with R2-R4 ; R3 excep- 
tionally present ; R2-R5 and MI connate or arising separately from 
cell. Hindwing : Sc + RI and Rs appressed for about half length of 
cell, sometimes even more ; Rs and MI always distinctly stalked ; M3 
and CuAl separate. Frenulum in both male and female absent. Tongue 
lacking, or sometimes rudimentary. Male antenna slightly dentate, 
ciliate (as e.g. in Dooabia), cilia only 1/5 width of flagellum. Antenna of 
© almost identical to that of male, somewhat less dentate. 3 genitalia : 
valvae asymmetric : spinulose harpe present on the right valva only ; 
basal coremata of valva small ; posterior edge of sternite 8 medially 
slightly notched ; tergite 8 simple. ® genitalia: lamella antevaginalis 
in female genitalia reduced. 


Aplasta ononaria (Fuessly, 1783) 


EXTERNAL FEATURES: See Fig. 7 and generic description. Palpus 
brown, bushy-scaled ; length of palpus in @ 0.9-1.2 mm (1.6-1.8 times 
diameter of eye), in female 1.1-1.3 mm (2.0-2.2 times diameter of eye) ; 
last segment as described for Holoterpna. Frons rather flat, dark brown. 
Vertex with greyish scales. Hindtibia (¢Q) with two pairs of spurs of 
nearly equal length. Relative length hindtibia/hindtarsus 1 : 1 (1.7- 
2.5mm). No constant differences in structure between the various 
populations within the area of distribution. 


GENITALIA @: see Figs. 42-44 (N. Israel). Considerable infrapopula- 
tional variability in shape of valvae. No constant differences between 
specimens examined from Germany, France, Greece, Turkey, Cyprus 
and Israel. 


GENITALIA © : see Fig. 106 (Israel). No significant and constant dif- 
ferences between specimens examined from Greece, Turkey and Israel. 


REMARKS : On the basis of more than 20 @ genitalia slides no constant 
correlation has been observed between structural details and flight 
season or habitus respectively (“f. berytaria Staudinger, 1901”; “f. fae- 
cataria Hübner, [1823]”). In Israel the smaller yellowish specimens tend 
to have narrower valva and smaller spines on the gnathos, but this 
is not a constant feature of all specimens examined. From some 
localities in Israel only dark coloured specimens are known, from other 
places only yellow ones (mainly surroundings of Mt. Hermon) ; how- 
ever sympatric occurrence is often found. The above mentioned “habi- 
tus forms” (4) are perhaps correlated with particular flight seasons 
(darker specimens emerging somewhat earlier ; compare Ellison & Wilt- 
shire, 1939 : 43). This observation has to be verified with more ex- 


19 


tensive material and by rearing the various “entities” (see also Haus- 
mann, 1994a). 


Tribus ARACIMINI Inoue, 1961 


DIFFERENTIAL FEATURES : Venation similar to that of Pseudoterpnini : 
Rs and MI of hindwing unstalked, M3 and CuAl unstalked, often 
separate. Frenulum present in male. Palpus comparatively short in both 
sexes. Hindtibia with four spurs in both sexes. Female antenna filiform. 
Ansa of tympanal organ rather broad at its base, only slightly dilated 
in the central part, tapering to the end. Male genitalia characteristic 
with strongly sclerotised socius bearing large spinulose lobi at base. 
The simple uncus very short, almost wanting. Gnathos broken. Perhaps 
related to Geometrini. 


REMARKS : Type species of type genus (Aracima muscosa Butler, 1878) 
examined. Not represented in the study area. 


Tribus ARCHAEOBALBINI Viidalepp, 1981 


DIFFERENTIAL FEATURES : See Viidalepp (1981). 


REMARKS : “Archaeobalbini” is an emended spelling of “Archeobalbini 
Viidalepp, 1981”. Not represented in the study area. Some features, 
e.g. strongly sclerotised and elongate socius, indicate close relationships 
to Geometrini. Perhaps also related to Pseudoterpnini. 


Tribus GEOMETRINI Guenée in Duponchel, 1844 [1845] 


DIFFERENTIAL FEATURES : Venation similar to that of Pseudoterpnini : 
Rs and MI of hindwing separate (in Paramaxates on very short 
common stalk) ; M3 and CuAl separate in both wings (exceptionally 
connate). Frenulum present in male, absent in female, (male frenu- 
lum absent in Paramaxates). As in most species of Pseudoterpnini 
last segment of palpus slender (“naked”) and long, especially in the 
female. Q antenna filiform, with very short cilia. Hindtibia of both 
sexes with 4 spurs. Abdominal crests absent. Ansa of tympanal organ 
usually widened into hammer-headed plate at apex ; sometimes with 
big globular lacinia (e.g. Chlororithra, Neohipparchus, Tanaorhinus). 
Genitalia & : Uncus more or less reduced (present e.g. in Jotaphora, 
Ornithospila, Sphagnodela and Neohipparchus) ; gnathos tapering, 
strongly sclerotised ; socius greatly elongate, slender, strongly sclerotised 
and widely spread (in Ornithospila membranous and somewhat shorter). 


20 


TYPE SPECIES of the following genera examined : Geometra Linnaeus, 
1758, lotaphora Warren, 1894, Tanaorhinus Butler, 1879, Ornithospila 
Warren, 1844 ; Neohipparchus Inoue, 1944, Paramaxates Warren, 1894 ; 
Chlororithra Butler, 1889 ; Sphagnodela Warren, 1893 ; in addition 
Mixochlora Warren, 1897, according to Inoue (1961 : 38). 


REMARKS : Not represented in the study area. The family group name 
“Geometridi” originally attributed to “Guenée in hitt.”. In Synchlorini 
Ferguson, 1969, the uncus is lacking and soci widely diverging as in 
the Geometrini, but Rs and MI of hindwing distinctly stalked. Male 
hindtibia (Ferguson, 1985 : 81) with 4 spurs (exception : some Che- 
teoscelis spp.). According to Pitkin (1993 : 45) Synchlorini is perhaps 
synonymous with Nemoriini “Ferguson, 1969”. Presence or absence 
of uncus is an unstable character in Geometrini (as defined here) ; 
therefore “Neohipparchini Inoue, 1961” should be better included in 
Geometrini (syn. n.). Both Neohipparchus and lotaphora have a stout 
external spine on the aedeagus, which is probably homologous. 


Tribus COMIBAENINI Inoue, 1961 


DIFFERENTIAL FEATURES : Venation very variable, in many cases even 
within populations of the same species. Hindwing: M3 and CuAl 
typically unstalked (with a very few exceptions). @ antenna bipectinate 
with very long branches. Ansa of tympanal organ tapering to the free 
end. & genitalia: Saccus forked, valva inserted in the central part of 
clasper ; uncus forked (exception : Microbaena, Uliocnemis) ; socius 
present ; gnathos weak or absent ; aedeagus long and slender (exception : 
Microbaena, Uliocnemis). ® genitalia: Apophyses anteriores rather 
long ; ostium bursae closely connected with surface of sternite 8 ; corpus 
bursae weakly sclerotised and small. 


TYPE SPECIES of additional genus examined : Uliocnemis Warren, 1893. 


REMARKS: Unstalked M3 and CuAl of hindwing is reminiscent of 
Pseudoterpnini and Geometrini and may indicate relationships. Ge- 
nitalia characterise Comibaenini as a very homogenous and apparently 
monophyletic group. The uncus region has a similar aspect in the genera 
Dichordophora Prout, 1913 (“tribe Dichordophorini’; Nearctic) and 
Oospila Warren, 1897 (Neotropical), which however — according to 
Ferguson (1985 : 5) — are “presumably unrelated” genera (convergence). 


MICROBAENA gen. n. 


Type species: Phorodesma pulchra Staudinger, 1897, by original 
(= present) designation. 


21 


DIFFERENTIAL FEATURES : Venation see Fig. 8 : rather similar to Comi- 
baena. Forewing : R2-R5 and MI stalked. Frenulum present in male 
(plesiomorphy). Tongue absent. Hindtibia (4) not dilated, with two 
terminal spurs of unequal length, without terminal projection (main 
distinguishing character separating Microbaena from all other Comi- 
baena species). 6 Genitalia : Uncus not forked (secondary apomorphic 
fusion 4) ; gnathos weak, laterally strongly sclerotised ; costa of valva 
basally with long, inwardly directed projection, medially with two 
prominent and pointed spine-like processi; aedeagus short, thick ; 
sternite 8 anteriorly and posteriorly concave, the posterior edge with 
two lateral incurved chitinous lobes ; tergite 8 with bilobed posterior 
border. 


REMARKS: The necessity of establishing a new genus was mentioned 
by Prout (1935: 12). There are close relationships with Uliocnemis, 
which externally corresponds well to Microbaena (e.g. male hindtibia 
with 2 spurs, without terminal projection). The main differential feature 
lies in the male genitalia: Uliocnemis with both processi of uncus 
absent ; shape of valva simple as in Thetidia ; aedeagus similar to that 
of Microbaena. In the @ genitalia of the Indoaustralian species Comi- 
baena inductaria (Guenée, 1857) and the African Comibaena unduli- 
linea (Warren, 1905) the two projections of the uncus are closely ap- 
proximate, fused at their base. Valva and aedeagus are quite similar 
in shape to those of M. pulchra. However the tongue is developed 
and the hindtibia as described for Comibaena. 


Microbaena pulchra (Staudinger, 1897), comb. n. 


EXTERNAL FEATURES : See generic description. Length of palpus (@) 
1.3-1.5 mm (1.5-1.7 times diameter of eye), second segment bushy- 
scaled, third segment “naked” (1.e. with short adherent scales), short : 
in Holotype 0.35 mm. Frons in Holotype from Israel distinctly convex 
in the ventral half, with many dark brown scales. Vertex white. Antenna 
of male bipectinate, with longest branches ca. 0.9-1.0 mm (about 8 times 
width of flagellum), tapering, last quarter simple and ciliate. Relative 
length hindtibia/ tarsus 2.2/1.8 mm in Holotype. 


GENITALIA 6 : see Fig. 45 (C. Israel ; Holotype). Compare generic 
description. Little difference between specimens examined from Israel 
(M. p. pulchra), Sudan and Zaire (M. p. minor ; see below and Fig. 46) : 
The valva in the nominate subspecies is somewhat broader. More 
extensive material is needed to verify this observation. 


GENITALIA © : unknown. 


2», 


REMARKS: Holotype (coll. ZMB) examined. Three NE. Sudanese 
males and one from Zaire are somewhat different : 


Microbaena pulchra minor ssp. n. 


HoLoTYPE : 4, Sudan sept. or., Kassala Prov., Erkowit, 1000-1300 m, 
25.V1.1962, leg. Remane, coll. ZSM. 


PARATYPES : 26, id. ; 14, Belgisch Kongo [Zaire], Aequator Provinz, 
Liboko, V.1945, coll. ZSM. 


DESCRIPTION (Fig. 146): Colour and pattern of all wings as in the 
nominate subspecies. Forewing length 6.8-9.1 mm only (in Holotype 
Of M. p. pulchra from Israel 10.6 mm). Third segment of palpus shorter 
(0.20 mm). Frons flat, very slightly convex in the ventral half, whitish 
with ochreous scales. Hindtarsus not shortened, length of both hindtibia 
and tarsus 1.5-2.0 mm. 


REMARKS: The populations from SW. Saudi Arabia and S. Oman 
may belong to the nominate subspecies (compare Wiltshire, 1985 : 10). 


COMIBAENA Hiibner, [1823] 


DIFFERENTIAL FEATURES: Venation see Fig. 9: Forewing: R2-R5 
and MI stalked. In some African species and the Indoaustralian 
C. inductaria M3 and CuAl of hindwing distinctly stalked. Frenulum 
present in male (plesiomorphy ; exception: C. undulilinea), absent in 
female. Hindtibia with two proximal and two terminal spurs (plesio- 
morphy), pencil and one terminal projection (main differential feature 
shared by Comibaena and the following genus). 4 Genitalia: Uncus 
forked ; aedeagus long, slender, straight ; sternite 8 and often tergite 
8 with specific sclerotisations ; typical group with posterior edge of 
sternite 8 strongly sclerotised, crown-shaped. 


REMARKS: This is a rather heterogeneous group with many species ; 
perhaps some distinct subgenera should be separated from typical 
Comibaena : e.g. the African C. leucospilata Warren group and the 
C. inductaria/undulilinea group. The loss of the frenulum in C. unduli- 
linea could have taken place analogously (convergence) to the phylo- 
genetic line of Proteuchloris/ Thetidia, as the very different genitalia 
imply. The western Palaearctic species C. pseudoneriaria Wehrli, 1926, 
and C. serrulata Fletcher, 1963, are closely related to C. bajularia 
(Comibaena s.str.). 


23 


Comibaena bajularia (|Denis & Schiffermüller], 1775) 


EXTERNAL FEATURES : Venation see Fig. 9. Tongue developed, length 
about 2.5 mm. Palpus 1.8-2.0 times diameter of eye, second segment 
bushy-scaled, third segment “naked”. Frons flat. Male antenna long 
bipectinate to 4/5 of length of flagellum, longest branches 1.3-1.5 mm 
(about 10 times width of flagellum); female antenna filiform with 
short cilia, their length 1/3 width of flagellum. Terminal projection 
of hindtibia (4) covering 1/3 to 1/2 of tarsus. Little structural variation 
within the area of distribution: Longest branches of male antenna in 
the Anatolian populations up to 1.7 mm. 


GENITALIA @ : see Figs. 47-48 (C. Turkey and N. Italy). No significant 
difference between specimens examined from S. Germany, N. Italy, 
N. and C. Turkey except slight variation in sternite 8, which in C. 
Turkish populations is more broadly sclerotised than in European and 
N. Turkish populations ; projections of uncus basally fused in N. 
Turkey. Genitalia very similar to those of C. serrulata, but uncus more 
deeply forked, costa of valva with one prominent basal spine. Sternite 8 
in C. serrulata as figured for the C. Turkish populations. 


GENITALIA 9: see Fig. 107 (C. Turkey). No significant difference 
observed between specimens examined from S. Germany and C. Turkey. 
Without significant difference from C. serrulata and C. pseudoneriaria. 


Proteuchloris gen. n. 


TYPE SPECIES : [Geometra (Phorodesma)| neriaria (Herrich-Schäffer, 
1852), by original (= present) designation. 


DIFFERENTIAL FEATURES: Venation see Fig. 10: Forewing: R2-R5 
and M1 unstalked. Frenulum absent in male and female (synapomorphy 
of this and the following genus). Hindtibia with two proximal and 
two terminal spurs (plesiomorphy), furthermore with one terminal 
projection as in Comibaena. & Genitalia : Uncus forked ; valva simple, 
very similar to that of Thetidia; aedeagus long, slender, straight ; 
posterior and anterior edge of sternite 8 strongly sclerotised, notched 
medially on posterior edge ; tergite 8 rounded posteriorly. 


Remarks: Loss of the frenulum in the Comibaena inductaria/ 
undulilinea group is presumably analogous (see above). Male genitalia, 
sternite 8 and tergite 8 of Proteuchloris exactly as in Thetidia, but 
the structure of the hindtibia does not allow inclusion in that genus. 
The Asiatic species Thetidia albicostaria (Bremer, 1864) is somewhat 
similar in habitus, but structurally better placed in Thetidia. 


24 


Proteuchloris neriaria (Herrich-Schäffer, 1852), comb. n. 


EXTERNAL FEATURES : Venation see Fig. 10. Tongue developed, length 
2.0-2.3 mm. Palpus long, 2.2-2.8 times diameter of eye, second segment 
bushy-scaled, third segment “naked” somewhat !onger (0.5-0.7mm) than 
in C. bajularia. Frons flat. Male antenna long bipectinate to 4/5 of 
length, longest branches about 1.5 mm (about 10 times width of flagel- 
lum) ; female antenna dentate with very short forked branches (length 
of cilia 1/4 width of flagellum). Projection of male hindtibia covering 
1/3 to 1/2 of tarsus. There is no structural variation within the area 
of distribution. 


GENITALIA @ : see Fig. 49 (N. Israel). See generic description. There is no 
significant difference between the specimens examined from N. Greece, 
Turkey and N. Israel. 


GENITALIA © : see Figs. 108-109 (N. Israel and Macedonia). There are 
slight differences between the specimens examined from N. Greece, 
Turkey and N. Israel in the shape of the posterior opening of the ostium 
bursae, which is wider in the populations from Israel. 


THETIDIA Boisduval, 1840 
subgenus ANTONECHLORIS Raineri, 1994, stat. n. 


DIFFERENTIAL FEATURES : Venation see Figs. 11-12 ; unusually variable 
in all species of the genus. Forewing : R2-R5 and M1 usually unstalked. 
Frenulum absent in both sexes (often present, but weak in & of 7: 
anomica (Prout, 1935)). Tongue developed, but short and with tendency 
to reduction. Hindtibia of both & and © with four spurs ; proximal 
spurs of female with tendency to reduction in some species ; tibia 
without terminal projection (distinguishing generic character of Thetidia 
from Comibaena and Proteuchloris). @ Genitalia: Valva without 
processi or spines ; aedeagus curved and slender ; sternite 8 and tergite 8 
as described for Proteuchloris, median excavations on sternite shallow 
or lacking. 


In the type species 7. plusiaria Boisduval, 1840, Sc and RI are usually 
fused, but not always ; R2-R5 and MI of forewing are distinctly stalked, 
whereas in the subgenera Antonechloris and Aglossochloris they are 
usually unstalked ; the stalk of R2-RS and MI is rather stout, perhaps 
indicating a secondary fusion. In some species of Aglossochloris R2-R5 
and MI are shortly stalked. Exceptionally these veins are also stalked 


25 


in T. smaragdaria ; therefore full generic separation of the “green 
Thetidia” (Antonechloris) species from T. plusiaria (corresponding to 
the significant difference in wing colour) is not adopted here. Anto- 
nechloris Raineri, 1994, is downgraded from genus to subgenus rank. 


Thetidia (Antonechloris) smaragdaria (Fabricius, 1787) 


EXTERNAL FEATURES: Venation see Fig. 11 (Italy). Forewing: RI 
arising quite separately from cell angle; R2-R5 and MI sometimes 
shortly stalked. Hindwing : Rs and M1 usually unstalked with tendency 
to be stalked, especially in the Spanish populations (where sometimes 
distinctly stalked) ; M3 and CuAl usually connate, but quite variable. 
Length of tongue 1.2-1.6 mm. Length of & palpus 1.5 times diameter 
of eye, in 9 twice diameter of eye; second segment of palpus long, 
bushy-scaled, third segment “naked” (smooth with adherent scales). 
Frons flat, green, lower part white. Vertex green. Antenna of male 
bipectinate to 3/4 of length of flagellum only ; longest branches 0.8- 
0.9 mm (about 8 times width of flagellum) ; antenna of female filiform 
or (exceptionally) slightly bipectinate with extremely short branches 
in basal half of flagellum (somewhat varying even within populations) ; 
length of cilia 1/3 width of flagellum. Hindtibia of both sexes with 
four spurs. 


GENITALIA @ : see Fig. 50 (C. Italy). Projections of uncus long, some- 
times even longer than socius. Valva usually somewhat more slender 
than in 7! persica. Posterior edge of sternite 8 slightly tapering. Only 
small differences observed between the specimens examined from 
C. Italy, Sicily, Bulgaria, N. Greece, and NW. Caucasus. 


GENITALIA © : see Figs. 110-111 (C. Italy, NW. Caucasus). Vaginal plate 
posteriorly rounded. Chitinisation of ostium bursae narrow. Specimens 
examined from C. Italy, N. Greece and NW. Caucasus. 


REMARKS: Not represented in the study area (S. Turkish populations 
see T. persica), but discussed here with regard to possible confusion 
with the following and the preceding species. Main differential features 
between T. smaragdaria, T: volgaria and T. persica: Pectination of 
female antenna, stalk of Rs and M1 in hindwing, length of uncus pro- 
jections in @ genitalia, shape of vaginal plate in Q genitalia. According 
to Vudalepp (1976 : 845) the two following taxa and 7: anomica should 
be considered conspecific with T° smaragdaria. Since there are significant 
and constant differences not only in habitus, but also in structure, they 
are better considered as species unless interfertility 1s proved. 


26 


Thetidia (Antonechloris) volgaria (Guenée, 1857) 


EXTERNAL FEATURES: As described for T. smaragdaria with the fol- 
lowing differences : Venation of forewing: RI often arising very near 
cell apex. Hindwing : Rs and MI shortly stalked or sometimes connate ; 
M3 and CuAl separate. In the male antenna length of longest branches 
0.6-0.8 mm (5-7 times width of flagellum); female antenna slightly 
bipectinate to 1/2 length of flagellum with very short branches. 


GENITALIA & : see Fig. 51 (S. Russia). Length of projections of uncus 
variable, usually nearly as long as socius. Valva shorter than in T. sma- 
ragdaria, terminally broader. 


GENITALIA 9 : see Figs. 112-113 (S. Russia, Armenia). Anterior edge 
of tergite 8 with two tapered projections. Vaginal plate posteriorly, 
anteriorly and laterally tapering. Chitinisation of ostium with deep 
caudal excavation. 


REMARKS : Not represented in the study area (S. Turkish populations 
see T. persica), but discussed here with regard to the possible confusion 
with the following and the preceding species. Main differential features 
from T. smaragdaria are wing colour (mainly underside), pectination of 
antenna, shape of vaginal plate in © genitalia. Concerning the possible 
occurrence in Armenia see remarks to T. persica. The data of T. volgaria 
genitalia (“G 4762, N. Iran”), figured in Hausmann, 1991 (Fig. 9) are 
erroneously attributed ; the figure shows a female from S. Ural (Russia), 
prep!)G' 1523. 


Thetidia (Antonechloris) persica sp. n. 


HoLorTyPE : 9, N. Iran, Elburs mts. c.s., Tacht 1 Suleiman, Sardab 
Tal (Vandarbad), 2500-2700 m, 14.-18.V11.1937, leg. Pfeiffer & Forster, 
coll. ZSM. 


PARATYPES : 44, N. Iran, Elburs mts. c.s., Tacht 1 Suleiman, Sardab 
Tal (Vandarbad), 2500-2700 m, 14.-18.V11.1937, leg. Pfeiffer & Forster, 
coll. ZSM ; 24, N. Iran, Elburs mts. c.s., Tacht 1 Suleiman, Vandarban- 
Tal., 1900-2200 m, 1.-3. VII.1936, leg. Pfeiffer, coll. ZFMK ; 14, N. Iran, 
Elburs mts. c., Ort Demavend, Tar Tal, 2200-2500 m, 13.-17.V11.1936, 
leg. Pfeiffer, coll. ZFMK. 


EXTERNAL FEATURES (Fig. 147): Venation (Fig. 12): Forewing: Ri 
arising quite separately from cell angle; R2-R5 and MI connate. 
Hindwing : Rs and MI always stalked, common stalk sometimes very 
long ; in Anatolian populations usually on short common stalk ; M3 


2a 


and CuAl arising separately. Frenulum absent in both sexes. Tongue 
somewhat reduced, length 0.5-1.0 mm (often invisible between the 
palpi). Palpus longer than in the preceding species: & about twice 
diameter of eye, © 2.3-2.5 times diameter of eye. Frons and vertex as 
described for T. smaragdaria. Antenna of male bipectinate up to tip (!), 
longest branches about 0.80 mm (= about 7 times width of flagellum) ; 
antenna of female bipectinate to 2/3 or 3/4 length of flagellum, 
branches narrow, longest branches 0.3 mm (twice width of fiagellum). 
Hindtibia of both sexes with four spurs. Structural features (also 
venation !) are constant for all specimens examined from N. Iran and 
E. Turkey (Hakkari, Van), with the exception of the somewhat shorter 
branches of antenna (0.65) and the short palpus (1.7 times diameter 
of eye) in the single male from Van. In Anatolian populations length 
of branches of female antenna usually about 0.10-0.15 mm (= width 
of flagellum). One male examined from S. Turkey (Zeitun) is structurally 
as described for 7! persica, but forewing veins R2-5 and MI slightly 
separate, palpus shorter (1.4 mm = 1.9 times diameter of eye), length 
of tongue about | mm, longest branches of antenna 0.75 mm (7 times 
width of flagellum). 


Larger than T° volgaria, length of forewing 16.7-18.3 mm in the typical 
population from Elburz Mountains. Wing colour as in 7: smaragdaria. 
Both antemedian and postmedian lines more strongly marked than 
in 7? smaragdaria, but not as strongly as in 7: volgaria. Postmedian 
line of forewing rather straight, slightly curved on the underside. Post- 
median line of hindwing more prominent on the underside towards 
the margin than in T. smaragdaria. 


GENITALIA & : see Figs. 52-53 (N. Iran, S. Turkey). Length of pro- 
jections of uncus about half length of socius. Valva broad. Posterior 
edge of sternite 8 broad. Specimens examined from N. Iran, E., C. 
and S. Turkey, Armenia. 


GENITALIA Q : see Figs. 114-116 (N. Iran, C. and E. Turkey). Pro- 
jections on anterior edge of tergite 8 usually rounded. Vaginal plate 
posteriorly rounded, laterally tapering. Chitinisation of ostium bursae 
very broad, narrower in C. Anatolian populations. Apophyses anteriores 
rather short in one female from C. Turkey (Fig. 114). Specimens 
examined from N. Iran, E. and C. Turkey, Armenia. 


REMARKS: Two males and two females from Armenia have been 
examined : one female with antenna typical of 7! persica, also with 
genitalia similar to those of 7! persica. The other one with very short 
antennal branches as in 7: volgaria/smaragdaria and genitalia (Fig. 113) 


28 


corresponding with those of T: volgaria. From this it can be concluded 
that two species occur sympatrically in Armenia. In habitus the pre- 
sumed “7: volgaria” is more similar to T° smaragdaria. Rs and MI of 
hindwing in the Armenian pair of “7! persica” are only shortly stalked 
(4) or unstalked (Q). More material and experimental research is 
required to establish whether hybridisation occurs in the contact zones 
of the various taxa. Anatolian populations are included in T! persica 
provisionally for reasons of genitalic similarity. The differences from 
the other above mentioned taxa lie in the 9 antenna (see above), wing 
shape and colour (often deep green), pointed forewing apex, smaller 
than typical 7: persica. These populations perhaps constitute a distinct 
(sub)species ; more material and experimental research are required. 


Thetidia (Antonechloris) silvia Hausmann, 1991 


EXTERNAL FEATURES : Venation as figured for T! persica. Hindwing : Rs 
and M1 stalked, length of stalk 1/5-1/4 of distance between cell angle 
and termen ; M3 and CuAl sometimes slightly separated only. Tongue 
rudimentary, length 0.5-0.7 mm. Length of palpus & 1.6-1.8 times 
diameter of eye, ® twice diameter of eye. Frons flat, pale green, lower 
part white. Vertex whitish, with pale green scales. Antenna of male 
long bipectinate to 5/6-7/8 of length of flagellum, length of longest 
branches 0.80-0.95 mm (7-8 times width of flagellum); antenna of 
female shortly bipectinate, branches thicker than in 7: persica, length 
0.15-0.20 mm (1.5 times width of flagellum). Hindtibia of male with 
four short spurs, of female with two terminal spurs of unequal length. 


GENITALIA @ : see Fig. 54 (C. Jordan). Projections of uncus comparative- 
ly long and slender. Valva short and broad. Posterior edge of sternite 8 
broad. 


GENITALIA 9 : see Fig. 117 (C. Jordan). Vaginal plate almost round. 
Chitinisation of ostium bursae anteriorly rounded. 


REMARKS : Holotype (coll. SMNK) re-examined. There are significant 
and constant differences from congeneric species in wing pattern. 


Thetidia (Antonechloris) bilineata Hausmann, 1991 


EXTERNAL FEATURES : Venation as figured for T. persica. Forewing : 
R2-R5 and MI connate. Hindwing: Rs and MI stalked, length of 
stalk 1/5 of distance between cell angle and termen. Tongue rudimen- 
tary, length 0.6 mm. Length of palpus (©) twice diameter of eye. Frons 


29 


and vertex as described for T. silvia. Antenna of female shortly 
bipectinate, branches more slender than in 7. silvia, longest branches 
0.25 mm (twice width of flagellum). Hindtibia of female with two 
terminal spurs of unequal length ; proximal spurs present, but very 
rudimentary and almost invisible. 


GENITALIA @ : unknown. 


GENITALIA © : see Fig. 118 (C. Jordan). Vaginal plate laterally tapering, 
anteriorly forked. 


REMARKS : Holotype (coll. SMNK) re-examined. There are significant 
differences from congeneric species in wing pattern. 


subgenus AGLOSSOCHLORIS Prout, 1912, stat. n. 


DIFFERENTIAL FEATURES : Features as described for Thetidia, mainly 
distinguished by the absence of proximal spurs in the male hindtibia 
(only one terminal pair of short spurs present) ; however in some species 
rudimentary proximal spurs are visible ; female hindtibia with two 
terminal spurs which tend to be reduced. Tongue absent or rudimentary 
(the latter e.g. in some species from Afghanistan). Venation (compare 
Fig. 13) rather variable in all species. @ genitalia in the type species 
T: fulminaria (Lederer, 1870) very similar to the subgenera Thetidia and 
Antonechloris, in the other species projections of uncus long, narrow, 
widely separated from each other. ® genitalia: chitinous opening of 
ostium bursae situated on the anterior half of the vaginal plate. 


REMARKS: In all the generic features mentioned by Prout (1913 : 28) 
there tend to be transitional characters (to Thetidia and Antonechloris). 
Therefore I propose to reduce the status of Aglossochloris to subgeneric 
rank. In a similar way Viidalepp (1988 : 31, 181) considers Aglossoch- 
loris as invalid at generic rank. Nevertheless the possibility remains 
that in future larval morphology or other features will reveal valuable 
apomorphic characters. 


Thetidia (A glossochloris) euryrithra Prout, 1935 


EXTERNAL FEATURES : Venation (Fig. 13) similar to that of 7! bilineata. 
Forewing : R2-R5 and M1 connate or sometimes stalked. Hindwing : 
Rs and MI stalked, length of stalk 1/4-1/3 distance from cell angle 
to termen; M3 and CuAÏ very shortly stalked. Tongue absent. 
Structure of palpus as described for T. smaragdaria, third segment 
somewhat shorter ; length in male 1.5 times, in female twice diameter 


30 


of eye. Frons flat. Antenna of male bipectinate to 4/5 of length of 
flagellum ; longest branches about 0.8 mm (— about 7 times width of 
flagellum) ; antenna of female shortly bipectinate to 2/3 of length of 
flagellum ; longest branches 0.20-0.25 mm (= twice width of flagellum). 
Hindtibia of male with two spurs, of female sometimes with short 
(rudimentary) terminal spurs, sometimes without spurs. 


GENITALIA @: see Fig. 55 (N. Jordan). Projections of uncus long, 
slender, well separated from each other. 


GENITALIA 9 : see Fig. 119 (N. Jordan). Border of vaginal plate shallow- 
ly notched posteriorly ; vaginal plate laterally tapering and anteriorly 
forked. Chitinisation of ostium bursae narrow. 


Tribus TIMANDROMORPHINI Inoue, 1961 


DIFFERENTIAL FEATURES : Venation : Forewing : M3 and CuAl slightly 
separate. Hindwing: Rs and MI unstalked. Frenulum in & present, 
in @ lacking. Frons convex. Hindtibia of both sexes with four spurs. 
Abdominal crests absent. @ genitalia (main differential feature) : Uncus 
broad, with terminal excavation ; socius strongly sclerotised, broad at 
base, tapered at apex ; gnathos strongly sclerotised ; valva with spinu- 
lose harpe and cucullus ; sternite 8 round with two lateral projections 
(lobes). 


TYPE SPECIES of type genus Timandromorpha Inoue, 1944, examined. 


REMARKS : Not represented in the study area. Perhaps related to Nemo- 
riini. 


Tribus NEMORIINI Gumppenberg, 1887 


(Ochrognesiini Inoue, 1961, syn. n.) 


Nemoriini Gumppenberg, 1887 (and not “Ferguson, 1969”), originally 
proposed as “Nemorunae” with the (N. American) type genus Nemoria 
Hübner, [1818], available according to article 36 ICZN. Gumppenberg 
took into account not only the Palaearctic, but also the Nearctic fauna. 
Therefore the hypothesis, that Gumppenberg’s concept of “Nemoria” 
concerns the Palaearctic Chlorissa species belonging to the Hemitheini 
(hypothetic homonymy “Nemoria Gumppenberg, 1887, nec Hübner, 
[1818]”) has to be rejected and article 39 ICZN cannot be applied. 


DIFFERENTIAL FEATURES : Venation: Forewing: M3 and CuAl se- 
parate or connate. Hindwing: M3 and CuAl usually stalked (very 
rarely connate or separate); Rs and MI stalked. Frenulum present 


31 


in @, absent in female. Third segment of 2 palpus often smoothly 
scaled and distinctly elongate. Hindtibia of both sexes with four spurs 
(exceptionally with two hindtibial spurs in the Nearctic genus Chlorosea 
Packard, 1874). Abdominal crests absent. @ genitalia (main differential 
feature) : Uncus slender, fairly long, often forked terminally ; socius 
semi-membranous, short; valva slender; costa of valva strongly 
sclerotised, terminally slightly twisted inside ; basally often with processi ; 
posterior edge of sternite 8 with median excavation. 


TYPE SPECIES of the following additional genera examined : Ochrognesia 
Warren, 1894, Osteosema Warren, 1894, Chloromachia Warren, 1899, 
Lophomachia Prout, 1912, Anisozyga Prout, 1911, Chlorodontopera 
Warren, 1893 (the latter genus with Rs and MI of the hindwing 
unstalked). According to Ferguson (1985) and Pitkin (1993) Chlorosea 
Packard, 1874, Nemoria, Hiibner, [1818], Phrudocentra Warren, 1895, 
Dichorda Warren, 1900, Lissochlora Warren, 1900, Chavarriella Pitkin, 
1993, Paromphacodes Warren, 1897, Rhodochlora Warren, 1894, and 
probably many others from the Nearctic and Neoptropical regions 
belong here. 


REMARKS: Not represented in the study area. Genitalic and external 
morphology as stated and figured in Ferguson (1985) and Pitkin (1993) 
for the Nearctic and Neotropical Nemoriini reveal close relationships 
to the (above mentioned) Asiatic genera examined by the author. 
Therefore Ochrognesiini Inoue, 1961, must be placed in the synonymy 
of Nemoriini Gumppenberg, 1887. 


Tribus THALASSODINI Inoue, 1961 


DIFFERENTIAL FEATURES : Cell of hindwing very short, discocellular 
vein oblique. Abdominal crests wanting. 


REMARKS: Several species of type genus Thalassodes Guenée, 1857 
examined. Not represented in the study area. Thalassodes much 
resembles Prasinocyma (Hemistolini 4), in external structure, except 
stalked M3 and CuAl and oblique discocellulars of hindwing. However 
the genitalia are very different. It will be necessary to examine a lot 
of species to determine whether Thalassodes can be placed in Hemistolini 
or not. 


Tribus HEMISTOLINI Inoue, 1961 


DIFFERENTIAL FEATURES: Venation: Forewing: R2-R5 and MI 
typically stalked ; M3 and CuAl usually arising at same point or slightly 


32 


separate (exception: Xenochlorodes). Hindwing : Sc + RI and Rs 
fused or appressed at one point only (exception : Xenochlorodes) ; Rs 
and MI stalked ; M3 and CuAl usually connate with tendency to be 
shortly stalked. Frenulum often absent in both sexes, but present in 
male of Gnophosema, Prasinocyma, Victoria, Gonochlora, Aporan- 
dria. Palpus very slender and typically short in both sexes. @ antenna 
bipectinate. & hindtibia not dilated, number of spurs varying. Abdo- 
minal crests weak (Hemistola) or absent. Ansa of tympanum medially 
broadened, distally tapering. 5 genitalia with simple stout uncus, which 
is terminally forked in some species (perhaps indicating relationships 
to Nemoriini) ; socius semi-membranous and broad, often of medium 
length, but sometimes nearly as long as uncus or rudimentary (as in 
most Prasinocyma spp.) ; gnathos elongate, usually strongly sclerotised, 
indicating relationships to Comostolini/ Jodiini ; saccus typically simple, 
but forked in Xenochlorodes ; tergite 8 simple, sternite 8 usually forked 
posteriorly (only slightly in Lophostola ; tapered in Victoria). 9 geni- 
talia: Papillae anales and surface between characteristically folded 
(compare Figs. 120-126). 


TYPE SPECIES of the following genera examined : Lophostola Prout, 
1912, Prasinocyma Warren, 1897, Chlorocoma Turner, 1910, Anthar- 
mostes Warren, 1899, Lophorrhachia Prout, 1916, provisionally also 
Aporandria Warren, 1894, Gonochlora Swinhoe, 1904, Neurotoca 
Warren, 1897, Euproutia Fletcher, 1979. Some additional possibly 
related genera are given under the generic description of Victoria. 


REMARKS: The group of genera included here is probably poly- 
phyletic ; presumably monophyletic groups are Hemistola/ Lophostola/ 
Prasinocyma/ Chlorocoma and Victoria] Lophorrhachia/ Antharmostes. 
Aporandria in its external appearance resembles some genera of Geo- 
metrini (e.g. Ornithospila), however it is clearly different in the tym- 
panum and venation and therefore better included here. 


HEMISTOLA Warren, 1893 


DIFFERENTIAL FEATURES : Venation (compare Fig. 14) : Forewing : Sc 
often anastomosing with RI ; R2-R5 and MI typically stalked ; M3 
and CuAl usually connate or slightly separate. Hindwing: Rs and 
MI stalked. M3 and CuAl connate or slightly stalked (e.g. in type 
species). Frenulum absent in both sexes. Hindtibia of both sexes with 
four spurs. 4 genitalia: see description of tribe. Anterior margin of 
valva medially notched as in many Jodiini. Base of valvula with one 
more or less sclerotised processus. Coremata at the base of the valva 


33 


developed in some species including the type species, but modified to 
stoutly sclerotised spines in many others. ® Genitalia: corpus bursae 
pear-shaped, rather strongly sclerotised ; lamella antevaginalis often 
divided into two spinulose fields (very similar in Lophostola). 


REMARKS: The genitalic morphology of the generic type species, H. 
rubrimargo Warren, 1893, shows it to be closely related to H. chryso- 
prasaria (see below), but basal coremata of valva are present (instead 
of a basal spine). The genitalic morphology of Chlorocoma and some 
Prasinocyma species is rather similar to that of Hemistola. The loss 
of the frenulum in Hemistola (as compared with these species) is 
evidently not sufficient as a separating feature at tribal level. Loss of 
the frenulum must have taken place several times independently in 
the phylogenesis of Geometrinae (compare Ferguson, 1985 : 4). 


Hemistola chrysoprasaria (Esper, 1795) 


EXTERNAL FEATURES: Venation see Fig. 14 and descriptions of tribe 
and genus. Forewing: Sc and RI anastomosing for a short distance 
or appressed; R2-R5 and MI stalked or (exceptionally) connate. 
Hindwing : M3 and CuAl usually connate, but sometimes separated 
by a short distance or on short common stalk. Discocellular veins 
strongly curved. In the populations from S. Turkey (Taurus Mountains) 
R2-R5 and MI of forewing connate or separate. Tongue developed, 
thick, its length usually about 2-3 mm, sometimes to 4mm. Palpus 
very slender, length nearly diameter of eye or somewhat exceeding it 
(especially in Q). Frons flat, reddish brown. Vertex white. Antenna 
of & bipectinate nearly to tip, longest branches ca. 0.5 mm only (four 
times width of flagellum) ; antenna of © shortly bipectinate nearly until 
tip, longest branches ca. 0.2mm (somewhat exceeding width of 
flagellum). Little difference in the external structural features between 
the nominate subspecies and ssp. occidentalis Wehrli, 1929, ssp. lissas 
Prout, 1912, and the populations from Sicily (see remarks). 


GENITALIA @: see Fig. 56 (S. Turkey). Little difference between 
Specimens examined from N. and C. Italy, Macedonia and S. Turkey. 
In S. Turkey the membranous processus of valvula is narrower than 
in European populations. The ssp. (4) lissas from C. Asia has basal 
processus of valva much smaller than in the nominate subspecies, in 
addition the membranous processus of the valvula and the fields of 
cornuti in the aedeagus are very small, and sternite 8 has two rounded 
elevations posteriorly ; it may be a distinct species. 


34 


GENITALIA © : see Fig. 120 (S. Turkey). Almost no differences between 
the specimens examined from C. Italy and S. Turkey. Lamella post- 
vaginalis rounded in C. Italy, tapering to an angle of 90° in S. Turkey. 


REMARKS : In one specimen from C. Italy the venation is quite unusual 
(individual aberration) : M1 and M2 of forewing stalked, and Rs, M1 
and M2 of hindwing stalked! Genitalia of the populations from Sicily 
show specific differences (Hemistola siciliana Prout, 1935, stat. n.), e.g. 
basal processus of valva not connected with anterior margin of valva 
by sclerotised ridge (connected in H. chrysoprasaria) ; socius and 
membranous processus of valvula narrower ; aedeagus without fields 
of cornuti ; sternite 8 with two tapering projections well separated from 
each other (see Fig. 57). 


XENOCHLORODES Warren, 1897 


DIFFERENTIAL FEATURES : Venation see Fig. 15 (valid for X. olympiaria 
and X. nubigena (Wollaston, 1858)) : Forewing : both RI and R2 fused 
with Sc, distally not reaching costa; R2 exceptionally free; R1 and 
(short) common stalk of R2-R5 and MI connate, RI sometimes arising 
from distal end of common stalk or between; MI very rarely un- 
stalked ; M3 and CuAl stalked. Hindwing : Sc + RI and Rs fused 
over 2/3-3/4 length of cell; Rs and MI distinctly stalked ; M3 and 
CuAl stalked. Discocellular veins strongly angled. Frenulum absent 
in both sexes. Tongue developed, length ca. 2 mm (lost in Hissarica). 
Palpus slender and short. Eyes quite distant from each other. Frons 
flat, reddish brown. Vertex white. Antenna bipectinate in both sexes, 
branches in ® very short. Hindtibia in both sexes not dilated, with two 
long terminal spurs of unequal length (in Hissarica short spurs of nearly 
equal length). Tergite 2 of male interiorly with two long processi, which 
are often club-shaped ; their function totally unknown. @ genitalia : 
uncus and gnathos strongly sclerotised, pincer-shaped ; gnathos strong- 
ly tapering ; socius membranous, but very weakly developed ; saccus 
forked (rounded in Hissarica); valva ventrally excavated, without 
spinulose processi ; basal coremata present ; aedeagus long and slender, 
with some external spines ; posterior edge of sternite 8 bilobed with 
narrow sinus. © Genitalia rather simple : Lamella antevaginalis poorly 
sclerotised ; ductus bursae strongly sclerotised. 


REMARKS: The systematic position remains unclear. The venation is 
reminiscent of Comostolini. See remarks to Hierochthonia. The genus 
Hissarica Viidalepp, 1988, with type species Hissarica postalbida Vuda- 
lepp, 1988 (specimens examined from Afghanistan and Tadzhikistan), 


35 


is very near Xenochlorodes in venation and genitalia. The status of 
Hissarica must be downgraded to subgenus of Xenochlorodes (stat. n.). 
Xenochlorodes (Hissarica) gilvescens Wiltshire, 1966, is probably the 
right name and combination for the species called “ Hissarica postalbida” 
in Vudalepp, 1988. Further studies are required to verify this hypothesis. 
The type of Phorodesma graminaria Kollar, 1850, has to be examined 
with regard to possible relationships with X. (H.) postalbida. 


“ Xenochlorodes” albicostaria Brandt, 1938, obviously belonging to an- 
other genus, can provisionally be maintained in Xenochlorodes (Hissa- 
rica) ; one “Paratypoid” examined. 


Xenochlorodes olympiaria cremonaria (Staudinger, 1897) 


EXTERNAL FEATURES : For venation see generic description and Fig. 15 ; 
almost no variation between the populations from Spain, France, 
N. Africa, Italy, Greece, Turkey, Israel etc. (more than 50 specimens 
examined). Palpus reddish brown fading to ochreous, their length in 
& 0.40-0.50 mm only (0.8-0.9 times diameter of eye), in 9 0.50-0.60 mm 
(= diameter of eye). Antenna of & bipectinate nearly to tip, length 
of longest branches 0.50-0.60 mm (about five times width of flagel- 
lum) ; in Q very shortly bipectinate or almost filiform, “branches” not 
exceeding width of flagellum, length of cilia about 2/3 width of flagel- 
lum. Relative length tibia/ tarsus in the hindleg (49) 1.8/2.0 mm. Other 
features as described for the genus. 


GENITALIA @ : see Fig. 58 (N. Israel). Only very slight differences 
observed between the specimens examined from Israel, Lebanon and 
S. Turkey (X. 0. cremonaria) and those from Spain, Algeria, Tunisia, 
S.Italy, Croatia, S.Greece, Crete, N. Turkey (X. o. olympiaria) : In X. 
o. olympiaria projections of sternite 8 posteriorly somewhat more 
rounded. 


GENITALIA © : see Fig. 121 (N. Israel). No constant differences observed 
between the specimens examined from Israel, Lebanon, S. Greece, 
C. Italy and Morocco. 


Remarks: Xenochlorodes beryllaria (Mann, 1853), syn. to X. olym- 
piaria (Herrich-Schäffer, 1852) see Hausmann, 1997. The populations 
from S. Turkey, Lebanon and Israel differ from those of N. Turkey 
and Europe in their pale green (almost white) wing colour. All the 
deep green specimens from the Levant examined by the author proved 
to be Hierochthonia species. Two syntypes (“Original”) of cremonaria 
examined. 


36 


GNOPHOSEMA Prout, 1912 


DIFFERENTIAL FEATURES : For generic description see revision of Ebert 
(1969). Venation (compare Fig. 16) : Forewing : R2 usually fused with 
R1. Hindwing : M3 and CuAl on short common stalk or connate ; 
hindwing termen very slightly angled at both M1 and M3. Frenulum 
present in male, absent in female. Palpus in both sexes short and 
slender. Tongue absent. Antenna in both sexes bipectinate. Hindtibia 
in both sexes with two spurs, not dilated. & genitalia : uncus and gnathos 
strongly sclerotised ; socius present ; ventral part of valva medially 
excavated ; basal processus of valva strongly developed in G. isometra 
(Warren, 1888), small or absent in G. palumba ; basal coremata absent ; 
posterior edge of sternite 8 with two flat projections. 9 genitalia : Folds 
between papillae anales as in many other Hemistolini. 


REMARKS : Genitalic characters such as the long simple uncus, clearly 
show that Gnophosema is not a genus of “Pseudoterpnini” (as suggested 
in Ebert, 1969)! Certain relationships with Hemistola are indicated, 
particularly in the @ genitalia (structure of uncus, socius etc.). The 
external spines of the aedeagus and shape of the slender valva are 
reminiscent of those of Xenochlorodes. 


Gnophosema palumba kurdistana Ebert, 1969 


EXTERNAL FEATURES : see generic description and Ebert (1969 : 193ff.). 


GENITALIA @ : compare Fig. 59 (S. Iran ; G. p. palumba Brandt, 1938). 
Basal processus in subspecies from Kurdistan “totally reduced” longer 
in nominate subspecies. According to Ebert (loc. cit.) the chitinous 
ridge of the aedeagus has five teeth. Compare the figure of the right 
valva of G. p. kurdistana in Ebert (1969 : 183, Fig. 11). 


GENITALIA 9 : compare Fig. 122 (S. Iran ; G. p. palumba Brandt, 1938). 
© genitalia of G. p. kurdistana unknown. 

VICTORIA Warren, 1897 

subgenus VICTORIA Warren, 1897 


DIFFERENTIAL FEATURES : External morphology as described for Victo- 
rinella subgen. n. (see below), but palpus somewhat longer : slightly 
exceeding diameter of eye in 4, nearly twice diameter of eye in 9. 
Distinctly differing in & genitalia from the following subgenus : Uncus 
long, terminally broadened and slightly forked in V. fuscithorax ; socius 


37) 


broad and short ; spinulose harpe lacking, ventral projection of valva 
in V. gordoni probably not homologous to that of Victorinella, as in- 
dicated by lack of spines and different shape ; valva medially excavated 
on ventral margin (similar to equivalent in Hemistola) ; posterior edge 
of sternite 8 with two lobes (V. albipicta, V. immunifica) or two small 
projections, situated very near to each other (V. fuscithorax, V. gordoni, 
but rudimentary in the latter). 


SPECIES INCLUDED : Victoria albipicta Warren, 1897 (type species), and 
Victoria fuscithorax Warren & Rothschild, 1905 ; provisionally also 
Victoria immunifica Prout, 1912, and Victoria gordoni, Prout, 1912. 


REMARKS: The structure of uncus, length of socius and shape of 
posterior edge of sternite 8 in V. fuscithorax correspond well to morpho- 
logical characteristics of Nemoriini ; Victoria perhaps links both tribes. 
Venation almost identical with that of Bustilloxia (except stalked M1 
of forewing) ; the concave hindwing termen between MI and M3 and 
red frons are reminiscent of Thalerini. The number of spurs on the @ 
hindtibia is complete as in most species of the tribes Hemistolini/ 
Comostolini. The basal coremata of the valva are as e.g. in Lophostola. 
The systematic position has not been ascertained, but there may be 
further relationships to some African genera such as Prosomphax 
Warren, 1911, Celidomphax Prout, 1912, Omphax Guenée, 1857, and 
Heterorachis Warren, 1898. Victoria immunifica Prout, 1912, and 
Victoria gordoni, Prout, 1912, differ from both subgenera Victoria and 
Victorinella in their male genitalia with ventral part of valva basally 
membranous (modified coremata), tongue present etc. Perhaps they 
constitute a well defined third subgenus of Victoria. 


subgenus VICTORINELLA subg. n. 


TYPE SPECIES : Victoria sematoperas Prout, 1916, by original (= present) 
designation. 


DIFFERENTIAL FEATURES : See Hausmann (1993 : 53f.). Venation see 
Fig. 17 (V. sematoperas) : Venation of forewing quite simple : Sc free ; 
RI free; R2-R5 stalked ; R2-R5 and MI connate; M3 and CuAl 
connate. Hindwing : Sc + RI and Rs fused at one point; Rs and MI 
stalked ; M3 and CuAl stalked. 4 frenulum strongly developed, absent 
in Q. Palpus comparatively short in both sexes. Tongue lacking. 
Antenna bipectinate in both sexes. Abdomen with dorsal crests. Hind- 
tibia with four spurs in both sexes. @ genitalia: Uncus shorter than 
in Victoria s.str., tapering to end, not forked terminally ; length of 
socius nearly that of uncus ; spinulose harpe present ; posterior edge 


38 


of sternite 8 with one central spinulose projection (simple in V. triplaga). 
© genitalia: Papillae anales transversally folded (similar to equivalent 
in Hemistola). 


FURTHER SPECIES EXAMINED: V. sematoperas Prout, 1916, V. oma- 
nensis (Wiltshire, 1980), V. triplaga Prout, 1915. 


REMARKS : Victorinella corresponds better in its genitalic morphology 
(4) to some species of the African genus Antharmostes Warren, 1899 
(e.g. A. interalbicans Warren, 1902) than to Victoria s.str.. However 
in Antharmostes the gnathos is weaker, tongue present, female antenna 
simple, hindtibia of male dilated and hind wing termen at M3 distinctly 
prominent (not or only slightly so at M1). 


Victoria (Victorinella) wiltshirei sp. n. 


Victoria sematoperas Hausmann, 1993 : Spixiana 16 (1) : 59 (nec Prout, 1916: 
Proc. Zool. Soc. London, 1916 : 142, pl. II, Fig. 26). 


HoıotyprE: 6, NE. Sudan, Ed Damer, Hudeiba, 24.1.1962, leg. 
Remane, coll. ZSM. 


EXTERNAL FEATURES : Holotype figured in Hausmann (1993: Fig. 4; 
“Victoria sematoperas (?)”). Wing colour green. White cell spot of all 
wings larger than in V. sematoperas, encircled by a broad red-brown 
ring. Apical spot of forewing large, round (in V. sematoperas much 
smaller and more or less subdivided into 3 spots). Spots in the anal 
angle of both fore and hindwing larger than in V. sematoperas. Post- 
median line present, but hardly visible (in V. sematoperas only one cross- 
line very near the termen). Venation exactly as figured for V. semato- 
peras (Fig. 17). Length of palpus 2/3 diameter of eye, ochreous, second 
segment with outstanding scales, third segment extremely short. Length 
of longest branches of & antenna 0.92 mm (8 times width of flagellum). 
Hindtibia (3) apparently with two proximal spurs of unequal length 
and one terminal spur (both legs present and examined), but this may 
be an artifact. Distance between terminal and proximal spurs very small. 


GENITALIA @ see Fig. 60 (NE. Sudan) : Similar to that of V. semato- 
peras (compare Fig. 61). Harpe oval. Spines situated more medially 
than in V. sematoperas. Posterior edge of sternite 8 not tapering as 
in V. sematoperas, less sclerotised and with teeth less concentrated. 


GENITALIA © : unknown, but presumably similar to genitalia of V. 
sematoperas (compare Fig. 123). 


REMARKS: The author was able to borrow a male from Herr Kuchler, 
München, of true V. sematoperas from Kaputir, W. Kenya, “near” the 


39 


locus typicus of the species (Mandere, NE. Kenya), which corresponds 
fairly well in external appearance to the original description and is 
genitalically identical with the holotype. Dedicated to Mr. E. P. Wilt- 
shire, who sent me drawings of the genitalia of the @ holotype and 
Q paratype of V. sematoperas, and kindly helped also by his extensive 
scientific discussions on the genus Victoria. 


Victoria (Victorinella) plantei Herbulot, 1976 


EXTERNAL FEATURES : This and the following species are characterised 
by smoothly scaled second segment of palpus (3). Third segment of 
Q palpus somewhat elongate ( 0.3, & 0.1 mm). 


GENITALIA @ see Fig. 62 (C. Israel), of © see Fig. 124 (C. Israel). 


REMARKS: Holotype re-examined by M. Herbulot, Paris (personal 
communication). For data on the morphology and differential features 
from the other species of the genus see Hausmann (1993 : 53f.). 


Victoria (Victorinella) eremita Hausmann, 1993 


GENITALIA @ see Fig. 63 (S. Israel). New material collected in April 
confirms that the genitalia of specimens emerging in April are similar 
to those of V. plantei. Rearing experiments are necessary to resolve 
the problem of possible crossfertilisation between the various “seasonal 
units” of V. eremita (compare Hausmann, 1993) and V. plantei. 


GENITALIA Q see Fig. 125 (S. Israel). 


REMARKS: Holotype re-examined. Data on the morphology and 
differential features from the other species of the genus see Hausmann 
(CIO Bi ESS) 


Victoria (Victorinella) fifensis Wiltshire, 1994 


GENITALIA @ see Fig. 64 (W. Saudi Arabia). 

REMARKS: Data on the morphology and differential features from 
the other species of the genus see Hausmann (1993 : 59 ; as “Victoria 
spec.”) and Wiltshire (1994 : 115f.). 

Tribus RHOMBORISTINI Inoue, 1961 


DIFFERENTIAL FEATURES : Venation: Forewing: M3 and CuAl se- 
parate. Hindwing: venation corresponding well with that of Berta 


40 


(Jodiini) and Comostola (Comostolini) particularly in the extraordinary 
course of the discocellular vein. Frenulum present in @, absent in 9. 
Tongue present. Palpus elongate, third segment of 9 palpi very long. 
Antenna of @ and © as described for Comostolini. Abdominal crests 
absent. Hindtibia with two spurs in 4, with four in Q. 


TYPE SPECIES examined of Rhomborista subgenus Spaniocentra. 


Remarks: Not represented in the study area. Closely related to both 
Comostolini and Jodiini. Some correspondences in SQ genitalia. 


Tribus COMOSTOLINI Inoue, 1961 


DIFFERENTIAL FEATURES: Venation: In Comostola, Comostolopsis 
and Pyrrhorachis R1-R5 and MI of forewing stalked, in Eucrostes 
and Mujiaoshakua however M1 arising below cell apex ; M3 and CuAl 
in both wings stalked; Rs and M1 of hindwing stalked ; hindwing 
termen not or slightly prominent on M3. Frenulum absent in both 
sexes. Tongue generally present. Palpus very slender; in female 3rd 
segment often elongate. Male antenna bipectinate, last 1/4-1/2 simple 
and ciliate. Female antenna filiform (very shortly bipectinate in 
Eucrostes). Hindtibia with four spurs in both sexes (two in Eucrostes), 
tarsus short. Ansa of tympanal organ very long and narrow. Abdominal 
crests wanting. @ genitalia: Uncus long, simple, or forked at apex ; 
gnathos present, tapering, often long ; basal coremata of valva absent 
(exception : Pyrrhorachis) ; base of valva with spinulose lobes ; aedeagus 
long and slender ; sternite 8 and tergite 8 usually simple. © genitalia 
often with signum. 


TYPE species of following additional genera examined : Comostola 
Meyrick, 1888, and Comostolopsis Warren, 1902 (perhaps should be 
included in Comostola, compare Inoue, 1961: 101), Pyrrhorachis 
Warren, 1896. According to Inoue (1961 : 79) Mujiaoshakua Inoue, 
1955 also belongs here. 


REMARKS: Some features are reminiscent of Hemistolini (e.g. 4 
genitalia). A rather close relationship to (and monophyly with) Jodun 
is indicated by the structure of antenna, hindtarsus etc., see below. 
In Syndromodes & genitalia are similar, e.g. uncus and tip of gnathos 
long and slender, basal coremata of valva absent etc., but relationship 
to Comostolini is not confirmed (male frenulum present; see also 
Microloxiini). Allochrostes Prout, 1912, despite similar external ap- 
pearance does not belong to Comostolini (perhaps near Microloxiini). 


41 


EUCROSTES Hübner, [1823] 


DIFFERENTIAL FEATURES: Venation see Fig. 18: Forewing: Sc and 
RI fused ; Ri arising from below cell apex ; R2-R5 and MI usually 
separate ; M3 and CuAI stalked. Hindwing : Sc + RI and Rs touching 
(not fused) at one point; Rs and MI usually stalked ; M3 and CuAl 
stalked ; discocellulars of all wings often weak between M2 and M3. 
Frenulum absent in both sexes. Frons flat. Palpus very slender, second 
and third segment elongate in ©. Tongue present, but weak. @ antenna 
bipectinate with long branches, © with very short branches or in some 
species flagellum filiform. Hindtibia in both sexes with two spurs, not 
dilated. 4 genitalia: Uncus slender, terminally forked ; socius semi- 
membranous, long; gnathos distinctly sclerotised, slender, tapering ; 
valva with chitinous crest instead of basal coremata ; aedeagus long, 
slender, without cornuti; sternite and tergite 8 simple. © genitalia: 
Apophyses posteriores 2.0-2.3 times length of apophyses anteriores (the 
latter 0.25 mm only) ; corpus bursae rather small and tender ; lamella 
antevaginalis sclerotised to a longitudinally folded ribbon; ostium 
bursae funnel-shaped. 


FURTHER SPECIES EXAMINED : Eucrostes disparata Walker, 1861 (Nige- 
ria, Tanzania). 


REMARKS: The male genitalia are strikingly similar to Comostolopsis ! 
The relationship is unquestionable, although forewing venation and 
some other features are quite different. 


Eucrostes indigenata (de Villers, 1789) 


EXTERNAL FEATURES: Venation (Fig. 18) see generic description : 
Forewing : R2-R5 and MI separate or connate, exceptionally on very 
short common stalk ; M3 and CuAl stalked ; forewing venation quite 
constant throughout the area of distribution. Hindwing: Rs and MI 
in West Mediterranean populations shortly stalked or sometimes 
connate ; distinctly stalked in populations from Tunisia, Sicily, S. Italy 
and East Mediterranean with only a very few exceptions, transitional 
in C. Italy. Frons reddish brown or orange, vertex white with some 
yellowish or greenish scales. Length of weak tongue 1.2-2.0 mm. Palpus 
whitish ochreous, their length in & 0.50-0.60 mm (= diameter of eye), 
in the Middle East populations somewhat exceeding diameter of eye ; 
in © 0.70-0.80 mm (nearly 1.5 times diameter of eye), first segment 
bushy-scaled, third segment “naked” with adherent scales. Antenna of 
& bipectinate over 2/3 of length of flagellum, length of longest branches 


42 


0.42-0.50 mm (4-5 times width of flagellum), in ® very shortly bi- 
pectinate (almost lamellate) over 1/2-2/3 of length of flagellum, length 
of longest branches 0.12-0.15, exceptionally 0.17 mm (slightly exceeding 
width of flagellum at same point). In hindleg of both sexes relative 
length tibia/ tarsus 1.2-1.7/1.5-2.0 mm. 


GENITALIA @: see Fig. 65 (N. Jordan) and generic description. No 
remarkable and constant difference between specimens examined from 
S. Spain, S. France, C. Italy, Sicily, Malta, Tunisia, Hungary, Crete, 
Lebanon, Syria, Israel and Jordan. 


GENITALIA © : see Fig. 126 (Israel) and generic description. No re- 
markable and constant difference between specimens examined from 
S. Spain, C. Italy, Sicily, Tunisia, Turkey, Lebanon, Israel and Jordan. 


REMARKS : The close relative E. disparata differs in (absolute) length 
of antennal branches (@ : about 0.35-0.40 mm = four times width of 
flagellum), and in the hindwing where Rs and MI are distinctly stalked 
as in East Mediterranean populations of E. indigenata. In the @ geni- 
talia the valva is somewhat shorter. Perhaps only separable at sub- 
specific rank. At the ZSM there are specimens of E. indigenata from 
S. Spain that differ from the nominate subspecies. These populations 
are probably isolated from the rest of the area of distribution, since 
the species is not recorded from Morocco and W. Algeria in the south ; 
in the north the Pyrenees could be another barrier : 


Eucrostes indigenata lanjeronica ssp. n. 


HoLotypPe: 9, Hispania mer., Andalusia, Lanjeron, 11.V1.1895, leg. 
Korb, coll. ZSM. 


PARATYPES : 14, id. ; 14, id., “1894”; 19, Hispania mer., Las Alpujarras, 
Orgiva, 1000 m, 13.V11.1962, leg. Sattler, coll. ZSM ; 14, Hispania 
mer., Andalusia, Ronda, 26.VI.1926, leg. Wehrli, coll. ZFMK; 16, 
id., 26.VIL.1926. One further female from “Castilia” (C. Spain; coll. 
ZSM) has a wrongly attached head (of a Chlorissa sp.), therefore is 
not included in the type series. 


DESCRIPTION (Fig. 148) : Length of the antennal branches in 9 : 0.22- 
0.27 mm (2.5 times width of flagellum) ; other external features and 
venation as described above for West Mediterranean populations. 
Yellowish postmedian line somewhat more distal than in E. i. indigenata. 
No distinguishing characters in genitalia (IP). 


43 


REMARKS : Females of the nominate subspecies have been examined 
from SE. France, Majorca, Italy, Sicily, Malta, Tunisia, Hungary, 
Croatia, Bosnia, Macedonia, Greece, Crete, Bulgaria, Turkey, Lebanon, 
Israel, Jordan. 


Tribus JODIINI Inoue, 1961 


DIFFERENTIAL FEATURES : Most features as described for Comostolini. 
However hindwing termen prominent at M3. Frenulum usually absent, 
but present in Gelasma (plesiomorphy 4). Both @ and © palpus 
elongate. Hindtarsus short. Abdominal crests wanting. @ genitalia : 
Valva slender, typically with ventral excavation ; basal coremata of 
valva absent in Jodis, but present in Gelasma and Berta (plesiomorphy 
G). Posterior edge of sternite 8 typically, but not always, strongly 
sclerotised with median notch. 


TYPE SPECIES of the following genera examined : Jodis Hübner, [1823], 
Gelasma Warren, 1893 (Gelasma s.str., Thalerura Warren, 1894), Berta 
Walker, [1863]. 


REMARKS: Not represented in the study area. Gelasma shows some 
presumably plesiomorphic features ; relationships with various other 
tribes (e.g. Hemitheini) are not excluded, perhaps a genus of great 
evolutionary age. Gelasma belongs to Hemitheini according to Viidalepp 
(1981 : 95). 


Tribus THALERINI Herbulot, 1963 
(Chlorochromini Duponchel, [1845], nom. invalid.) 


DIFFERENTIAL FEATURES: Venation: Forewing: R2-R5 and MI 
usually stalked. Hindwing : Rs and MI stalked ; M3 and CuAl usually 
unstalked ; margin of hindwing strongly concave between M1 and M3. 
Frenulum absent in both sexes (present in male of Culpinia and Bustil- 
loxia). Frons always red, orange or reddish brown. Palpus very slender. 
GQ antenna usually bipectinate. Hindtibia with two terminal spurs in 
both sexes (female exceptionally with four spurs : Culpinia). genitalia : 
Uncus simple ; socius present, long, sclerotisation often comparatively 
strong ; gnathos weak or absent ; valva long and slender ; costa medially 
with strongly sclerotised processus ; a second processus at base of valva 
(exception: Bustilloxia) ; basal coremata of valva present ; sternite 8 
and tergite 8 simple (exception : Bustilloxia). Q genitalia : Sclerotisations 
of lamella ante- and postvaginalis ring-shaped. 


AA. 


TYPE SPECIES of following additional genera examined : Dyschloropsis 
Warren, 1895 ; Heteroculpinia Hausmann, 1994, Dolosis Prout, 1912. 
Bustilloxia Exposito, 1979, has some features indicating relationships 
to Thalerini (compare Hausmann, 1994), but its systematic position is 
isolated (e.g. in @ genitalia transtilla differs strongly from that of typical 
Thalerini species). 


REMARKS: The tribes Hemistolini, Comostolini, Jodiini, Thalerini, 
Hemitheini and Microloxiini are linked by various (mostly exotic) 
genera, which show combinations of transitional features. E.g. in 
Culpinia the number of female hindtibial spurs and the presence of 
male frenulum correspond well with typical Hemitheini features, but 
genitalia, wing colour and hindwing margin reveal close relationships 
to Thalerini. 


THALERA Hübner, [1823] 


DIFFERENTIAL FEATURES: Venation compare Fig. 19 and tribal de- 
scription. Forewing: RI often fused with R2 for a short distance ; 
R2-R5 and MI stalked ; M3 and CuAl distinctly separate. Hindwing : 
Sc + R1 and Rs fused over about 1/4 of length of cell (not fused 
in all other Thalerini genera) ; M3 and CuAl unstalked as in Dyschlo- 
ropsis. Tongue present (absent in Dyschloropsis). Fringes chequered. 
& genitalia : saccus elongate. Q genitalia : Apophyses anteriores in type 
Species comparatively long: 0.6mm = 1/2 of length of apophyses 
posteriores (shorter in Dyschloropsis and Heteroculpinia) ; corpus 
bursae very long and slender. 


REMARKS: T. lacerataria Graeser, 1889, has somewhat different fore- 
wing venation : RI fused with Sc ; R2-R5 and MI connate. 


Thalera fimbrialis magnata A. F uchs, 1903 


EXTERNAL FEATURES : Venation see Fig. 19, compare also tribal and 
generic description : Hindwing : CuAl and M3 arising at same point 
or separated by a short distance only (exceptionally shortly stalked). 
Tongue developed, length 3.2-4mm. Palpus (@@) slender, short 
(nearly = diameter of eye). Frons flat. Antenna of male bipectinate 
to tip, longest branches 0.8-0.9 mm (five times width of flagellum) ; 
antenna of female shortly bipectinate almost to tip, longest branches 
0.3 mm (twice width of flagellum), in Greek populations about 0.4 mm. 
High constancy of external structural features throughout the area of 
distribution of both 7! f. fimbrialis and T: f. magnata. 


45 


GENITALIA @ : see Fig. 66 (C. Turkey). There are almost no remarkable 
and constant differences between the European and the Asiatic popu- 
lations (specimens examined from Germany, S. Italy and Macedonia ; 
C. Turkey, N. Iran, C. Asia) : In the latter valva slightly narrower. 


GENITALIA © : see Fig. 127 (N. Iran). No differences within the area 
of distribution. 


REMARKS :Perhaps T: fimbrialis var. magnata A. Fuchs, 1903, should 
be better considered as a synonym of Thalera fimbrialis. Specimens 
from the Asiatic populations are said to be larger than the nominate 
ones ; but this is a very slight difference with many transitions and 
exceptions. 


CULPINIA Prout, 1912 


DIFFERENTIAL FEATURES : Venation see Fig. 20 : Forewing : R1 usually 
fused with Sc for short distance. Hindwing : Sc + RI and Rs touching 
at one point, not fused ; M3 and CuAl shortly stalked as in Hetero- 
culpinia. Frenulum present in male, absent in female. Hindtibia of male 
with two, of female with four spurs. Fringes chequered (not in Dyschlo- 
ropsis and Heteroculpinia). Q genitalia: Apophyses anteriores com- 
paratively long (0.6 mm = half length of apophyses posteriores) ; corpus 
bursae comparatively short, rounded. 


REMARKS: The genus has many transitional features linking Thalerini 
with Hemitheini. 


Culpinia prouti (Thierry-Mieg, 1913) 


EXTERNAL FEATURES: Venation see Fig. 20: Forewing: RI often 
free ; R2-R5 and MI shortly stalked (in type species Culpinia diffusa 
(Walker, 1861) usually connate) ; M3 and CuAl of both wings on short 
common stalks. Tongue developed, length ca. 3.5-5 mm (absent in the 
type species and Dyschloropsis). Palpus slender, its length somewhat 
exceeding diameter of eye in male, nearly twice diameter of eye in 
female (Q palpus not elongate in type species) ; first segment bushy- 
scaled. Frons flat, reddish brown. Antenna of male bipectinate nearly 
to tip, length of longest branches ca. 0.6 mm (four times width of flagel- 
lum); of female very shortly bipectinate, length of longest branches 
ca. 0.1 mm (nearly = width of flagellum). Proximal pair of spurs on 
female hindtibia rudimentary. No structural variation within the area 
of distribution. | 


46 


GENITALIA @ : see Fig. 67 (S. Turkey). 


GENITALIA 9: see Fig. 128 (C. Israel). No remarkable difference 
between the specimens examined from S. Turkey and C. Israel. 


REMARKS: The @ genitalia seem to show a closer relationship to 
Thalera lacerataria Graeser, 1889 and to the genus Heteroculpinia, 
than to the type species of the genus, Culpinia diffusa, but the male 
frenulum and structure of the © hindtibia do not allow the species 
to be transferred. 


Tribus HEMITHEINI Bruand, 1846 


DIFFERENTIAL FEATURES: Venation: Hindwing: Sc + RI and Rs 
fused or appressed at one point ; Rs and MI stalked ; M3 and CuAl 
usually stalked. Frenulum present in male, absent in female ; according 
to Prout (1912: 21) “@ frenulum weak”. Palpus of female often 
elongate. Male with one pair of metathoracic tibial spurs ; female with 
two pairs (e.g. most Chlorissa species, Phaiogramma, Hethemia) or 
one pair distally (some “Chlorissa” species, Kuchleria, Pamphlebia, 
Neromia and Mesothea). Both @ and © antenna usually filiform, ciliate, 
but bipectinate in @ of Kuchleria, Nipponogelasma, Chlorochlamys, 
Chloropteryx and Xerochlora, in 9 of Kuchleria. & genitalia : Uncus 
simple, comparatively long ; gnathos weak or absent ; length of semi- 
membranous socius about that of uncus ; transtilla often narrow and 
U-shaped as in Thalerini, but sometimes extended ; valva with basal 
coremata ; aedeagus divided into a slender basal “stalk” and a broad 
distal half, very often with cornuti or sclerites ; sternite 8 in male 
sometimes with specific characters ; sternite 2 typically with lateral 
apophyses, which are strongly sclerotised, slender and tapered distally. 


TYPE species of the following additional genera examined : Hemithea 
Duponchel, 1829, Diplodesma Warren, 1896, Pamphlebia Warren, 
1897. According to Inoue (1961 : 57) and Viidalepp (1981 : 95) Nippo- 
nogelasma Inoue, 1946, and Parachlorissa Inoue, 1961 also belong here. 
According to Ferguson (1985 : 102f.) the N. American genera Chloro- 
chlamys Hulst, 1896, Chloropteryx Hulst, 1896, Xerochlora Ferguson, 
1969, Hethemia Ferguson, 1969, and Mesothea Warren, 1901, belong 
to Hemitheini. 


REMARKS: In the literature “Hemitheini” has often been treated as 
a polyphyletic “tribe” in which authors include those genera that cannot 
be placed in the other established tribes. Hemitheini (as defined here) 
is rather closely related to Thalerini. E.g. genitalic morphology of Diplo- 


47 


desma, Nipponogelasma, Hethemia and Mesothea reveal transitional 
features (mainly to Culpinia). 


CHLORISSA Stephens, 1831 


DIFFERENTIAL FEATURES : see Dantart (1990). Venation see Fig. 21: 
Forewing: Sc and RI often fused, length of fusion variable, excep- 
tionally Ri free ; R1 free from R2-R5. R2-R5 and MI shortly stalked, 
exceptionally connate or only separated by a short distance ; M3 and 
CuAl often stalked. Hindwing: M3 and CuAl always distinctly 
stalked ; termen at M3 usually angled. Palpus of female long. Antenna 
of both male and female filiform without pectinations, ciliate. Hind- 
tarsus of male reduced, except Chlorissa gigantaria (Staudinger, 1892) 
according to Viidalepp (1988, Fig. 1/4). & genitalia with socius very 
narrow ; transtilla U-shaped, narrow; processus of costa of valva 
prominent ; aedeagus with sclerites ; sternite 8 with one more or less 
prominent, strongly sclerotised median tooth. Q genitalia: Lamella 
antevaginalis a single plate ; osttum bursae posteriorly with two hairy 
lobes. 


FURTHER SPECIES EXAMINED : C. pretiosaria Staudinger, 1877, C. gelida 
Butler, 1889, C. gigantaria Staudinger, 1892, C. talvei Vudalepp, 1988, 
C. arkitensis Viidalepp, 1988, C. rubripicta (Warren, 1893) and C. 
distinctaria (Walker, 1866). 


REMARKS: The genera Hemithea and Diplodesma are closely related 
to Chlorissa. “Chlorissa should perhaps be sunk to subgeneric rank 
of Hemithea” (Prout, 1930 : 27). Some genitalic features of Hemithea 
(e.g. narrow socius, contents of aedeagus) lead to the opinion, that 
Hemithea/Chlorissa are more closely related to each other than to 
Phaiogramma] Chlorissa. Some other features, e.g. shape of valva and 
harpe, hindwing termen, structure of female genitalia confirm the 
generic distinction between Chlorissa and Hemithea. The palpus and 
antenna of the C. pretiosaria group are somewhat similar to those 
of Phaiogramma. In some African species RI-R5 of forewing stalked, 
with R1 fused to Sc. According to the generic diagnosis given by Prout 
(1934 : 117) they have to be transferred to Diplodesma : Diplodesma 
approximans (Warren, 1897), comb. n., Diplodesma subrufibasis (Prout, 
1930), comb. n., Diplodesma eborilitoris (Fletcher, 1958), comb. n. nec 
(Prout, 1930 : unavailable) and probably many others. 


48 


Chlorissa viridata (Linnaeus, 1758) 


EXTERNAL FEATURES : Venation see Fig. 21 : As described for the 
subgenus. Tongue developed, length 3-3.5 mm. Length of palpus of 
male somewhat exceeding diameter of eye, of female up to 1.5 times 
diameter of eye. Frons slightly convex. Antenna of male and female 
filiform, ciliate to tip of flagellum, length of longest cilia in male 
0.09 mm (= width of flagellum), in female about 0.06 mm (2/3 width 
of flagellum). Relative length tibia/tarsus in the foreleg (@Q) 1.3/ 
2.3mm; in the hindleg 3.0/1.2 mm (8) or 2.3/1.5 mm (©). Male 
hindtibia slightly dilated. Little structural variation within the area of 
distribution : in the Central Asian subspecies C. v. insigniata (Stau- 
dinger, 1901) R2-R5 and MI of forewing always separate ; in Trans- 
caucasian and Georgian specimens (n=2 X 5: both wings examined) 
R2-R5 and MI of forewing connate, M3 and CuAl separate, in the 
hindwing discocellular vein joining M3 + CuAl very near CuA2. 


GENITALIA @ : see Figs. 68-69 (N. Italy, Transcaucasia). For distin- 
guishing characters from C. cloraria see Dantart (1990). Terminal 
sclerotisation of vesica short, well separated from the patch of micro- 
cornuti by a hyaline area of the vesica. Specimens examined from S. 
Germany, N. Italy, Macedonia, Armenia, Transcaucasus, C. Asia. In 
Transcaucasian specimens shape of processus of costa of valva variable 
and transitional to C. cloraria as it is also in C. Asiatic C. v. insigniata. 


GENITALIA Q : see Fig. 129 (Transcaucasia). For distinguishing cha- 
racters from C. cloraria see Dantart (1990). Specimens examined from 
S. Germany, N. Italy, Armenia, Transcaucasus, C. Asia. 


REMARKS : Not represented in the study area, but treated here as the 
Species occurs in Transcaucasia, and because separation from the 
following species is difficult. The Transcaucasian populations however 
need taxonomic revision with more extensive material. Perhaps sepa- 
ration of the two taxa (viridata/cloraria) is not complete in some 
regions. 


Chlorissa cloraria (Hübner, [1813]) 


EXTERNAL FEATURES : In venation, structure of tongue, palpus, frons, 
antenna and hindtibia as described for C. viridata. In the populations 
examined from C. Turkey R2-R5 and MI of forewing distinctly stalked, 
M3 and CuAl (forewing) shortly stalked. Last segment of palpus often 
darker. Apart from the venation little structural variation within the 
area of distribution. 


49 


GENITALIA @ : see Figs. 70-71 (C. Greece, C. Turkey). For distinguishing 
characters from C. viridata see Dantart (1990). Terminal sclerotisation 
of vesica often long and tapering, touching the patch of microcornuti 
(with exceptions). Specimens examined from Austria, Corsica, C. Italy, 
N. and C. Greece and C. Turkey. For Spanish populations see Dantart 


(lee): 


GENITALIA 9 : see Fig. 130 (C. Greece). For distinguishing characters 
from C. viridata see Dantart (1990). Specimens examined from Corsica, 
C. and S. Italy, N. and C. Greece. For Spanish populations see Dantart 
(l.c.). Determination may be impossible even after genitalic dissection. 


REMARKS: see remarks to C. viridata. 
Chlorissa asphaleia Wiltshire, 1966 


EXTERNAL FEATURES : Venation as figured for C. viridata, but R1 of 
forewing free from both Sc and R2-R5; R2-R5 and MI connate. 
Length of tongue 4-4.5 mm. Last segment of palpus of male very short ; 
length of palpus 1.7 times diameter of eye, in female somewhat longer 
(twice diameter of eye). Frons slightly convex. Antenna as described 
for C. viridata, but longest cilia in female about half width of flagellum. 
Relative length tibia/tarsus in the foreleg (3%) 1.3/3.0 mm; in the 
hindleg 3.0/1.2 mm (@) or 2.8/2.1 mm (©). 


GENITALIA @ : see Fig. 72 (N. Turkey). The big rounded basal pro- 
cessus, costal projection on distal end of valva, and large basal core- 
mata are unique within the genus. Reminiscent of Thalerini, e.g. genus 
Thalera. 


GENITALIA @ : see Fig. 131 (N. Turkey). Sclerites of lamella antevaginalis 
and surface between papillae anales reminiscent of Thalerini, e.g. genus 
Thalera. 


REMARKS: Not yet recorded in the study area. 
PHAIOGRAMMA Gumppenberg, 1887, stat. n. 


TYPE SPECIES : Nemoria faustinata Milliere, 1868, by subsequent mono- 
typy. 

DIFFERENTIAL FEATURES: Venation (see Fig. 22) similar to that of 
Chlorissa, but hindwing termen somewhat more rounded, less angled 
at M3. Palpus of female extremely long. Antenna of male dentate ; 
cilia grouped in tufts ; 9 antenna scarcely ciliate. & hindtibia with two, 


50 


© with four spurs. @ hindtarsus short as in Chlorissa : relative length 
tibia/tarsus about 2.7/1.5 mm. @ genitalia with socius rather broad, 
especially subapically ; transtilla expanded ; processus of costa of valva 
absent or small; aedeagus without chitinous plates, with patches of 
cornuti; posterior edge of sternite 8 slightly concave. ® genitalia: 
Lamella antevaginalis divided into two sclerites ; surface somewhat 
wrinkled. Main differential characters from Chlorissa : shape of socius, 
aedeagus and sternite 8 (@). 


ADDITIONAL SPECIES : The whole Afrotropical species complex sur- 
rounding C. faustinata/ C. stibolepida (Butler, 1879) has to be included 
in Phaiogramma. 


REMARKS : Compare remarks to Neromia and Neromia simplexa. Until 
now Phaiogramma has been regarded as a synonym of Chlorissa. 


Phaiogramma pulmentaria (Guenée, 1857), comb. n. 


EXTERNAL FEATURES: Venation (see Fig. 22) similar to that of C. 
viridata, but in the forewing RI typically free (in a very few specimens 
however fused with Sc) ; R2-R5 and MI usually connate, sometimes 
shortly stalked or separated from each other, in S. Spain, Morocco, 
Italy and Sicily always stalked, in Turkey usually stalked but some- 
times connate ; M3 and CuAl of forewing usually shortly stalked, but 
variable. Length of tongue 3.5-4 mm. Palpus of male long, length about 
1.5-2 times diameter of eye ; of female very long : 2.0-2.6 times diameter 
of eye. Frons flat. Antenna of male dentate almost to tip, longest cilia 
about 0.15 mm (1.5-1.7 times width of flagellum) ; of female filiform, 
scarcely ciliate, longest cilia only 1/3 width of flagellum. Hindtibia (49) 
as described for C. viridata. Apart from the venation no structural 
variation occurs within the area of distribution. 


GENITALIA 6: see Fig. 73 (N. Israel). No remarkable and constant 
differences between populations examined from S. Italy, S. Greece, 
Cyprus, S. Russia, Turkestan, Morocco, Israel, Jordan. For Spanish 
populations see Dantart (1.c.). 


GENITALIA 9: see Fig. 132 (N. Israel). No remarkable and constant 
differences between populations examined from S. Italy, S. Greece, 
Cyprus, Tunisia, Israel, Jordan. For Spanish populations see Dantart 
(L.c.). 

REMARKS : “C. p. palaestinensis Fuchs, 1903” is often regarded as valid 
at subspecific rank for the populations of the Middle East. Neither 


SI 


genitalic morphology nor external appearance reveal any indication 
of isolation of populations and separation into distinct subspecies. 


“Chlorissa” solidaria (Guenée, 1857) 


REMARKS: Recorded for S. Iraq in Wiltshire (1957 : 101) as “Chlorissa 
solidaria Guen.”. This is a reference to Prout (1921 : 187), who mentions 
“solidaria Guen.” from Fao, S. Iraq as a species “closely allied to Chlo- 
rissa pulmentaria”: This, however, is obviously concerning Symmacra 
solidaria Gn., one of the few green species belonging to Sterrhinae : 
it is transferred to the genus Symmacra by Prout (1935 : 26). 


Phaiogramma faustinata (Milliére, 1868), comb. n. 


EXTERNAL FEATURES : Venation very similar to C. pulmentaria : Fore- 
wing: RI typically free (in a very few specimens however fused with 
Sc); R2-R5 and MI connate, exceptionally very shortly stalked or 
separate, M3 and CuAl usually connate, but variable. Hindwing : 
discocellular vein arising from M3 + CuAl medially between CuAl 
and CuA2. In C. African populations M3 and CuAl of forewing 
sometimes distinctly stalked, discocellular vein of hindwing arising from 
M3 + CuAl very near CuA2. Length of tongue 3-3.5 mm. Palpus of 
male long, length about 1.5 times diameter of eye ; of female very long : 
2.3-2.8 times diameter of eye. Frons flat. Antenna (@Q) as described 
for P. pulmentaria, flagellum of male antenna slightly dentate. Hindtibia 
(SQ) as described for C. viridata, spurs of & rather short, in © the 
length of the longer terminal and longer proximal spur more than twice 
that of the shorter. Apart from the venation there is no structural 
variation within the area of distribution. 


GENITALIA @: see Figs. 74-76 (N. and S. Israel; Cyprus). No re- 
markable and constant differences observed between populations 
examined from Tunisia, Libya, Egypt, S. Greece, Cyprus and S. Oman. 
For Spanish populations see Dantart (l.c.). However in populations 
examined from Armenia, N. and C. Israel there are three patches of 
cornuti present instead of two, with only a very few exceptions (n = 35 
& slides). The only @ examined from S. Israel (Yotvata ; Fig. 75) has 
apparently two patches of cornuti, but this may be due to positioning 
on the slide. In Egypt there are regularly two patches. 


GENITALIA Q: see Fig. 133 (Israel). No remarkable and constant 
differences observed between the populations examined from Tunisia, 


52 


Egypt, S. Greece, Crete, Cyprus, C. Lebanon, Israel, N. Jordan, N. 
Oman, Sudan. For Spanish populations see Dantart (l.c.). 


Remarks: The N. African “Eucrostes” rhoisaria (Chrétien, 1909) is 
a synonym (compare Hausmann, 1994a). In C. Africa male genitalia 
and venation (see above) seem to justify the separation of some sub- 
species (or species 4). The whole complex should be revised separately. 


Phaiogramma discessa (Walker, 1861), comb. n. 


EXTERNAL FEATURES : unknown. 


GENITALIA 4: see Fig. 77 (Bahrein) : According to Wiltshire (1990 : 
110) differs from the next species in the aedeagus. Fig. 77 is redrawn 
from a sketch sent to the author by Mr. E. P. Wiltshire in a personal 
communication : central patch of external spines (“cornuti”) replaced 
by a sclerotised ridge. However, conspecifity of this Bahrein specimen 
with typical P discessa from India (“N.-Hindustan”) awaits verification. 


GENITALIA 9 : unknown. 


REMARKS: In 1985 Rose & Devinder described “Chlorissa patialensis” 
(Phaiogramma patialensis (Rose & Devinder, 1985), comb. n.) from 
NW. India. The genitalia figured in the original description are quite 
similar to those of P faustinata. As far as can be judged from this 
illustration, the processus of the costa of the valva is lacking. As the 
locus typicus of patialensis and that of discessa are almost the same 
and no differential diagnosis has been made, the two names may be 
synonymous. If this is the case, the & from Bahrein would belong 
to another species or it could be an extremely abnormal mutation of 
P. faustinata. 


Phaiogramma polemia (Prout, 1920), comb. n. 


GENITALIA & : see Wiltshire (1967: pl. VI, Fig. 4). Conspecifity not 
ascertained. 


REMARKS: For data on the morphology see Hausmann (1995 : 590). 
No authentic material available, photograph of holotype (Q ; BMNH) 
examined. 

NEROMIA Staudinger, 1898 


DIFFERENTIAL FEATURES : Venation see Fig. 23: similar to that of 
Chlorissa and Phaiogramma. Forewing: RI free; R2-R5 and MI 


53 


stalked, sometimes distinctly ; M3 and CuAl shortly stalked or connate. 
Hindwing termen rounded as in Phaiogramma. Palpus comparatively 
short and slender in both sexes (in N. simplexa 2 palpus elongate). 
Antenna of & ciliate with cilia grouped in tufts ; antenna of © scarcely 
ciliate. Hindtibia of female with two spurs. Hindtarsus of male long 
in N. pulvereisparsa (in N. simplexa shortened). 4 genitalia somewhat 
resembling that of Hemithea (e.g. postition of harpe in N. pulvereis- 
parsa), but socius broader ; uncus slightly split terminally (but not in 
N. simplexa) ; basal coremata of valva present ; transtilla expanded. 
© genitalia more or less of the Phaiogramma type : lamella antevaginalis 
divided and posterior (caudal) part of corpus bursae wrinkled, somewhat 
resembling female genitalia of the Microloxiini genus Thelycera. 


REMARKS : The female hindtibia of some African “Chlorissa” species 
has two spurs (Prout, 1930 : 27). These species should be re-examined 
and incorporated in a revision of the generic differences between 
Chlorissa, Phaiogramma, Hemithea, Diplodesma and Neromia. Since 
N. simplexa shows transitional features to Phaiogramma, it should be 
considered whether downgrading of Neromia to a subgenus of Phai- 
ogramma may be necessary. Phaiogramma and Neromia are more 
closely related than Phaiogramma and Chlorissa. 


Within the Hemitheini the combination of two spurs on female hind- 
tibia, short female palpus (these features being the main differential 
characters according to Prout 1913: 26), dorsoventrally split uncus 
and harpe arising from ventral part of valva are quite unusual and 
may indicate certain relationships to Microloxiini. However the filiform 
antenna of 49, long socius (@ genitalia), and presence of basal coremata 
of valva are quite unusual for Microloxuni. 


Neromia pulvereisparsa pulvereisparsa (Hampson, 1896) 


EXTERNAL FEATURES: Venation see Fig. 23 and generic description. 
Length of tongue 2-3 mm. Length of palpus in @ somewhat exceeding 
diameter of eye, in Q somewhat longer : 1.2-1.4 times diameter of eye. 
Frons flat, dark brown. Vertex greyish. Antenna of male filiform, 
slightly notched between segments, with two rows of tufts of cilia, longest 
cilia (0.20 mm) about twice width of flagellum ; antenna of female 
filiform, ciliate, longest cilia somewhat exceeding width of flagellum. 
Relative length tibia/tarsus in foreleg (49) about 1.1/2.3 mm; in 
hindleg of both male and female about 1.8-2.0/1.7-1.8 mm. Male and 
female hindtibia with two long spurs of unequal length. 


54 


GENITALIA @: see Fig. 78 (N. Oman). No remarkable differences 
observed between specimens examined from N. Oman and S. Iran. 


GENITALIA 9: see Fig. 134 (N. Oman). No remarkable differences 
observed between specimens examined from N. Oman and S. Iran. 


REMARKS: The descriptions given above concern material from N. 
Oman and S. Iran (no topotypic material examined). The external 
appearance of one S. Oman specimen (nearer to locus typicus) figured 
in Wiltshire (1990: Fig. 61) is identical to the specimens examined 
(see remarks to N. p. jodisata). 


Neromia pulvereisparsa jodisata Staudinger, 1898 


EXTERNAL FEATURES : Little structural difference from the nominate 
subspecies as described above : Tongue slightly shorter, length ca. 1.7- 
2.5 mm. Scales of frons ochreous, paler than in nominate subspecies, 
as is the colouring of the whole insect, even the vertex is paler. 


GENITALIA @ : see Fig. 79 (C. Israel). No remarkable and constant 
structural differences from the nominate subspecies. Specimens exami- 
ned from Israel and Jordan. The male genitalia of an Egyptian specimen 
figured by Wiltshire (1949: Fig. 83) obviously match well those of 
Palestinian populations. 


GENITALIA 9 : see Fig. 135 (C. Israel). No remarkable and constant 
structural differences from the nominate subspecies. Specimens exam- 
ined from Israel and Jordan. 


REMARKS: Differs from the nominate subspecies mainly in the much 
brighter wing colour. As the species is widely distributed in the Arabian 
peninsula, perhaps there is no real isolation between the typical popu- 
lations of jodisata and pulvereisparsa, in which case the name jodisata 
would sink in synonymy. 


Neromia simplexa Brandt, 1938 


EXTERNAL FEATURES: Venation (compare Fig. 23) as described in 
generic diagnosis. Forewing : R2-R5 and M1 on short common stalk ; 
M3 and CuAl connate (constant in three specimens examined). Length 
of tongue 2.5-3.0 mm. Palpus brown, length in & 0.80 mm (1.2 times 
diameter of eye), in ® 0.90-0.95 mm (1.7-1.8 times diameter of eye). 
Frons flat or slightly convex, brown. Vertex whitish. Antenna of 4 
filiform, ciliate to tip, longest cilia 0.25-0.30 mm (about three times 
width of flagellum) ; antenna of © scarcely ciliate, length of cilia about 


2) 


half width of flagellum. Relative length tibia/tarsus in foreleg (49) 
0.9/1.8 mm. Tarsus shortened in hindleg (49) : relative length tibia/ 
tarsus 1.7-1.8/1.2-1.3 ; hindtibia of both sexes with two spurs of unequal 
length. 3 


GENITALIA @ : see Fig. 80 (N. Oman). Socius subapically very broad. 
Central part of valva strongly sclerotised with two stout projections. 
Aedeagus with two patches of very small cornuti. The genitalia figured 
here correspond to sketches of specimens from C. Iraq (Baghdad), 
which Mr. E. P. Wiltshire sent to the author in a personal communi- 
cation. 


GENITALIA Q : see Fig. 136 (N. Oman). 


REMARKS: N. simplexa shows transitional features between Neromia 
and Phaiogramma (especially P faustinata) in length of © palpus, length 
of hindtarsus, shape of harpe (4 genitalia), breadth of socius, position 
of cornuti in aedeagus (3 genitalia) and lamella antevaginalis (© 
genitalia). Perhaps it would be better transferred there. 


KUCHLERIA Hausmann, 1995 


DIFFERENTIAL FEATURES : For generic differential analysis see Haus- 
mann (1995). Venation see Fig. 24. Sternite 2 without lateral apophyses 
(differential feature from most other Hemitheini genera). 


Kuchleria gisisi Hausmann, 1995 


GENITALIA @ see Fig. 81 (C. Israel), of Q see Fig. 137 (C. Israel). 


REMARKS: For data on the morphology and distinguishing characters 
from the other species of the genus see Hausmann (1995 : 589). 


Tribus MICROLOXIINI tribus n. 


TYPE GENUS: Microloxia Warren, 1893, by original (= present) de- 
signation. 


DIFFERENTIAL FEATURES : Venation : see Figs. 25-29 and Hausmann 
(1995) : Hindwing : Termen rounded ; Rs and MI usually very distinctly 
stalked ; very long fusion of Sc + Rl and Rs in some genera. Frenulum 
sometimes present in male (tending to be weakly developed), but often 
absent ; absent in female. Palpus short and slender in both sexes. Frons 
flat. Tongue short or absent, rarely well developed. Antenna in both 
sexes bipectinate (except Thelycera). Hindtibia of female with two spurs 


56 


only. Abdominal crests absent. & genitalia: Distal end of uncus often 
split dorsoventrally ; socius present, but often broad and shortened ; 
gnathos typically weak or absent ; valva without basal coremata (except 
Thelycera) ; harpe arising from ventral part of valva. 


GENERA INCLUDED here after examination of their type species : Micro- 
loxia Warren, 1893, Mixocera Warren, 1901, Thelycera Prout, 1912, 
Acidaliastis Hampson, 1896, Acidromodes gen. n., Tropicollesis Prout, 
1930, Hemidromodes Prout, 1912, Allochrostes Prout, 1912, Para- 
prasina Warren, 1897, and perhaps also Syndromodes Warren, 1897 
(despite complete number of spurs and filiform antenna ; see Comosto- 
lini). Hierochthonia Prout, 1912 placed here only provisionally. 


REMARKS: For possible relationships with Neromia see above. The 
presence of basal coremata on the valva in Thelycera is perhaps to 
be interpreted as a plesiomorphic feature. Reduction (as e.g. in Thely- 
cera) and elongation of branches of antenna are presumably processes 
taking place with (comparatively) high frequency in the phylogenesis 
of the Geometrinae. The frenulum must surely have been lost many 
times independently not only in the Microloxiini, but also in other 
tribes of Geometrinae. 


MICROLOXIA Warren, 1893 


DIFFERENTIAL FEATURES: For generic description see Hausmann, 
1995. Venation see Fig. 25. 


Microloxia herbaria (Hübner, [1813]) 


GENITALIA @ see Fig. 83 (C. Turkey), © see Fig. 138 (SE. Turkey). 


REMARKS : For description of morphology see Hausmann (1995 : 573). 
Microloxia ruficornis Warren, 1897 


GENITALIA @ see Fig. 84 (S. Israel), © see Fig. 139 (S. Israel). 


REMARKS : For description of morphology see Hausmann (1995 : 576). 
ACIDALIASTIS Hampson, 1896 
DIFFERENTIAL FEATURES : Venation see Fig. 26: Forewing: RI fused 


with Sc, not reaching costa as in Microloxia ; R2-R5 distinctly stalked ; 
R3 lacking, exceptionally present in wing apex; origins of R2, R4 


sf 


and RS on the stalk very near to each other, sometimes connate ; R2, 
R4 and R5 widely diverging ; R2-RS and MI unstalked ; M3 and CuAl 
unstalked, points of origin from cell often well separated. Hindwing : 
Sc + RI and Rs fused over 3/4 length of cell, sometimes even 4/5, 
in Israel and Jordan up to 7/8 ; Rs and M1 on extremely long common 
stalk, distally widely diverging ; M3 and CuAl unstalked, sometimes 
separate, sometimes on very short common stalk. Frenulum lacking 
in both sexes. Frons flat. Palpus short in both sexes. Tongue lacking. 
Antenna bipectinate in both sexes. Hindtibia with two spurs in both 
sexes, not dilated in male. À genitalia : Uncus broad ; gnathos lacking ; 
socius shortened ; valva ventrally with long, stout, outwardly directed 
and curved spine (harpe) ; aedeagus without cornuti ; sternite 8 simple, 
its anterior margin strongly convex. © genitalia very simple with small 
and weakly sclerotised corpus bursae. Apophyses very short, broad, 
triangular. 


REMARKS : Studies of the differential features of the genus Acidaliastis 
necessitate the description of a new genus (see below). Acidaliastis 
curvilinea (Prout, 1912) according to Janse (1935 : 403) has two spurs 
on the male hindtibia, not four as stated in Prout (1930 : 48). Genitalia 
(e.g. shape of valva) quite similar to A. micra. The generic placement 
of Acidaliastis prophanes Prout, 1922, is confirmed by genitalic 
morphology. 


Acidaliastis micra Hampson, 1896 
Euchloris dissimilis Warren & Rothschild, 1905 : Novitates Zool. 12 : 26, pl. 4, 
Fig. 27, syn. n. 


EXTERNAL FEATURES : Venation see Fig. 26 and generic description. 
Palpus white, length in both sexes ca. 0.45 mm (= diameter of eye). 
Frons white or brigthly ochreous. Antenna bipectinate nearly to tip 
in both sexes, longest branches in male 0.30-0.38 mm (6-8 times width 
of flagellum), in female about 0.20 mm (three times width of flagellum), 
branches broad, arising quite distantly from each other. Relative length 
hindtibia/tarsus in @ about 1.0/0.8 mm, in Q about 1.4/1.1 mm. 


GENITALIA @: see Fig. 85 (S. Israel). See also generic description. 
Infrapopulational variability rather small. No structural difference with- 
in the area of distribution, neither between the whitish and the brownish 
forms. 


GENITALIA 9: see Fig. 140 (S. Israel). See also generic description. 
No structural difference within the area of distribution, not even be- 
tween the whitish and the brownish forms. 


58 


REMARKS: Locus typicus of A. micra is Aden. According to Prout 
(1930 : 48) characterised by small size, straight postmedian line and 
black cell spots. From Sudan, the locus typicus of “Euchloris” dissimilis 
Warren & Rothschild, 1905, and from Israel large numbers of specimens 
(several hundreds from each country) were examined by the author : 
Brownish coloured specimens occur only occasionally, cell spot often 
rather small, specimens often very small (compare distinguishing cha- 
racters mentioned by Prout, l.c.). As there are no constant differences 
in structure or habitus within the area mentioned above, the taxon 
dissimilis has to sink into synonymy. 


ACIDROMODES gen. n. 


TYPE species: Acidaliastis nilotica Wiltshire, 1985, by original 
(= present) designation. 


DIFFERENTIAL FEATURES : Venation see Fig. 27: Forewing: RI fused 
with Sc for a short distance only, not reaching costa, as in Microloxia 
and Acidaliastis ; R2-R5 and MI shortly stalked, all veins present ; 
R2-R5 not widely diverging as in Acidaliastis ; M3 and CuAl separate. 
Hindwing : Sc + R1 and Rs fused over somewhat more than 1/2 length 
of cell only (in Acidromodes saharae 3/4); Rs and MI distinctly 
stalked, but shorter than in Acidaliastis (in Acidromodes saharae 
distally diverging as in Acidaliastis) ; M3 and CuAl on a short common 
stalk, in Acidromodes saharae distinctly stalked. Frenulum absent in 
both sexes (present in & of Tropicollesis). Tongue absent. Male palpus 
quite long, length about 0.75 mm (1.2 times diameter of eye). Antenna 
bipectinate to tip, longest branches about 0.65 mm (5-6 times width 
of flagellum). Hindtibia of male strongly dilated, with four spurs. Hind- 
tarsus short, relative length tibia/tarsus about 2.0/1.1 mm. @ genitalia 
(see Fig. 82): Uncus simple, narrow; socius broad, nearly reaching 
length of uncus ; gnathos present, but weak ; harpe arising from ventral 
part of valva, somewhat resembling that of Hemidromodes robusta 
(see below) ; aedeagus with one short stout cornutus ; the long stout 
processi of sternite 8 are unique. © genitalia unknown. 


EXAMINED MATERIAL : Holo- and Paratype of Acidromodes nilotica 
(Wiltshire, 1985) from Sudan, also one male of Acidromodes saharae 
(Wiltshire, 1985) from Burkina Faso (Upper Volta). 


REMARKS: Not represented in the study area. According to Prout 
(1930 : 48) the male hindtibia of the African species “Acidaliastis” 
subbrunnescens Prout, 1916, and “Acidaliastis” bicurvifera Prout, 1916, 
have four spurs ; therefore they probably should be included in this 


59 


new genus. This genus has a combination of various particular features 
evidently showing close relationships to Hemidromodes and Tropicol- 
lesis : male hindtibia dilated, with four spurs ; fusion of Sc + Ri and 
Rs ; stout cornutus ; shape of harpe etc. 


HEMIDROMODES Prout, 1916 


DIFFERENTIAL FEATURES : Venation see Fig. 28: Forewing: Sc and 
R1 free, without fusions ; R2-R5 and M1 shortly stalked, exceptionally 
connate ; M3 and CuAl connate, sometimes very shortly stalked or 
separate. Hindwing : Sc + RI and Rs fused for a long distance ; Rs 
and MI distinctly stalked ; stalk of M3 and CuAl somewhat shorter, 
sometimes connate. Venation of hindwing according to Prout (1935: 
16) similar to that of Hierochthonia and that of Syndromodes (Prout, 
1930). Frenulum present in male, but often somewhat weak, absent 
in female. Frons flat. Palpus whitish, in @ very short and slender, in 
Q slender, but somewhat elongate. Tongue lacking. Antenna short 
(ca. 2-3 mm), bipectinate in both sexes, branches of & long, of © short. 
Hindtibia of 4 greatly dilated, with four spurs, proximal pair of spurs 
long, the terminal ones very short. Q hindtibia very slender, only 
terminal pair of spurs present. 5 genitalia : uncus simple, stout ; socius 
broad, somewhat shorter than uncus ; gnathos more strongly sclerotised 
than in other Microloxiini genera ; juxta and transtilla expressed as 
two extended sclerites ; the stout basal processus of valva is unique ; 
harpe arising from ventral part of valva; aedeagus with strongly 
sclerotised cornutus (cornuti) ; sternite and tergite 8 simple. © genitalia : 
Lamella antevaginalis strongly sclerotised ; corpus bursae with a 
constriction separating a stiff, longitudinally ribbed, posterior part from 
a smaller smooth anterior part. 


REMARKS: Its nearest ally in Africa is perhaps the genus Tropicollesis. 
There may also be some relationships to “Lophochoristini” of North 
America. One further new species from C. Africa will be described 
in a separate article. 


Hemidromodes robusta robusta (Prout, 1913) 


EXTERNAL FEATURES: Venation see Fig. 28 (Sudan): Variability as 
described in generic diagnosis. Length of palpus in & 0.30-0.45 mm 
(0.5-0.7 times diameter of eye), in 2 0.70-0.80 mm (about 1.2 times 
diameter of eye). Frons pale brown or ochreous, becoming brighter 
towards ventral part. Antenna (4Q) bipectinate nearly to tip ; length 


60 


of longest branches in ¢ 0.40-0.50 mm (about 6 times width of flagel- 
lum), in 2 about 0.20 mm. Terminal pair of spurs in & hindtibia extreme- 
ly short. Relative length tibia/tarsus 1.7/1.2 mm ; © exceptionally with 
very short rudiments of proximal spurs. 


GENITALIA 6 : see Fig. 86 (NE. Sudan). Little infrapopulational 
variability within the long series from Sudan at the ZSM. Length of 
cornutus somewhat variable. In two slides (BMNH ; @ from Assuan, 
S. Egypt) harpe more irregularly shaped. 


GENITALIA Q : see Fig. 141 (NE. Sudan). Corpus bursae strongly 
tapering anteriorly. Ostium bursae with deep notch posteriorly. 


REMARKS: The species name used in many publications is “affinis” 
while “robusta” has often (erroneously) been used as the species name 
for the eastern eremic species H. sabulifera. The populations of S. and 
C. Egypt probably belong to the nominate subspecies, but further 
material is necessary to resolve this question. 


Hemidromodes robusta affinis (Rothschild, 1915) 


EXTERNAL FEATURES : No material available. 
GENITALIA @ : unknown. 
GENITALIA © : unknown. 


REMARKS: Subspecies rank is contested by Herbulot (1965 : 27), who 
sinks it into the synonymy of H. robusta. Since the author also includes 
triforma as a subspecies of H. robusta, the validity of the differential 
features used is doubtful. Synonymy however is not excluded. According 
to Speidel & Hassler (1989 :51) two species occur in the Sahara. The 
record of H. robusta “sensu Wiltshire” (= H. sabulifera triforma) how- 
ever has to be regarded as erroneous. The genitalia of “A. subbrun- 
nescens Prout, 1915” (Speidel & Hassler, l.c.) are figured under the 
name “Acidaliastis subbrunnescens Prout, 1916” in Wiltshire (1985, 
pl. 19, Fig. 4). Wiltshire’s figure is strikingly similar to the genitalia 
of Acidaliastis micra (see above). 


Hemidromodes robusta galala Wiltshire, 1949 


EXTERNAL FEATURES : No material available. 


GENITALIA @ : see Wiltshire (1949 : Fig. 84) ; in the illustrated genitalia 
slide (WA. 102) aedeagus with cornutus accidentally lost. Apparently 
not different from Sudanese populations. According to Wiltshire (1949 : 


61 


402) green “affinis” from Assuan (S. Egypt) “with identical genitalia”; 
those genitalia examined by the author (see above under A. r. robusta). 


GENITALIA © : unknown. 


REMARKS : On account of the apparently constant red wing colour 
this is maintained provisionally as a subspecific taxon. One male from 
Marigat, Kenya is somewhat different. 


Hemidromodes unicolorata sp. n. 


HOLOTYPE : 4, Kenya, Marigat, 21.VIII.1973, leg. Politzar, coll. Som- 
merer. Genitalic morphology of one specimen from “Abessinia” (Prp. 
Geom. 8125) in the BMNH corresponds well. 


DESCRIPTION : Deep green like H. robusta, crosslines on upperside of 
wings almost invisible. Length of forewing 7.7 mm. External structure 
without differences from A. robusta. 4 genitalia (Fig. 87): basal 
processus of valva much longer than in H. robusta; harpe tapering 
distally ; cornutus of aedeagus broader ; posterior edge of sternite 8 
strongly convex. More detailed description and figures of the adult 
will be given in a subsequent paper. 


Hemidromodes sabulifera sabulifera Prout, 1922 


EXTERNAL FEATURES : For venation see generic description and compare 
Fig. 28 : Hindwing : Sc + RI and Rs fused over 1/3-1/2 of length of 
cell; M3 and CuAl shortly stalked. Palpus in & short (0.5 mm = 
0.7 times diameter of eye), in Q comparatively long (0.9-1.0 mm = 
1.2-1.4 times diameter of eye). Frons pale brown, brighter towards 
ventral part. Antenna as described for H. robusta, longest branches 
of 4 antenna somewhat longer (0.5-0.6 mm). Terminal pair of spurs 
in & hindtibia short, but usually distinctly visible, occasionally absent. 
© hindtibia without rudiments of proximal spurs. Relative length tibia/ 
tarsus in hindleg about 1.6/1.0 mm. 


GENITALIA @ : see Fig. 88 (E. Iran). Basal processus of valva longer than 
in H. robusta, Juxta more rounded, Harpe more strongly sclerotised, 
strongly tapering dorsally, Cornutus longer. As yet no specimen found 
with second cornutus present (compare H. s. hessa). No remarkable 


and constant difference between specimens examined from E. Iran and 
N. Oman. 


62 


GENITALIA 9 : see Fig. 142 (E. Iran). Ostium bursae posteriorly only 
a little concave in E. Iranian specimen, however rather deeply notched 
in one 9 from N. Oman. 


REMARKS: Holotype (Q) re-examined in the BMNH (Prp. Geom. 
4062). It is strikingly different from the following subspecies in the 
white ground colour and the comparatively straight (and dotted) 
postmedian lines on all wings. N. Oman specimens not distinguishable 
in external appearance from material from E. Iran. According to 
Wiltshire (1990 : 111) the distribution of nominate subspecies includes 
NW. India, S. Iran, C. and E. Saudi Arabia, but in C. Arabia there 
are transitions to A. s. hessa ; real isolation between the populations 
is doubtful ; eventually the taxa may have to be synonymised. 


Hemidromodes sabulifera hessa Prout, 1935 


EXTERNAL FEATURES : For venation see generic description and compare 
Fig. 28: Hindwing: fusion of Sc + RI and Rs over about 1/3-1/4 
of length of cell; Rs and MI on a very long stalk ; M3 and CuAl 
rather distinctly stalked ; these features expressed with high constancy. 
External features as described for A. sabulifera, but palpus of © slightly 
shorter than equivalent in nominate subspecies (1.1-1.3 times diameter 
of eye). Relative length tibia/tarsus about 1.9/1.2 mm. Terminal spurs 
in @ hindtibia always present and distinctly visible. 


GENITALIA @ : see Figs. 89-91 (C. and S. Israel). High infrapopulational 
variability in the shape of harpe and cornutus. Even some variation 
in the number of cornuti : out of 75 dissected @@ of H. s. hessa (from 
Israel) 11 @4 had two cornuti present (this feature commoner in S. 
than in C. Israel). Little difference between the genitalia of specimens 
examined from Israel/Jordan and genitalia of the nominate subspecies : 
Harpe often with one or two small additional lateral teeth, mainly 
in specimens from the northern part of the area of distribution. 


GENITALIA © : see Fig. 143 (S. Israel) : Resemble those of A. s. sabu- 
lifera, but as in the N. Oman specimen the ostium bursae has a deep 
notch posteriorly. 


REMARKS: Dissected Paratype (4) of H. s. hessa re-examined in the 
BMNH (Prp. Geom. 8117). Differs from the nominate subspecies in 
the brownish, not dotted ante- and postmedian lines, which are strongly 
undulate ; wings often sandy coloured. 


63 


Hemidromodes sabulifera triforma Wiltshire, 1949 


EXTERNAL FEATURES : As described for A. s. hessa. 


GENITALIA @: see Fig. 92 (W. Saudi Arabia). Specimens examined 
from Saudi Arabia and Yemen. No difference from & genitalia of HA. s. 
hessa. 


GENITALIA © : unknown. 


REMARKS: Holotype of “H. robusta triforma” re-examined in the 
BMNH (Prp. Wilts. 827). Populations of H. s. hessa and A. s. triforma 
are not clearly separated from each other : there are no differences 
in genitalic morphology, but in external appearance there are transitional 
features in S. Israel and S. Jordan. The latter name may have to be 
considered a synonym of A. s. hessa. 


HIEROCHTHONIA Prout, 1912 


DIFFERENTIAL FEATURES: Venation see Fig. 29: Forewing: Sc and 
RI anastomosing ; length of fusion variable, often touching at one 
point only ; length of fusion between RI and R2 variable too; R3 
lost (total fusion with R4); R2-R5 and M1 distinctly stalked ; M3 
and CuAl connate or exceptionally stalked. Hindwing : long fusion 
of Sc + RI and Rs (2/3 to 3/4 of length of cell); Rs and MI on 
very long common stalk ; M3 and CuAl distinctly stalked, length of 
stalk variable, in one specimen M3 and CuAl connate. Frenulum 
present in @, absent in ©. Frons flat, reddish brown. Vertex white. 
Tongue lacking. Palpus very small and slender. Antenna of both sexes 
bipectinate, branches long in @, short in Q. Hindtibia of both sexes 
not dilated, with two terminal spurs. 4 genitalia : uncus short, slightly 
split dorsoventrally ; socius present, but often hardly visible, being inside 
fold over uncus ; gnathos rather strongly sclerotised ; basal coremata 
absent ; aedeagus with one stout cornutus. ® genitalia: lamella ante- 
and postvaginalis characteristically sclerotised ; posterior (caudal) half 
of corpus bursae stiff, longitudinally ribbed, divided by a constriction 
from membranous anterior half which has small, horizontally positioned 
signum bursae. | 


REMARKS: The systematic position is unclear. Venation, structure of 
harpe in @ genitalia and other features probably indicate relationships 
to Microloxiini. Transverse subdivision of corpus bursae in © genitalia 
as in Hemidromodes. Some other features, but especially the habitus 
reminiscent of Xenochlorodes. Possibly forms a link between Micro- 


64 


loxuni and Xenochlorodes (actually placed in Hemistolini). The African 
genus Rhodesia Warren, 1905, corresponds in many features (e.g. 49 
antenna, frenulum, $Q genitalia) ; however the structure of palpus and 
hindtibia is rather different. 


Hierochthonia pulverata (Warren, 1901) 


EXTERNAL FEATURES : For venation see generic description (Fig. 29). 
Palpus ochreous with some reddish-brown scales, length 0.45-0.50 mm 
(3/4 diameter of eye) in both sexes. Length of longest antennal branches 
in @ about 0.55 mm (4-5 times width of flagellum), in ® about 0.25 mm 
(twice width of flagellum). Relative length tibia/tarsus in the hindleg 
of both sexes about 1.8/2.0 mm. Further features see generic description. 


GENITALIA @ : see Fig. 93 (C. Lebanon). Valva basally much broader 
han in the following species. Harpe strongly sclerotised, somewhat 
tapering, placed in central part of valva. Cornutus crest-shaped. 
Posterior edge of sternite 8 convex. Only little structural variation 
within the area of distribution (specimens examined from N. and C. 
Lebanon). 


GENITALIA Q : see Fig. 144 (C. Lebanon). Resembling very much those 
of following species. Lamella postvaginalis almost rectangular. Length 
of signum bursae somewhat variable. There is little structural variation 
within the area of distribution (specimens examined from S. Turkey, 
N. and C. Lebanon). 


REMARKS : H. pulverata and the following species are probably allo- 
patric vicariants. To date no sympatric occurrence is known. Although 
very closely related to each other, striking differences in @ genitalia 
do not allow them to be considered conspecific. 


Hierochthonia semitaria (Piingeler, 1901) 


EXTERNAL FEATURES : Venation and other external features without 
any remarkable and constant difference from H. pulverata. See de- 
scription above. One Jordanian male has antennal branches slightly 
shorter (about 0.45 mm). 


GENITALIA À : see Fig. 94 (C. Israel). Valva more slender than in the 
preceding species. Harpe less tapering, with one small subapical tooth ; 
harpe placed on ventral edge of valva. Cornutus crest-shaped with 
central teeth much elongate. Posterior edge of sternite 8 concave. There 


65 


is little structural variation within the area of distribution (specimens 
examined from N. and C. Israel, C. Jordan). 


GENITALIA © : see Fig. 145 (C. Israel). Very similar to genitalia of 
preceding species. Lamella postvaginalis posteriorly tapering, almost 
triangular. Length of signum bursae somewhat variable. Only little 
structural variation within the area of distribution (specimens examined 
from N. and C. Israel, N. and C. Jordan). 


REMARKS : Compare preceding species. Both original types examined 
(4Q) ; male labelled as “type” (= holotype), female as “cotype” (= para- 


type). 
Acknowledgements 


I wish to express my gratitude to the following colleagues, who helped by 
loaning or donating material: E. Arenberger, Vienna, O. Karsholt, Copen- 
hagen, K. Kuchler, Munich, M. Krüger, Pretoria, M. Lödl, Vienna, H. 
Malicky, Lunz (Austria), W. Mey, Berlin, B. Müller, Berlin, M. NuB, Berlin, 
P. Schaider, Munich, M. Scoble, London, B. Skule, Rodovre (Denmark), 
D. Stiining, Bonn, L. Weigert, Deggendorf (Germany), J. Wimmer, Steyr 
(Austria). For friendly scientific correspondence I have to thank E. P. Wilt- 
shire, Berks (U.K.) and C. Herbulot, Paris. For scientific cooperation 
concerning the fauna of Israel (e.g. organisation of field collecting, scientific 
discussions) I am deeply indebted to G. Miller, Jerusalem, R. Ortal, Jerusalem, 
and A. Freidberg, Tel Aviv. M. Sommerer, Munich, A. Olivier, Antwerpen, 
and S. Whitebread, Magden (Switzerland) helped to arrange publication. 
Special thanks also to M. Corley, Oxford, and W. Sauter, Illnau (Switzerland), 
for supporting the editorial process and revising the manuscripts. 


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Lissochlora and Chavarriella, with particular reference to the species 
of Costa Rica (Lepidoptera : Geometridae, Geometrinae). Bull. nat. Hist. 
Mus. Lond. (Ent.) 62 (2) : 39-159. 

Prout, L. B., 1913. Die spannerartigen Nachtfalter. /n : Seitz, A. [1912-1916] : 
Die Gross-Schmetterlinge der Erde, Bd. 4. Verlag A. Kernen, Stuttgart. 

Prout, L. B., 1921. Moths of Mesopotamia and N.W. Persia (Pt. II). J. 
Bombay nat. Hist. Soc. 28 : 187-191. 


67 


Prout, L. B., 1930. Die spannerartigen Nachtfalter (Fauna Africana). In: 
Seitz, A., Groß-Schmetterlinge der Erde, Bd. 16. Verlag A. Kernen, 
Stuttgart. 

Prout, L. B., 1934. Die indoaustralischen Spanner. Jn : Seitz, A. [1920-1941], 
GroB-Schmetterlinge der Erde, Bd. 12. Verlag A. Kernen, Stuttgart. 
Prout, L. B., 1935. Brephinae, Oenochrominae, Hemitheinae, Sterrhinae, 
Larentiinae. In: Seitz, A. [1934-1939]: Die Gross-Schmetterlinge der 

Erde, Suppl. 4. Verlag A. Kernen, Stuttgart. 

ROSE, H. S. & DEVINDER, 1985. A new species of the genus Chlorissa Stephens 
from North India (Geometrinae : Geometridae : Lepidoptera). J. Bombay 
nat. Hist. Soc. 82 : 614-616. 

SCOBLE, M. J., 1994. A taxonomic revision of the genera Phrygionis Hübner 
and Pityeja Walker (Geometridae : Ennominae, Palyadini). Zool. J. 
Linn. Soc. 111 : 99-160. 

SPEIDEL, W. & HassLer, M., 1989. Die Schmetterlingsfauna der südlichen 
algerischen Sahara und ihrer Hochgebirge Hoggar und Tassili n’Ajjer 
(Lepidoptera). Nachr. ent. Ver. Apollo Suppl. 8 : 1-156. 

VuDALEPP, J., 1976. A list of Geometridae (Lepidoptera) of the USSR, I. 
Ent. Obozr. 55 (4) : 842-852. 

VIIDALEPP, J., 1981. The systematic of the genera of the subfamily Geometrinae 
(Lepidoptera, Geometridae). Trudy vses ent. Obshch. 63 : 90-95. 

VIDALEPP, J., 1988. Geometridae fauna of the Central Asian mountains. 
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Vives Moreno, A., 1994. Catalogo sistematico y sinonimico de los Lepi- 
dopteros de la Peninsula Iberica y Baleares II, Madrid. 

Vosnits, A. M., 1976. Geometrinae and Sterrhinae from Mongolia (Lepi- 
doptera, Geometridae). Annis hist.-nat. Mus. natn. hung. 68 : 169-174. 

WILTSHIRE, E. P., 1948. Middle East Lepidoptera, IX : Two new forms or 
species and thirty-five new records from Cyprus. Entomologist’s Rec. 
J. Var. 60 : 79-87. 

WILTSHIRE, E. P., 1949. The Lepidoptera of the Kingdom of Egypt, Pt. 2. 
Bull. Soc. Fouad I Ent. 33 : 381-457. 

WILTSHIRE, E. P., 1985. New Geometridae from Africa, with a list of additions 
to the Lepidoptera fauna of the Sudan (Lepidoptera, Geometridae). 
Entomofauna 6 (2) : 9-22. 

WILTSHIRE, E. P., 1990. An illustrated, annotated catalogue of the macro- 
heterocera of Saudi Arabia. Fauna of Saudi Arabia 11 : 91-250. 

WILTSHIRE, E. P., 1994. Arabian Lepidoptera : a Supplement to the Catalo- 
gue of Saudi Arabian Macro-Heterocera. Fauna of Saudi Arabia 14: 
113-136. 


68 


Figs 1-4. Venation (4) 

1) Myinodes shohami Hausmann, 1994 
2) Eumegethes tenuis Staudinger, 1898 
3) Orthostixis cribraria (Hübner, [1799]) 
4) Pingasa lahayei (Oberthür, 1887) 


69 


Figs 5-9. Venation (@) 

5) Pseudoterpna coronillaria halperini ssp. n. 

6) Holoterpna pruinosata (Staudinger, 1898) 

7) Aplasta ononaria (Fuessly, 1783) 

8) Microbaena pulchra minor ssp. n. 

9) Comibaena bajularia ({Denis & Schiffermüller], 1775) 


70 


Figs 10-13. Venation (4) 

10) Proteuchloris neriaria (Herrich-Schäffer, 1852) 

11) Thetidia (Antonechloris) smaragdaria (Fabricius, 1787) 
12) Thetidia (Antonechloris) persica sp. n. 

13) Thetidia (Aglossochloris) euryrithra Prout, 1935 


71 


17 


A 
X 
dA 


Figs 14-17. Venation (@) 

14) Hemistola chrysoprasaria (Esper, 1795) 

15) Xenochlorodes olympiaria cremonaria Staudinger, 1897 
16) Gnophosema palumba Brandt, 1938 

17) Victoria sematoperas Prout, 1916 


72 


Figs 18-23. Venation (@) 

18) Eucrostes indigenata (de Villers, 1789) 

19) Thalera fimbrialis magnata A. Fuchs, 1903 
20) Culpinia prouti (Thierry-Mieg, 1913) 

21) Chlorissa viridata (Linnaeus, 1758) 

22) Phaiogramma pulmentaria (Guenée, 1857) 
23) Neromia pulvereisparsa (Hampson, 1896) 


Figs 24-29. Venation (@) 

24) Kuchleria gisisi Hausmann, 1995 

25) Microloxia herbaria (Hiibner, [1813]) 
26) Acidaliastis micra Hampson, 1896 

27) Acidromodes nilotica (Wiltshire, 1985) 
28) Hemidromodes robusta (Prout, 1913) 
29) Hierochthonia semitaria (Püngeler, 1901) 


74 


Figs 30-35. & Genitalia (scale bar = 1 mm) 

30) Myinodes shohami Hausmann, 1994 (N. Israel ; Paratype) 

31) Eumegethes tenuis Staudinger, 1898 (C. Algeria) 

32) Orthostixis cribraria cribraria (Hiibner, [1799]) (Hungary) 

33) Orthostixis cribraria amanensis Wehrli, 1932 ( S. Turkey ; Paralectotype) 
34) Orthostixis calcularia Lederer, 1853 (C. Turkey) 

35) Orthostixis cinerea Rebel, 1916 (Cyprus ; Holotype) 


75 


Figs 36-41. & Genitalia (scale bar = 1 mm) 

36) Pingasa lahayei (Oberthür, 1887) (Saudi Arabia) 

37) Pseudoterpna coronillaria axillaria Guenée, 1857 (C. Lebanon) 
38) Pseudoterpna coronillaria halperini ssp. n. (NE. Israel ; Paratype) 
39) Pseudoterpna coronillaria cinearascens (Zeller, 1847) (C. Italy) 
40) Holoterpna pruinosata (Staudinger, 1898) (C. Israel) 

41) Holoterpna pruinosata (Staudinger, 1898) (N. Israel) 


76 


Figs 42-49. 4 Genitalia (scale bar = 1 mm) 
42) Aplasta ononaria (Fuessly, 1783) (N. Israel ; f. faecataria Hiibner) 
43) Aplasta ononaria (Fuessly, 1783), with sternite 8 (N. Israel ; f. berytaria Staudinger) 
44) Aplasta ononaria (Fuessly, 1783), genital aberration 
(N. Israel ; f. berytaria Staudinger) 
45) Microbaena pulchra (Staudinger, 1897) (C. Israel ; Holotype) 
46) Microbaena pulchra minor ssp. n., with sternite 8 (above) and tergite 8 (below) 
(NE. Sudan ; Holotype) 
47) Comibaena bajularia ([ Denis & Schiffermüller], 1775), with sternite 8 (C. Turkey) 
48) Comibaena bajularia ([Denis & Schiffermiiller], 1775), sternite 8 only (N. Italy) 
49) Proteuchloris neriaria (Herrich-Schäffer, 1852) (N. Israel) 


Figs 50-55. à Genitalia with sternite 8 (scale bar = 1 mm) 

50) Thetidia (Antonechloris) smaragdaria (Fabricius, 1787) (C. Italy) 

51) Thetidia (Antonechloris) volgaria (Guenée, 1857) (S. Russia) 

52) Thetidia (Antonechloris) persica sp. n. (N. Iran ; Paratype) 

53) Thetidia (Antonechloris) persica sp. n. (S. Turkey) 

54) Thetidia (Antonechloris) silvia Hausmann, 1991 (C. Jordan ; Paratype) 
55) Thetidia (Aglossochloris) euryrithra Prout, 1935 (N. Jordan) 


78 


Figs 56-61. & Genitalia with sternite 8 (scale bar = 1 mm) 

56) Hemistola chrysoprasaria (Esper, 1795) (S. Turkey) 

57) Hemistola siciliana Prout, 1935 (Sicily) 

58) Xenochlorodes olympiaria cremonaria (Staudinger, 1897) (N. Israel) 
59) Gnophosema palumba Brandt, 1938 (S. Iran) 

60) Victoria wiltshirei sp. n. (NE. Sudan ; Holotype) 

61) Victoria sematoperas Prout, 1916 (Kenya) 


Figs 62-67. & Genitalia with sternite 8 (scale bar = 1 mm) 
62) Victoria plantei Herbulot, 1976 (C. Israel) 

63) Victoria eremita Hausmann, 1993 (S. Israel ; Paratype) 
64) Victoria fifensis Wiltshire, 1994 (W. Saudi Arabia) 

65) Eucrostes indigenata (de Villers, 1789) (N. Jordan) 

66) Thalera fimbrialis magnata A. Fuchs, 1903 (C. Turkey) 
67) Culpinia prouti (Thierry-Mieg, 1913) (S. Turkey) 


80 


Figs 68-77. 4 Genitalia with sternite 8 (scale bar = 1 mm) 

68) Chlorissa viridata (Linnaeus, 1758) (N. Italy) 

69) Chlorissa viridata (Linnaeus, 1758), right valva only (Transcaucasia) 
70) Chlorissa cloraria (Hübner, [1813]) (C. Greece) 

71) Chlorissa cloraria (Hübner, [1813]) (C. Turkey) 

72) Chlorissa asphaleia Wiltshire, 1966 (N. Turkey) 

73) Phaiogramma pulmentaria (Guenée, 1857) (N. Israel) 

74) Phaiogramma faustinata (Milliére, 1868) (N. Israel) 

75) Phaiogramma faustinata (Milliere, 1868), aedeagus only (S. Israel) 
76) Phaiogramma faustinata (Milliere, 1868), aedeagus only (Cyprus) 
77) Phaiogramma discessa & (Walker, 1861), aedeagus only (Bahrein) 


81 


Figs 78-85. & Genitalia with sternite 8 (scale bar = 1 mm) 

78) Neromia pulvereisparsa pulvereisparsa (Hampson, 1896) (N. Oman) 
79) Neromia pulvereisparsa jodisata Staudinger, 1898 (C. Israel) 

80) Neromia simplexa Brandt, 1938 (N. Oman) 

81) Kuchleria gisisi Hausmann, 1995 (C. Israel ; Holotype) 

82) Acidromodes nilotica (Wiltshire, 1985) (Sudan ; Paratype) 

83) Microloxia herbaria (Hübner, [1813]) (C. Turkey) 

84) Microloxia ruficornis (Warren, 1897) (S. Israel) 

85) Acidaliastis micra Hampson, 1896 (S. Israel) 


82 


Figs 86-94. 4 Genitalia (scale bar = 1 mm) 

86) Hemidromodes robusta robusta (Prout, 1913) (NE. Sudan) 

87) Hemidromodes unicolorata sp. n. ; valva, aedeagus and sternite 8 only 
(Kenya ; Holotype) 

88) Hemidromodes sabulifera sabulifera Prout, 1922 (E. Iran) 

89) Hemidromodes sabulifera hessa Prout, 1935 (C. Israel) 

90) Hemidromodes sabulifera hessa Prout, 1935 ; aedeagus only (S. Israel) 

91) Hemidromodes sabulifera hessa Prout, 1935 ; aedeagus only (S. Israel) 

92) Hemidromodes sabulifera triforma Wiltshire, 1949 ; valva and aedeagus only 
(W. Saudi Arabia ; Holotype) 

93) Hierochthonia pulverata (Warren, 1901) (C. Lebanon) 

94) Hierochthonia semitaria (Püngeler, 1901) (C. Israel ; Holotype) 


83 


Figs 95-102. © Genitalia (scale bar — 1 mm) 
95) Myinodes shohami Hausmann, 1994 (N. Israel ; Paratype) 
96) Eumegethes tenuis Staudinger, 1898 (C. Algeria) 
97) Orthostixis cribraria (Hübner, [17991 (C. Turkey) 
98) Orthostixis calcularia Lederer, 1853 (C. Turkey) 
99) Orthostixis cinerea Rebel, 1916 (Cyprus) 
100) Pingasa lahayei (Oberthür, 1887) (Nigeria) 
101) Pseudoterpna coronillaria axillaria Guenée, 1857 (C. Lebanon) 
102) Pseudoterpna coronillaria halperini ssp. n. (NE. Israel ; Paratype) 


Figs 103-109. © Genitalia (scale bar = 1 mm) 

103) Pseudoterpna coronillaria cinearascens (Zeller, 1847) (C. Italy) 
104) Pseudoterpna rectistrigaria Wiltshire, 1948 (Cyprus) 

105) Holoterpna pruinosata (Staudinger, 1898) (N. Israel) 

106) Aplasta ononaria (Fuessly, 1783) (N. Israel ; f. faecataria Hübner) 
107) Comibaena bajularia ([Denis & Schiffermüller], 1775) (C. Turkey) 
108) Proteuchloris neriaria (Herrich-Schäffer, 1852) (N. Israel) 

109) Proteuchloris neriaria (Herrich-Schäffer, 1852) (Makedonia) 


85 


Figs 
110) 
111) 
112) 
113) 
114) 
115) 
116) 
117) 
118) 
119) 


nz 


110-119. © Genitalia (scale bar = 1 mm) 

Thetidia (Antonechloris) smaragdaria (Fabricius, 1787) (C. Italy) 

Thetidia (Antonechloris) smaragdaria (Fabricius, 1787) (NW. Caucasus) 
Thetidia (Antonechloris) volgaria (Guenée, 1857) (S. Russia) 

Thetidia (Antonechloris) volgaria (Guenée, 1857) ; vaginal plate only (Armenia) 
Thetidia (Antonechloris) persica sp. n. (N. Iran ; Holotype) 

Thetidia (Antonechloris) persica sp. n. (E. Turkey ; vaginal plate only) 
Thetidia (Antonechloris) persica sp. n. (C. Turkey ; vaginal plate only) 
Thetidia (Antonechloris) silvia Hausmann, 1991 (C. Jordan ; Paratype) 
Thetidia (Antonechloris) bilineata Hausmann, 1991 (C. Jordan ; Holotype) 
Thetidia (Aglossochloris) euryrithra Prout, 1935 (N. Jordan) 


Figs 120-126. © Genitalia (scale bar — 1 mm) 

120) Hemistola chrysoprasaria (Esper, 1795) (S. Turkey) 

121) Xenochlorodes olympiaria cremonaria (Staudinger, 1897) (N. Israel) 

122) Gnophosema palumba Brandt, 1938 (S. Iran) 

123) Victoria sematoperas Prout, 1916 (Kenya ; Paratype) 

124) Victoria plantei Herbulot, 1976 ; ostium-bursae-region only (C. Israel ; Holotype) 
125) Victoria eremita Hausmann, 1993 (S. Israel ; Paratype) 

126) Eucrostes indigenata (de Villers, 1789) (C. Israel) 


Figs 127-136. 9 Genitalia (scale bar = 1 mm) 

127) Thalera fimbrialis magnata A. Fuchs, 1903 (N. Iran) 

128) Culpinia prouti (Thierry-Mieg, 1913) (C. Israel) 

129) Chlorissa viridata (Linnaeus, 1758) (Transcaucasia) 

130) Chlorissa cloraria (Hiibner, [1813]) (C. Greece) 

131) Chlorissa asphaleia Wiltshire, 1966 (N. Turkey) 

132) Phaiogramma pulmentaria (Guenée, 1857) (N. Israel) 

133) Phaiogramma faustinata (Milliere, 1868) (N. Israel) 

134) Neromia pulvereisparsa pulvereisparsa (Hampson, 1896) (N. Oman) 
135) Neromia pulvereisparsa jodisata Staudinger, 1898 (C. Israel) 
136) Neromia simplexa Brandt, 1938 (N. Oman) 


88 


Figs 137-145. 9 Genitalia (scale bar = | mm) 

137) Kuchleria gisisi Hausmann, 1995 (C. Israel ; Paratype) 

138) Microloxia herbaria (Hübner, [1813]) (SE. Turkey) 

139) Microloxia ruficornis (Warren, 1897) (S. Israel) 

140) Acidaliastis micra Hampson, 1896 (S. Israel) 

141) Hemidromodes robusta robusta (Prout, 1913) (NE. Sudan) 
142) Hemidromodes sabulifera sabulifera Prout, 1922 (E. Iran) 

143) Hemidromodes sabulifera hessa Prout, 1935 (S. Israel) 

144) Hierochthonia pulverata (Warren, 1901) (C. Lebanon) 

145) Hierochthonia semitaria (Püngeler, 1901) (C. Israel ; Paratype) 


89 


Figs 146-148. 


146) Microbaena pulchra minor ssp. n., 6, holotype 
147) Thetidia persica sp.n., 2, holotype 


148) Eucrostes indigenata lanjeronica ssp.n., 4, paratype 


Nota lepid. 19 (1/2) : 91-106 ; 21.X1.1996 ISSN 0342-7536 


Systematic list of the geometrid moths of the Levant 
and neighbouring countries 


Part I : Orthostixinae and Geometrinae 


Axel HAUSMANN 


Zoologische Staatssammlung München, Münchhausenstr. 21, D-81247 München, Germany 


Summary 


Specific and subspecific names of the Orthostixinae and Geometrinae (Geo- 
metridae) occurring in the Levant and neighbouring countries are listed. In the 
study area 49 species group names refer to valid taxa : 43 species (occurrence 
of two of these is probable but they are not yet recorded) and six further 
taxa at subspecific level. Citation of original descriptions, locus typicus and 
some important synonyms are given. À brief survey of the distribution of 
the taxa within the study area is added. 


Zusammenfassung 


Art- und Unterartnamen der in der Levante und den umliegenden Ländern 
vorkommenden Arten der Unterfamilien Orthostixinae und Geometrinae 
(Geometridae) werden aufgelistet. Im Untersuchungsgebiet beziehen sich 49 
Namen der Artgruppe auf giiltige Taxa: Neben 43 Arten (in zwei Fallen ist 
das Vorkommen zu erwarten, aber nicht abgesichert) sind sechs weitere Taxa 
auf Unterart-Niveau verbreitet. Es werden Originalbeschreibung, locus typicus 
und einige wichtige Synonyme erwähnt. Ein kurzer Uberblick über die 
Verbreitung der Taxa im Untersuchungsgebiet wird angefiigt. 


Introduction 


This is the first paper in the second of three series of publications 
by the author on the Geometridae of the Middle East. The three series 
are 


1) Morphology of the species and subspecies occurring in the Levantine 
basin and neighbouring countries ; with figures of male and female 
genitalia of all taxa, of venation of all genera, description of external 
structure of the imago and discussion of relationships at subspecies, 
species, genus and tribus level. 


91 


2) Systematic list of these species with some important synonyms, 
citation of original descriptions, locus typicus and a short survey of 
the geographical distribution within the study area. 


3) Faunistic data, phenology and ecology of the species occurring in 
the state of Israel (administration frontiers of 1990). 


The present publication initiates the second series of publications. There 
are some differences from the fairly well accepted list of Leraut (1980). 
In addition to the data in the present paper there is information on 
the reasons for these changes in the discussions on tribal systematics 
in the first series (morphology). By including many additional Old 
World genera and tribes it should facilitate the systematic arrangement 
for any future Palaearctic Orthostixinae or Geometrinae Fauna. A 
systematic check list of species occurring in the Levant is provided 
in Tab. 1. 


Tab. 1. Systematic checklist of species and subspecies occurring in the study area 
(Subfamilies Orthostixinae and Geometrinae) 


Orthostixinae 
Myinodes shohami Hausmann, 1994 
Eumegethes tenuis Staudinger, 1898 
Orthostixis cribraria amanensis Wehrli, 1932 
Orthostixis cinerea Rebel, 1916 
Orthostixis calcularia Lederer, 1853 

Geometrinae 

PSEUDOTERPNINI Warren, 1893 
Pingasa lahayei multispurcata Prout, 1913 
Pseudoterpna coronillaria axillaria Guenée, 1857 
Pseudoterpna coronillaria halperini Hausmann, 1996 
Pseudoterpna rectistrigaria Wiltshire, 1948 
Holoterpna pruinosata (Staudinger, 1898) 
Aplasta ononaria (Fuessly, 1783) 

COMIBAENINI Inoue, 1961 
Microbaena pulchra (Staudinger, 1898) 
Comibaena bajularia ([Denis & Schiffermüller], 1775) 
Proteuchloris neriaria (Herrich-Schäffer, 1852) 
Thetidia (Antonechloris) persica Hausmann, 1996 
Thetidia (Antonechloris) silvia Hausmann, 1991 
Thetidia (Antonechloris) bilineata Hausmann, 1991 
Thetidia (Aglossochloris) euryrithra Prout, 1935 

HEMISTOLINI Inoue, 1961 
Hemistola chrysoprasaria (Esper, 1795) 
Xenochlorodes olympiaria cremonaria (Staudinger, 1897) 
Gnophosema palumba kurdistana Ebert, 1969 
Victoria (Victorinella) wiltshirei Hausmann, 1996 
Victoria (Victorinella) plantei Herbulot, 1976 
Victoria (Victorinella) eremita Hausmann, 1993 
Victoria (Victorinella) fifensis Wiltshire, 1994 

COMOSTOLINI Inoue, 1961 
Eucrostes indigenata (de Villers, 1789) 


92 


THALERINI Herbulot, 1963 
Thalera fimbrialis magnata A. Fuchs, 1903 
Culpinia prouti (Thierry-Mieg, 1913) 
HEMITHEINI Bruand, 1846 
Chlorissa cloraria (Hübner, [1813 |) 
Chlorissa asphaleia Wiltshire, 1966 
Phaiogramma pulmentaria (Guenée, 1857) 
Phaiogramma faustinata (Milliere, 1868) 
Phaiogramma discessa (Walker, 1861) 
Phaiogramma polemia (Prout, 1920) 
Neromia pulvereisparsa pulvereisparsa (Hampson, 1896) 
Neromia pulvereisparsa jodisata Staudinger, 1898 
Neromia simplexa Brandt, 1938 
Kuchleria gisisi Hausmann, 1995 
MICROLOXIINI Hausmann, 1996 
Microloxia herbaria (Hübner, [1813]) 
Microloxia ruficornis (Warren, 1897) 
Acidaliastis micra Hampson, 1896 
Hemidromodes robusta robusta (Prout, 1913) 
Hemidromodes robusta affinis (Rothschild, 1915) 
Hemidromodes robusta galala Wiltshire, 1949 
Hemidromodes sabulifera sabulifera Prout, 1922 
Hemidromodes sabulifera hessa Prout, 1935 
Hemidromodes sabulifera triforma Wiltshire, 1949 
Hierochthonia pulverata (Warren, 1901) 
Hierochthonia semitaria (Püngeler, 1901) 


The delimitation of the study area is based on the following zoogeo- 
graphical patterns : 


1) Many tropical and subtropical African species range northwards 
to C. Sudan and SW. Saudi Arabia. They are not the subject of this 
paper. For this reason the Sudan and also the Asir mountains of Saudi 
Arabia (northwards to Mecca) are excluded. The western limit of the 
study area (the political border between Egypt and Libya) is drawn 
arbitrarily as there are few zoogeographical discontinuities in the region. 


2) Many species occurring in S. Iran (e.g. some tropical Asiatic 
elements) find their western limit of distribution in the Oman. They 
are not the subject of this paper. For this reason the southeastern parts 
of the Arabian peninsula are excluded from the study area. 


3) The species inhabiting the W. Iranian heights are not the subject 
of this paper. They often extend to the E. Turkish mountains (e.g. 
Hakkari) east of the River Tigris. The study area is extended to the 
mountains of N. Iraq in order to allow the inclusion of the complete 
Iraqi fauna. 


4) Many C. European or N. Mediterranean species have their eastern 
limit of distribution in N. Turkey or Armenia. They are not the subject 
of this paper. 


93 


These zoogeographical patterns define the study area, which includes 
S. Turkey south of the summits of Taurus between Göksu and the 
River Tigris, Cyprus, Syria, Lebanon, Iraq, Kuwait, N. Saudi Arabia 
(north of a line between Medina and Kuwait), Israel, Jordan, Egypt 
(see Fig. 1). 


ARTA R (compare Wiltshire, 1990) : 
endemic 

FO Euroriental 

SE South European 

Al Anatolian-Iranian 

EM East Mediterranean 

HM  Holomediterranean 

SM South Mediterranean 

PE Pan-Eremic 

WE Western Eremic 

EE Eastern Eremic 

SA Saharo-Arabian 

SS Saharo-Sindian 

TA Tropical Asiatic 

PT  Palaeotropical 

Eth  Ethiopian (Afrotropical) 


N., W., E., S., C. = North, West, East, South, Central [country respectively] 


When part of a country is not specified as above, the taxon occurs 
throughout. 


SYSTEMATIC PART 
ORTHOSTIXINAE Meyrick, 1892 


Oenochrominae auct. nec Guenée, 1857 
MYINODES Meyrick, 1892 
Myinodes shohami Hausmann, 1994 
Myinodes shohami Hausmann, 1994 : Nota lepid. 17 (1/2) : 38 (loc. typ. : 
Qasr el Hallabad, Jordan) 
Pseudotagma interpunctaria sensu Staudinger, 1892 : Dt. Ent. Z. Iris 4: 
168 (from Beirut and Jerusalem) (nec Herrich-Schäffer, 1839: pl. 6 
and wrapper ; loc. typ. : Sicily). 
RANGE: S. Turkey, N. Irag, Lebanon, ? S. Syria, N. Jordan, 
N. + C. Israel (AI ; also in SW. Iran) 


94 


‘(1x91 995) taie Apnys ‘I SU 


95 


EUMEGETHES Staudinger, 1898 
Eumegethes tenuis Staudinger, 1898 
Thalpochares (Eumegethes) tenuis Staudinger, 1898 : Dt. Ent) ZATas0= 
268, Pl. 4, Fig. 6 (loc. typ. : Sfax, E. Tunisia) 
Lomosrapha petroffi Andres & Seitz, 1923 : Senckenbergiana 7 : 60 (loc. 
typ. : LEgypt]) 
Eumegethes picta Turati, 1934: Atti Soc. Ital. Sci. Nat. 73: 176, pl. 3, 
fig. 14 (loc. typ. : Porto Bardia, E. Libya) 
RANGE: W. Egypt (SM) 
ORTHOSTIXIS Hübner, [1823] 
Orthostixis cribraria (Hübner, [ 17991) 
Geometra cribraria Hübner, [1799]: Samml. Eur. Schmett. Geom. t. 16, 
f. 83 (loc. typ. : [Europe]). 
Phalaena laetata Fabricius, 1798: Suppl. Ent. Syst. p. 456 (loc. typ. : 
„S. Russia”; junior primary homonym to Phalaena laetata Fabricius, 
1784 : Ent. Syst. 3 (2) : 164) 
Orthostixis cribraria amanensis Wehrli, 1932 
Orthostixis cribraria Hb. amanensis Wehrli, 1932 : Mitt. münch. ent. Ges. 
22 : 3 (loc. typ. : Düldül Dagh, Amanus) 
RANGE : S. Turkey, N. and C. Lebanon (ssp. e; sp. EM, with 
Sicily, Sam Ce. Italy) 
Orthostixis cinerea Rebel, 1916 
Orthostixis cinerea Rebel, 1916 : Jahresb. Wien. Ent. Ver. 26 : 108 (loc. 
typ. : Nicosia, Cyprus) 
RANGE : Cyprus (e) 
Orthostixis calcularia Lederer, 1853 
Orthostixis calcularia Lederer, 1853 : Verh. Zool.-Bot. Ver. Wien 3 : 260 
(loc. typ. : Elisabethpol, Transcaucasus) 


RANGE: S. Turkey, ,,Syria” (Prout, 1912) (AJ) 
GEOMETRINAE Duponchel, [1845] 


PSEUDOTERPNINI Warren, 1893 


Pseudoterpninae Warren, 1893: Proc. Zool. Soc. London 1893: 349. 
Availability of the name “Pseudoterpnini” based on article 36 ICZN ; 
Type genus according to article 63 ICZN Pseudoterpna Hübner, [1823], 
although not listed in the original publication, but Pseudoterpna 
belongs to Pseudoterpninae as suggested by Warren. Next available 
name would be Pseudoterpnini Herbulot, 1963. 

Terpnini Inoue, 1961, nec Kusnetzov, 1904, homonym, not available. 
Based on Terpna Herrich-Schäffer, [1854], which is an incorrect sub- 
sequent spelling of Zerpne Hübner, 1822. In addition homonym to 
Terpnidae Kusnetzov, 1904 : Horae Ent. Soc. Ross. 37 : 53. The latter 


96 


based on Terpne Hbn., junior objective synonym to Geometra Linnaeus, 
1758. Terpnini Kusnetzov therefore synonym to Geometrini. 
PINGASA Moore, [1887] 
Pingasa lahayei (Oberthür, 1887) 
Hypochroma lahayei Oberthür, 1887 : Bull. Soc. Ent. Fr. 1887 : LEX (loc. 
typ. : Ain-Sefra, NW. Algeria) 
Pingasa lahayei multispurcata Prout, 1913 
Pingasa multispurcata Prout, 1913 : Novit. Zool. 20 : 397 (loc. typ. : Rawal 
Pindi, N. Pakistan) 
RANGE : W. Saudi Arabia, C. and S. Iraq (ssp. TA ; sp. PT) 
PSEUDOTERPNA Hübner, [1823] 
Pseudoterpna coronillaria (Hübner, [1817]) 
Geometra coronillaria Hübner, [1817]: Samml. Eur. Schmett. Geom. : 
pl. 93, fig. 479-482 (loc. typ. : [Europe : S. France, Corsica or Spain]) 
Pseudoterpna coronillaria axillaria Guenée, 1857 
Pseudoterpna axillaria Guenée, 1857 : Spec. Gén. Lép. 9 : 339 (loc. typ. : 
Beirut, Lebanon) 
RANGE : Lebanon, ? S. Turkey (Prout 1913 : 20 “Taurus”), ? Cyprus, 
? NW. Syria (ssp. EM, sp. HM) 
Pseudoterpna coronillaria halperini Hausmann, 1996 
Pseudoterpna coronillaria halperini Hausmann, 1996: Nota lepid. 19 
(1/2) : 16 (loc. typ. : Senir, NE. Israel) 
RANGE: N. Jordan, N. + C. Israel (ssp. e, sp. HM) 
Pseudoterpna rectistrigaria Wiltshire, 1948 
Pseudoterpna coronilleria Hb. (sic !) ab. rectistrigaria Rebel, 1939 : Mitt. 
münch. ent. Ges. 29: 531, pl. 15, fig. 12. (not available, see Haus- 
mann, 1994 : 81) 
Pseudoterpna rectistrigaria Wiltshire, 1948 : Entomologist’s Rec. J. Var. 
15 : 81, fig. B (loc. typ. : Cyprus) 
RANGE : Cyprus (e) 
HOLOTERPNA Püngeler, 1900 
Holoterpna pruinosata (Staudinger, 1898) 
Eucrostis (?) pruinosata Staudinger, 1898 : Dt. Ent. Z. Iris 10 : 303 (loc. 
typ. : Jerusalem, Israel) 
RANGE: Lebanon, N. + C. Israel (EM, furthermore NE. Italy: 
Trieste, probably introduced by ship) 
APLASTA Hübner, [1823] 
Aplasta ononaria (Fuessly, 1783) 
Phalaena ononaria Fuessly, 1783 : Arch. Insectengesch. 3: 1, t. 17, f. 4-6 
(loc. typ. : not stated, probably [S. Europe]) 
Geometra faecaturia Hübner, [1819]: Samml. eur. Schmett. 5, Geom. : 
pl. 97, fig. 503 (loc. typ. : [Europa]) 


97 


Aplasta faecataria Hübner, [1823]: Verz. Bek. Schmett. : 304 (justified 
emendation). 

Aplasta ononaria ab.? (v.?) berytaria Staudinger, 1901: Cat. Lep. 
Palaearct. 1 : 261 (loc. typ. : Beirut, Lebanon). 

RANGE: S. Turkey, Cyprus, N. Iraq, Lebanon, Syria, N. Israel, 
N. Jordan (EO) 


COMIBAENINI Inoue, 1961 

Euchlorini Herbulot, 1963 

MICROBAENA Hausmann, 1996 
Microbaena pulchra (Staudinger, 1897) 

Phorodesma pulchra Staudinger, 1897 : Dt. Ent. Z. Iris 10 : t. 4, fig. 27, 
(p. 302 : 1898) (loc. typ. : Jerusalem or Jaffa, Israel) 

RANGE : C. Israel (Eth) F 

COMIBAENA Hübner, [1823] 
Comibaena bajularia (| Denis & Schiffermüller], 1775) 

Phalaena pustulata Hufnagel, 1767 : Berlinisches Magazin : 520 (loc. typ. : 
Berlin distr. ; junior primary homonym to Phalaena pustulata O. F. 
Müller, 1764: Faun. Frid. 46 ; identity Hyphoraia aulica (Linnaeus, 
1758), Arctiidae) 

Geometra (Phalaena) bajularia [Denis & Schiffermiiller], 1775 : Schmett. 
Wien : 97 (loc. typ. : Vienna, Austria). 

RANGE: S. Turkey (EO) 

PROTEUCHLORIS Hausmann, 1996 
Proteuchloris neriaria (Herrich-Schäffer, 1852) 

neriaria Herrich-Schäffer, 1848 : Syst. Bearb. Schm. Eur. 3 : t. 70, fig. 429 
(uninominal, no description, not available). 

[Geometra (Phorodesma)] neriaria Herrich-Schäffer, 1852 : Syst. Bearb. 
Schm. Eur. 6 : 62 (nec “Friv.”; loc. typ. : Crete ; available : description 
in the supplements of Vol. 6 apparently uninominal with species name 
only, but there is precisely stated position in text of Vol. 6, where 
the description has to be inserted. Thus the combined generic and 
subgeneric indication that is given in text part of Vol. 6, is valid also 
for the supplementary description of neriaria). 

RANGE: S. Turkey, Lebanon, N. Israel (EM) 

THETIDIA Boisduval, 1840 
subgenus ANTONECHLORIS Raineri, 1994 
Thetidia (Antonechloris) smaragdaria (Fabricius, 1787) 

Phalaena smaragdaria Fabricius, 1787: Mant. Ins. 2: 192 (loc. typ. : 
Austria) 

RANGE : not represented in the study area, but discussed in Haus- 
mann (1996) with regard to T. persica. 


98 


Thetidia (Antonechloris) volgaria (Guenée, 1857) 
Geometra prasinaria Eversmann, 1837 : Bull. Soc. Imp. Nat. Mosc. 10 (2) : 
52 (loc. typ. : Orenburg, S. Russia ; junior primary homonym to Geo- 
metra prasinaria Linnaeus). 
Geometra volgaria Guenée, 1857: Spec. Gén. Lép. 9: 344 (loc. typ. : 
Orenburg, S. Russia). 
RANGE : not represented in the study area, but discussed in Haus- 
mann (1996) with regard to T. persica. 
Thetidia (Antonechloris) persica Hausmann, 1996 
Euchloris smaragdaria F. ssp. anomica sensu Schwingenschuss, 1940 (Ent. 
Z. 53 : 202 : Pelur, N. Iran) nec Prout, 1935 : 17 (loc. typ. : Issyk Kul, 
C. Asia). 
Thetidia persica Hausmann, 1996: Nota lepid. 19 (1/2): 27 (loc. typ. : 
Tacht i Suleiman, N. Iran). 
RANGE: S. Turkey, N. Iraq (AI) 
Thetidia (Antonechloris) silvia Hausmann, 1991 
Thetidia silvia Hausmann, 1991: Mitt. münch. ent. Ges. 81: 117, t. 1, 
figs. 10, 11, t. 10, fig. 71 (loc. typ. : Shaubak, Jordan). 
RANGE: C. Jordan (e) 
Thetidia (Antonechloris) bilineata Hausmann, 1991 
Thetidia bilineata Hausmann, 1991 : Mitt. münch. ent. Ges. 81: 117, t. 1, 
fig. 8, t. 10, fig. 72 (loc. typ. : Shaubak, Jordan). 
RANGE: C. Jordan (e) 
subgenus AGLOSSOCHLORIS Prout, 1912 
Thetidia (Aglossochloris) euryrithra Prout, 1935 
Aglossochloris crucigerata euryrithra Prout, 1935: in Seitz Macrolep. 
4 Suppl. : 18, fig. 3c (loc. typ. : Amman, Jordan) 
RANGE: N. Jordan (e) 


HEMISTOLINI Inoue, 1961 
HEMISTOLA Warren, 1893 
Hemistola chrysoprasaria (Esper, 1795) 

Phalaena (Geometra) vernaria {Denis & Schiffermiiller], 1775 : Ank. Verz. 
Schmett. Umg. Wien: 97 (loc. typ.: Vienna distr. ; junior primary 
homonym to Phalaena (Geometra) vernaria Linnaeus, 1761 : Fauna 
Svec. (2) : 323, identity Jodis lactearia (Linnaeus, 1758)) 

Phalaena (Geometra) chrysoprasaria Esper, 1795 : Schmett. in Abbild. 5: 
37, t. 5, fig. 1 (loc. typ. : first locality mentioned “Vienna” but “Originals 
from Frankfurt”) 

Hemistola immaculata auct. nec Thunberg, 1784 (“Geometra immaculata”; 
cf. Karsholt & Schmid Nielsen, 1985 : 448) 

Hemistola biliosata auct. nec de Villers, 1789 (“ Phalaena Geometra bilio- 
sata” Linn. Ent. 2: 386, pl. 6, fig. 22). Recently some authors have 


99 


treated biliosata as the valid species name for this taxon (e.g. Skou, 
1986 : 28 ; Rezbanyai-Reser, 1993 : Boll. Soc. Tic. Sci. Natur. 81 (1): 
47). Shape of postmedian line of forewing in original illustration of 
biliosata however clearly proves its true identity to be Hylaea fasciaria 
f. prasinaria. 
RANGE: S. Turkey (EO) 
XENOCHLORODES Warren, 1897 
Xenochlorodes olympiaria (Herrich-Schäffer, 1852) 
olympiaria Herrich-Schäffer, 1851 : Syst. Bearb. Schmett. Eur. 3: pl. 87, 
fig. 539 (loc. typ. : not stated ; uninominal, no text, not available) 
Geometra (Eucrostes) olympiaria Herrich-Schäffer, 1852 : Syst. Bearb. 
Schmett. Eur. 6 : 63 (loc. typ. : Olymp near Brussa, NW. Turkey) 
Geometra beryllaria Mann, 1853 : Verh. Zool.-Bot. Ver. Wien 3 : 76 (loc. 
typ. : Spalato, Dalmatia) 
Nemoria aureliaria Milliere, 1864 : Ann. Soc. Linn. Lyon (N.S.) 11 : 268 
(Icon. Chen. 2 : 37), pl. 55, fig. 1, 2 (loc. typ. : S. Italy) 
Xenochlorodes olympiaria cremonaria (Staudinger, 1897) 
Eucrostis olympiaria HS. var. ? cremonaria Staudinger, 1897 : Dtsch. Ent. 
Z. Iris 10 : 179 (loc. typ. : Haifa, Israel) 
Xenochlorodes pallida Warren, 1897: Novit. Zool. 4: 47 (loc. typ. : 
Lebanon, Beirut) 
RANGE: S. Turkey, Cyprus, Lebanon, N. + C. Israel (ssp. EM ; 
sp. HM) 
GNOPHOSEMA Prout, 1912 
Gnophosema palumba Brandt, 1938 
Gnophosema palumba Brandt, 1938 : Ent. Rndsch. 55 (49) : 572, figs. 238- 
243 (loc. typ. : Mian-Kotal, SW. Iran) 
Gnophosema palumba kurdistana Ebert, 1969 
Gnophosema palumba kurdistana Ebert, 1969: Arkiv för Zool. 22 (6): 
193, figs. 16, 17, gen.fig. 11 (loc. typ. : Shaqlawa, N. Iraq) 
RANGE : N. Irag (ssp. e ; sp. Al) 
VICTORIA Warren, 1897 
subgenus Victorinella Hausmann, 1996 
Victoria (Victorinella) wiltshirei Hausmann, 1996 
Victoria sematoperas sensu Wiltshire, 1949 : Bull. Soc. Fouad ler d’Ent. 
33 : 399 (from Wadi Aideb, S. Egypt). Not available, homonym to 
Victoria sematoperas Prout, 1916: Proc. Zool. Soc. London 1916: 
142, pl. I, Fig. 26 (loc. typ. : Mandere, S. Somalia). 
Victoria wiltshirei Hausmann, 1996: Nota lepid. 19 (1/2) : 39 (loc. typ. : 
Ed Damer, NE. Sudan) 


RANGE: S. Egypt, N. Sudan (e) 


100 


Victoria (Victorinella) plantei Herbulot, 1976 
Victoria plantei Herbulot, 1976: Alexanor 9: 290 (loc. typ. : Sodom, 
C. Israel) 
RANGE : C. Israel (e) 
Victoria (Victorinella) eremita Hausmann, 1993 
Victoria eremita Hausmann, 1993 : Spixiana 16 (1) : 55 (loc. typ. : Yotvata, 
S. Israel) 
RANGE : S. Israel (e) 
Victoria (Victorinella) fifensis Wiltshire, 1994 
Victoria fifensis Wiltshire, 1994: Fauna of Saudi Arabia 14: 115, pl. 1, 
figs. 5-6 (loc. typ. : Tarima, W. Saudi Arabia) 
RANGE : W. Saudi Arabia (e) 


COMOSTOLINI Inoue, 1961 
EUCROSTES Hübner, [1823] 
Eucrostes indigenata (de Villers, 1789) 
Phalaena indigenata de Villers, 1789: Caroli Linn. Ent. 2: 383, pl. 6, 
fig. 19 (loc. typ. : Europe, “Le Content”) 
Geometra fimbriolaria Hübner, [1817] 1796: Samml. Eur. Schmett. 5: 
pl. 91, fig. 468 (loc. typ. : [Europa]) 
RANGE : Lebanon, N. + C. Israel, Jordan (HM) 


THALERINI Herbulot, 1963 
Chlorochromites Duponchel, [1845], (“sub-tribe”), nom. invalid. : “Chloro- 
chromini” valid according to Seven (1991 : 44) and including also those 
species placed here in Hemitheini. Based on the type genus Chloro- 
chroma Duponchel, [1845], which is an objective junior synonym of 
Thalera Hübner, [1823] (same type species). Family group name 
Chlorochromites forgotten for about 100 years, but “Thalerini” accepted 
e.g. by Leraut (1980), Vives Moreno (1994) and Viidalepp (1981) (cf. 
also Holloway, 1993 : 4). 
Thalerini Herbulot, 1963 : Tribal name not introduced by Inoue, 1961, 
as suggested by Viidalepp (1981 : 95). 
THALERA Hübner, [1823] 
Thalera fimbrialis (Scopoli, 1763) 
Phalaena fimbrialis Scopoli, 1763 : Ent. Carniolica : 216 (loc. typ. : Car- 
niolia (Slowenia)) 
Thalera fimbrialis magnata A. Fuchs, 1903 
Thalera fimbrialis ab. (var. ?) magnata A. Fuchs, 1903 : Jahrb. Nass. Ver. 
Nat. 56 : 53 (loc. typ. : Tura, Transcaspia) 


101 


Thalera fimbrialis major Warnecke, 1930 : Int. Ent. Z. Guben 24: 133, 
pl. 1 figs. 10-11 (loc. typ. : Issyk Kul, Kirgizia) 
RANGE: S. Turkey, Lebanon (ssp. AI with C. Asian mountains, 
sp. EO) 
CULPINIA Prout, 1912 
Culpinia prouti (Thierry-Mieg, 1913) 
Thalera prouti Thierry-Mieg, 1913 : Feuille Jeun. Nat. 43 : 180 (loc. typ. : 
Akbes, S. Turkey) | 
Thalera feroniaria Oberthür, 1916: Lep. Comparée 12: 123 (loc. typ. : 
Akbes, S. Turkey) 


RANGE: S. Turkey, Lebanon, C. Israel (EM) 


HEMITHEINI Bruand, 1846 
CHLORISSA Stephens, 1831 
Chlorissa viridata (Linnaeus, 1758) 
Phalaena viridata Linnaeus, 1758 : Syst. Nat (10) 1: 523 (loc. typ. : not 
stated [Europe]) 
RANGE : Not represented in the study area, but discussed in Haus- 
mann (1996) with regard to C. cloraria. 
Chlorissa cloraria (Hübner, [1813]) 
Geometra cloraria Hübner, [1813]: Samml. Eur. Schmett. 5: pl. 68, 
fig. 352 (loc. typ. [Europe]) 
RANGE : C. Turkey, probably reaching the study area in the Taurus 
Mountains (EO) 
Chlorissa asphaleia Wiltshire, 1966 
Chlorissa asphaleia Wiltshire, 1966 : Zeits. Wien. Ent. Ges. 51: 30, pl. 2, 
fig. 3 (loc. typ. : Derbend, N. Iran). 
RANGE : E. Turkey, perhaps reaching the study area in the Taurus 
mountains (Al) 
PHAIOGRAMMA Gumppenberg, 1887 
Phaiogramma pulmentaria (Guenée, 1857) 
etruscaria Zeller, 1849 : Ent. Z. [Stettin] : 203 (loc. typ. : Livorno, N. Italy ; 
uninominal in the text, binominal in the index ; name forgotten for 
about 100 years) 
Nemoria pulmentaria Guenée, 1857 : Spec. Gén. Lép. Phal. 9 : 349 (loc. 
typ. : S.-Frankreich (Dalmatien, Italien)) 
Nemorla (sic !) pulmentaria Gn. var. palaestinensis Fuchs, 1903 : Societ. 
Entom. 18 : 51 (loc. typ. : Jerusalem) 
RANGE: S. Turkey, N. + NW. Irag, Lebanon, N. + C. Israel, N. 
Jordan (EO) 


102 


Phaiogramma faustinata (Milliere, 1868) 
Nemoria faustinata Milliere, 1868 : Ann. Soc. Linn. Lyon (N.S.) 17 : 26 
(Icon. Chen. 2 : 436) (loc. typ. : Barcelona, Spain) 
RANGE : Cyprus, C. Lebanon, Israel, N. Jordan, W. Saudi Arabia, 
Egypt (Eth) 
Phaiogramma discessa (Walker, 1861) 
lodis discessa Walker, 1861 : List. Lep. Ins. Brit. Mus. 22 : 544 (loc. typ. : 
North Hindustan) 
? Chlorissa patialensis Rose & Devinder, 1985 : Journ. Bomb. Nat. Hist. 
Soc. 82 : 614, figs. 1-5 (loc. typ. : Patiala, NW. India) 
RANGE : E. Saudi Arabia (north-western limit of distribution 
Kuwait ?) (TA) 
Phaiogramma polemia (Prout, 1920) 
Microloxia polemia Prout, 1920: Novit. Zool. Tring 27 : 300 (loc. typ. : 
Kut-al-Amara, C. Iraq) 
RANGE: C. + S. Iraq (e ; also SW. Iran) 
NEROMIA Staudinger, 1898 
Neromia pulvereisparsa (Hampson, 1896) 
Nemoria pulvereisparsa Hampson, 1896: Proc. Zool. Soc. Lond. 1896: 
268, pl. 10, fig. 27 (loc. typ. : Aden, S. Yemen) 
RANGE: C. + S. Iraq, Saudi Arabia (EE) 
Neromia pulvereisparsa jodisata Staudinger, 1898 
Nemoria ? (Neromia) jodisata Staudinger, 1898 : Dt. Ent. Ztschr. Iris 10: 
304, pl. 4, fig., 28 (loc. typ. : Israel/ Jordan, Jordan Valley) 
Chlorissa draudti Andres-Seitz, 1924: Senckenbergiana VI (1/2): 70, 
fig. 19 (loc. typ. Wadi Hof near Heluan, Egypt). 
RANGE : C. + E. Egypt, S. + C. Israel, S. Jordan (ssp. e ; sp. EE) 
Neromia simplexa Brandt, 1938 
Neromia simplexa Brandt, 1938 : Entom. Rundsch. 55 : 572, fig. 219 (loc. 
typ. : Fort Mian-Kotal, SW. Iran) 
RANGE: C. Iraq (according to Heydemann et al. 1963: 104), E. 
Saudi Arabia (near Kuwait) (EE) 
KUCHLERIA Hausmann, 1995 
Kuchleria gisisi Hausmann, 1995 
Kuchleria gisisi Hausmann, 1995: Atalanta 25 (3/4): 588, fig. 49 (loc. 
typ. : Tel Aviv, C. Israel). 
RANGE : C. Israel (e) 


MICROLOXIINI Hausmann, 1996 
MICROLOXIA Warren, 1893 
Microloxia herbaria (Hübner, [1813]) 


103 


Geometra herbaria Hübner, [1813]: Samml. Eur. Schmett. 5: pl. 79, 
fig. 407 (loc. typ. [Europa]) 
RANGE: S. Turkey, Cyprus, Lebanon, C. Iraq, 23 Israel, heights 
of C. Israel (EO) 
Microloxia ruficornis Warren, 1897 
Microloxia ruficornis Warren, 1897 : Novit. Zool. 4 : 42 (loc. typ. : Natal, 
S. Africa). 
Eucrostes halimaria Chretién, 1909: Le Naturaliste 31: 18 (loc. typ.: 
Biskra, Algeria). 
Microloxia indecretata auct. nec Walker, [1863] (List Specimens lepid. 
Insects Collin Br. Mus. 26 : 1555 (loc. typ. : South Hindustan)) 
RANGE: Egypt, Saudi Arabia, C. + S. Israel, S. Jordan, Kuwait, 
S. Iraq (PT) 
ACIDALIASTIS Hampson, 1896 
Acidaliastis micra Hampson, 1896 
Acidaliastis micra Hampson, 1896: Proc. zool. Soc. London 1896 : 267, 
pl. 10, fig. 20 (loc. typ. : S. Yemen, Aden) 
Euchloris dissimilis Warren & Rothschild, 1905 : Novitates Zool. 12 : 26; 
pl. 4, fig. 27 (loc. typ. : Nakheila, Sudan) 
Eucrostes ? desertoria Rebel, 1909: Verh. Nat. Ver. Karlsruhe 21: 137 
(loc. typ. : Sinai). 
RANGE: S. + E. Egypt, Saudi Arabia, S. + C. Israel, S. Jordan (SA) 
HEMIDROMODES Prout, 1916 
Hemidromodes robusta (Prout, 1913) 
Hierochthonia robusta Prout, 1913 : Novit. Zool. 20 : 435 (loc. typ. : Port 
Sudan) 
RANGE: C. + S. Egypt (also Sudan ; ssp. e ; sp. WE) 
Hemidromodes robusta affinis (Rothschild, 1915) 
Hemidromodes affinis Rothschild, 1915: Ann. Mag. Nat. Hist. 16 
(8th series) : 392 (loc. typ. : Tahihout, Hoggar, Algeria) 
Hierochtonia debonoi Kriiger, 1939: Ann. Mus. Libico Stor. Nat. 1: 
352, pl. 16, figs. 109-110 (loc. typ. : Gebel es-Soda, Tripolitania, Libya) 
RANGE : NW. Egypt ? (ssp. e ?, sp. WE) 
Hemidromodes robusta galala Wiltshire, 1949 
Hemidromodes affinis galala Wiltshire, 1949 : Bull. Soc. Fouad ler d’Ent. 
33 : 401 ; pl. 9, figs. 3, 4 (loc. typ. : Suez Road (km 100), E. Egypt). 
RANGE: E. Egypt (ssp. e ; sp. WE) 
Hemidromodes sabulifera Prout, 1922 
Hemidromodes sabulifera Prout, 1922 : Bull. Hill Mus. | : 257 (loc. typ. : 
Bombay, W. India) 
RANGE: C. and E. Saudi Arabia (also S. Iran ; TA) 


104 


Hemidromodes sabulifera hessa Prout, 1935 
Hemidromodes sabulifera hessa Prout, 1935 : in Seitz Macrolep. 4, Suppl. : 
16, fig. 6b (loc. typ. : Ghor el Safieh, S. Dead Sea, Jordan) 
RANGE: S. + C. Israel, S. + C. Jordan (ssp. e ; sp. TA) 
Hemidromodes sabulifera triforma Wiltshire, 1949 
Hemidromodes robusta triforma Wiltshire, 1949 : Bull. Soc. Fouad ler 
d’Ent. 33 : 363 ; figs. 10, 11 (loc. typ. : Hejaz, Saudi Arabien) 
RANGE: W. Saudi Arabia (ssp. e ; sp. TA) 
HIEROCHTHONIA Prout, 1912 
Hierochthonia pulverata (Warren, 1901) 
Microloxia pulverata Warren, 1901 : Novit. Zool. 8 : 193 (loc. typ. : Leba- 
non, Beirut) 
RANGE: S. Turkey, Lebanon (EM) 
Hierochthonia semitaria (Püngeler, 1901) 
Eucrostes semitaria Piingeler, 1901: Dtsch. Ent. Z. Iris 14: 333 (Lt.: 
Dead Sea, near Qumran, C. Israel) 


RANGE : N. + C. Israel, N. + C. Jordan (e) 
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REBEL, H., 1939. Zur Lepidopterenfauna Cyperns. Mitt. münch. ent. Ges. 
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SKOU, P., 1986. The Geometrid Moths of North Europe. Entomonograph, 
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VIIDALEPP, J., 1981. The systematic of the genera of the subfamily Geometrinae 
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Vives Moreno, A., 1994. Catalogo sistematico y sinonimico de los Lepi- 
dopteros de la Peninsula Iberica y Baleares II., Madrid. 

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species and thirty-five new records from Cyprus. Entomologist’s Rec. 
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106 


Nota lepid. 19 (1/2) : 107-112 ; 21.X1.1996 ISSN 0342-7536 


Sciopetris karsholti, eine neue Psychide aus Tunesien 
(Lepidoptera, Psychidae) 


Peter HATTENSCHWILER 


Seeblickstrasse 4, CH-8610 Uster. Schweiz 


Summary 


A Danish group of entomologists collected in a oak forest in Tunisia a series 
of male specimen of a primitive Psychidae moth of the subfamily Taleporiinae, 
tribus Taleporiüni. Sciopetris karsholti sp. n. is described and provisionally 
assigned to the genus Sciopetris Meyrick. 


Résumé 


Un groupe d’entomologistes danois a capturé dans une forêt de chênes en 
Tunisie une série de mâles d’une espèce primitive de Psychidae appartenant 
à la sous-famille des Taleporiinae, Tribus Taleporiini. Description de la nouvelle 
espèce, Sciopetris karsholti sp .n., provisoirement attribuée au genre Sciopetris 
Meyrick. 


Zusammenfassung 


Eine dänische Gruppe von Entomologen sammelte in einem Eichenwald in 
Tunesien eine Serie von primitiven Psychidae Männchen der Unterfamilie 
Taleporiinae, Tribus Taleporiini. Die neue Art, Sciopetris karsholti sp. n. wird 
beschrieben und provisorisch in die Gattung Sciopetris Meyrick gestellt. 


Einige Mitarbeiter des Zoologischen Museums in Kopenhagen sam- 
melten 1988 in Tunesien. Sie erbeuteten in einem Eichenwald im ,,Ain 
Draham“ Gebiet (Abb. 1) über mehrere Tage verteilt eine Serie von 
25 Männchen einer unbekannten Art, die als eine primitive Psychide 
erkannt wurde. Trotz sehr intensivem Suchen gelang es nicht die Säcke 
zu finden. Auch von den Weibchen fehlt jede Spur. Es ist anzunehmen, 
dass sie ungefliigelt sind und so in oder bei den Säcken bleiben. Falls 
sie gefliigelt sind und auch fliegen, ware die Wahrscheinlichkeit doch 
recht gross, dass neben den vielen Männchen auch einzelne Weibchen 
gefangen worden waren. Das sind aber alles Vermutungen, die Antwort 
kennen wir noch nicht. 


107 


Abb. 1. Eichenwald am Fundort der S. karsholti sp. n. im Ain Draham Gebiet in 
Tunesien. (Foto : ©. Karsholt). 


Nach den Männchen ist diese neue Art in die Unterfamilie Taleporiinae 
Tutt, 1900 und Tribus Taleporiini Tutt, 1900 zu stellen. Sie passt unge- 
zwungen in keine der bekannten Gattungen, steht jedoch den Gattungen 
Sciopetris Meyrick, 1891 und Kozhantshikovia Saigusa, 1961 nahe. 
Nachdem nur die Männchen bekannt sind und die Säcke, Raupen und 
auch die Weibchen fehlen, will ich davon absehen eine neue Gattung 
zu kreieren und stelle die neue Art provisorisch an den Schluss der 
Gattung Sciopetris, obwohl der neuen Art die Ocellen fehlen und auch 
einige andere Unterschiede vorhanden sind. Die neue Art soll Sciopetris 
karsholti sp. n. heissen, zu Ehren des aktiven Forschers im Gebiet der 
Lepidoptera und Sammlers der neuen Art, Ole Karsholt im Zoologi- 
schen Museum Kopenhagen. 


Sciopetris karsholti sp. n. 


MATERIAL : 25 Männchen am Tag gefangen durch Herrn Ole Karsholt, 
alle von Tunesien, Ain Draham area, 5. - 18.5.1988. Ein Exemplar wird 
als Holotypus bezeichnet, es befindet sich in der Sammlung des Zoo- 
logischen Museums der Universität Kopenhagen. Die anderen 24 Männ- 
chen sind als Paratypus bezeichnet, davon sind 16 ebenfalls in der 


108 


Sammlung des Zoologischen Museums Kopenhagen und 8 Exemplare 
in meiner Sammlung. 


FALTERBESCHREIBUNG : Männchen mit einer Flügel-Spannweite von 
9.5-10 mm. Im Vorderflügel 8 Adern aus der Discoidalzelle, im Hinter- 
fliigel deren 5 und eine eingeschobene Zelle, beide Fliigel schmal, nach 
aussen etwas erweitert, Apex spitz (Abb. 2). Beschuppung der Vorder- 
flügel Breitenklasse 4-5, (Sauter, 1956), hellbraun mit unregelmässigen 
blassbraunen Flecken, die bei einzelnen Tieren wie eine Gitterung 
wirken (Abb. 3). Hinterflügel : Beschuppung der Breitenklasse 2-3, ein- 
farbig hell braungrau. Fransen aller Flügel blass-braungrau. Fühler 
fadenförmig mit etwa 29 Gliedern, ringsum unregelmässig bewimpert. 
Ocellen fehlen, Labialpalpen 3-gliedrig, nach vorn gebogen. Augen 
kugelig, Abstand 1.2-1.5 mal Augenhöhe, Gesicht anliegend, der Kopf 
struppig mit gelbbraunen Haaren bedeckt. Vorderbeine mit kleiner 
Epiphyse an der Tibia, Mittelbeine mit einem Paar Endsporne, die 
Hinterbeine mit je einem Paar Mittel- und Endsporne an der Tibia. 
Das Genital (Abb. 4a) mit kurzem Saccus, dıe Valve überragt das 
Tegumen. Am distalen Ende des Sacculus der Valve sitzt ein Dorn 
(Abb. 4b). 


Der Lebensraum (Abb. 1) wird vom Finder der neuen Art wie folgt 
beschrieben : Der Typenfundort ist in NW Tunesien gelegen, etwa 
35 km südlich der Mittelmeerküste im Hügelgebiet südlich des Dorfes 
Ain Draham auf ca. 900 m ü/M. Das Biotop ist ein offener, trockener 
Eichenwald (Quercus suber und Quercus faginea ssp. baetica (= ©. 
mirbeckii)). Das Klıma ist im Einflussgebiet des Mittelmeeres, regen- 
reiche Winter (oder sogar Schnee !) und trocken im Sommer. 


Alle Falter wurden in der Zeit vom 5. Mai bis zum 18 Mai 1988 am 
Morgen mit dem Netz gefangen. In keinem Fall flogen sie zum Licht, 
jedoch waren einige in der Malaisefalle. Die Männchen flogen frei und 
recht hoch um die Eichen, nicht in Bodennähe. Wie schon erwähnt 


Abb. 2. Aderung der Flügel von S. karsholti sp. n. 


109 


war es trotz sehr gründlichem Suchen an Stämmen und Strünken der 
Eichen aber auch an Steinen nicht möglich die Säcke oder gar Weibchen 
zu finden. Es muss im Moment offen bleiben, wo die Entwicklung 
statt findet, in Bodennähe oder sogar oben in den Eichen. Bis jetzt 
ist die neue Art nur aus diesem Gebiet bekannt. 


Abb. 3. Männchen von S. karsholti sp. n. Die Flecken- oder Gitterzeichnung auf 
den Vorderflügeln ist variabel. (Foto : Museum Kopenhagen). 


Abb. 4. a = Genital des Männchens von S. karsholti sp. n., b = Dorn am distalen 
Ende des Sacculus (vergrössert). Zeichnungen von Prof. Dr. Willi Sauter. 
c = S. hartigi Sieder, 1976, d = vergrösserter Dorn des Sacculus, e = Aedeagus. 


110 


Abb. 5. Männliche Genitalien von Kozhantshikovia vernalis Saigusa, 1961 (nach 
Saigusa, 1961, p 307). a = Ansicht von ventral, b = von lateral. Der Saccus und 


der Aedeagus sind sehr lang. 


Flügel: 
Adern aus Discoidalzelle 
Vorderflügel 


Hinterflügel 
Eingeschobene Zelle 
im Hinterflügel 


Männliches Genital: 
Saccus 


Dorn am distalen 
Ende des Sacculus 


Bedornte Platten auf 
dem Diaphragma 


Aedeagus-Länge 


Ocellen 


S. karsholti spec.nov. 


8 
6 


vorhanden 


kürzer als Genitallange 


vorhanden 


fehlen 


Kürzer als Genitallange 
ohne Saccus 


fehlen 


Gen. Sciopetris 
S. hartigi SIEDER 


fehlt 


fehlt 


vorhanden 


fehlen 


Kürzer als Genitallänge 
ohne Saccus 


vorhanden 


Gen. Kozhantshikovia 
K. vernalis SAIGUSA 


fehlt 
sehr lang, länger als 
Genitalänge 


fehlt (?) 


sehr ausgeprägt 


1.5-2 Genitallängen 
ohne Saccus 


fehlen 


Abb. 6. Vergleich einiger Merkmale von S. karsholti sp. n. mit Merkmalen der Gat- 
tungen Sciopetris und Kozhantshikovia. 


Diskussion 


Die beiden Gattungen Sciopetris und Kozhantshikovia stehen einander 
in verschiedenen Aspeken recht nahe. Die neue Art steht dazwischen, 
sie hat Ubereinstimmungen mit Merkmalen beider Gattungen. Das 
männliche Genital der spec. nov. zeigt gesamthaft und in mehreren 
Details eine brauchbare Übereinstimmung mit Sciopetris, was für 


111 


Kozhantshikovia weit weniger zutrifft (Abb. 4 und 5). Im Vergleich 
mit Sciopetris fehlen der neuen Art die Ocellen und im Hinterflügel 
sind 6 Adern aus der Discoidalzelle mit einer eingeschobenen Zelle 
vorhanden, wogegen Sciopetris 5 Adern ohne eingeschobene Zelle 
aufweist. Die Tabelle Abb. 6 zeigt einige Vergleiche zwischen den beiden 
Gattungen und der neuen Art. Von den anderen Arten in der Gattung 
Sciopetris ist S. karsholti durch das Fehlen der Ocellen und von 
Kozhantshikovia durch den viel kürzeren Saccus leicht zu trennen. 


Verdankungen 


Ich danke den Herren Ole Karsholt (Kopenhagen) für die interessanten Falter- 
chen, die Fotos des Lebensraumes und des Männchens sowie für die zusätz- 
lichen Angaben über Klima, Fundort usw. und Prof. Dr. Willi Sauter für 
die Beratung, die Genitalzeichnung der neuen Art und die Durchsicht des 
Manuskriptes. 


Literatur 


MEYRICK, E., 1891. A fortnight in Algeria, with description of new Lepidoptera. 
Entomologist’s monthly Magazine 27 : 9, 55-62, p. 58. 

SAIGUSA, T., 1961. Systematic studies of Diplodoma and its allied genera 
in Japan. Sieboldia 2 (4) : 262-314. 

SAUTER, W., 1956. Morphologie und Systematik der schweizerischen Sole- 
nobia-Arten. Rev. Suisse Zool. 63 : (3) Nr. 27 : 451-559. 

SIEDER, L., 1959. Neue palaearktische Psychiden. Zeitschr. Wien. Ent. Ges. 
44 : 145-150. 

SIEDER, L., 1976. Eine neue Psychide aus Sardinien, Sciopetris hartigi spec. 
nov. Ent. Ber., Amst. 36, I, III : 43-45. 


112 


Nota lepid. 19 (1/2) : 113-128 ; 21.X1.1996 ISSN 0342-7536 


On the induction of metamorphosis of Lepidoptera 
by means of ecdysone and 20-hydroxyecdysone 


Data on 268 hybrid and non-hybrid Sphingidae and 14 Bombyces 


E. A. LOELIGER* & F. KARRER** 


* Hofdijck 48, NL-2341 Oegstgeest, The Netherlands 
** Rebbergstrasse 5, CH-4800 Zofingen, Switzerland 


Summary 


Metamorphosis and adult eclosion were induced by the injection of appropriate 
amounts of 20-hydroxyecdysone or ecdysone into the pupae of female hybrids 
of Sphingidae (Lepidoptera) which otherwise would have remained in diapause 
until death. Non-hybrid pupae were also treated. Ecdysone produced better 
results than 20-hydroxyecdysone. For hybrids, the optimum dose of hormone 
was assessed at six mg per g live weight; the dose might be considerably 
higher for non-hybrids. The duration of metamorphosis correlated weakly 
with the injected dose ; the correlation with temperature was pronounced. 
Interspecies differences were small. Ecdysone-induced imagines fed and mated 
normally, produced fertile sperm and eggs, and displayed normal longevity. 
Shipment of the injected pupae had a deleterious effect on metamorphosis 
in the majority of cases. 


Résumé 


La métamorphose et l’'éclosion de femelles adultes hybrides de Sphingidae 
(Lepidoptera) ont été causées par l’injection de quantités appropriées de 20- 
hydroxyecdysone ou ecdysone dans leurs chrysalides, sans quoi celles-ci 
seraient restées en diapause jusqu’à leur mort. Des chrysalides non-hybrides 
furent également traitées. L’ecdysone produisit de meilleurs résultats que la 
20-hydroxyecdysone. Pour les hybrides, la dose hormonale optimale était 
fixée à six ug par gramme de poids vivant ; la dose pourrait être considé- 
rablement supérieure pour les non-hybrides. Il y avait une corrélation faible 
entre la durée de la métamorphose et la dose injectée ; la corrélation avec 
la température était élevée. Les différences interspécifiques étaient faibles. Les 
imagos obtenus par l'injection d’ecdysone se nourissaient et s’accouplaient 
normalement, produisant du sperme et des oeufs fertils et montrant une 
longévité normale. L'envoi de chrysalides injectées avait un effet néfaste sur 
le déroulement de la métamorphose dans la majorité des cas. 


Key words : Ecdysone, ecdysteroids, metamorphosis, Lepidoptera, Sphingidae, 
hybrids, shipment 


113 


Introduction 


Female hybrids of certain species of the Sphingidae family enter 
metamorphosis spontaneously only in exceptional cases or not at all 
(FISCHER, 1932). The first successful attempt to break this barrier was 
accomplished by means of transfusion of haemolymph taken from 
normal metamorphosing pupal instars (MEYER, 1953). The transfusion 
procedure was, however, hazardous ; therefore this procedure was never 
developed and applied. It was only after purified hormones became 
available that experimental work on the induction of metamorphosis 
gained momentum (WILLIAMs, 1968 ; OHTAKI & WILLIAMs, 1970). The 
hormones were also injected into female hybrid pupae predestined 
to die in Papilionidae (Clarke and Willig, 1977) and in Bombyces (ADEs 
et al., 1989), but obviously not yet in Sphingidae (NıJHoUT, 1994). 


In 1983, our series of experiments was started. Initially, the results 
varied markedly. In 1989, by pure chance, we switched from 20- 
hydroxyecdysone to ecdysone ; immediately our results were more 
successful. We could now assess the dose/effect relationship, possible 
interspecies differences, and the relationship between the duration of 
metamorphosis and the effect of ambient temperature on pupae. 


In non-hybrid pupae we tried to synchronize the eclosion of males 
and females which, under mid-European climatological conditions, only 
haphazardly enter metamorphosis. 


Materials And Methods 


Experimental animals (Table 1) 


The majority of pupae referred to under Results (Tables 2 and 3) 
were derived from well-known hybrid and non-hybrid species (ADEs 
et al., 1989; DE FREINA, 1991 ; DE FREINA & Witt, 1987 ; DENso, 
1913 ; FISCHER, 1931 ; GEHLEN, 1933; JoHN, 1932; JorDAN, 1913; 
MEERMAN, 1988 ; MEERMAN & Smid, 1988 ; ROTHSCHILD & JORDAN, 
1903). Novel hybrids are indicated under Results as hybr.nov.Loeliger 
(Table 1). 


Most pupae were maintained on dry sand. Only pupae known to be 
susceptible to dessiccation (elpenor, gschwandneri, harmuthi, and 
rydbergi) were kept in an atmosphere of sufficient humidity. After the 
injection of ecdysteroid, the pupae were stored at room temperature 
unless otherwise stated. 


114 


Table 1 
Sphingid hybrids 


Combination of species 
Published (Denso, 1913 ; Jordan, 1913 ; John, 1932 ; Gehlen, 1933 ; De Freina, 1991) 


x H. vespertilio epilobii 
H. euphorbiae x P. elpenor harmuthi 
H. gallii x H. hippophaes fromkei 

x 

x 

x 


Ö 
H. euphorbiae 


H. gallii H. vespertilio carolae 

H. gallii P. elpenor gschwandneri 
H. hippophaes H. euphorbia hippophorbiae 
H. hippophaes x P. elpenor rydbergi 


Unpublished (Hybr. nov. Loeliger) 


2 
H. dahlii P. elpenor no eponym 
H. gallii H. tithymali himyarensis | gallyarensis 
H. nicaea H. euphorbiae conspicua | paranicaea conspicua 
H. nicaea H. euphorbiae cretica Paranicaea cretica 
H. nicaea H. euphorbiae gecki paranicaea gecki 
H. nicaea H. vespertilio nicertilio 


Table 1 : Synopsis of eponyms for the experimental hybrid female pupae, which usually 
remain in permanent diapause until they die. 


Other materials 


Injection needles: Microlance™, Becton, Dickinson & Company ; 
30 g% 0.3 X 13 bl Ib. 
Syringes : Jecton-S™, | ml, subdivided into 0.02 ml. 
Water and physiological saline : pyrogen-free and sterile. 
Alcohol : aethylalcohol (ethanol) and isopropylalcohol (isopropanol), 
> 99%. 
Collodium : 3%. 
Iodine : 1%, in ethanol 65%. 
Ecdysteroids : Ecdysone from SIGMA, St. Louis, USA; FLUKA, 
Buchs, CH ; 
ROHTO, Osaka, J. Abbreviations used : SIGMA = S, 
FLUKA = F, and ROHTO = R. 
20-hydroxyecdysone from SIGMA and from ROHTO. 


Injection fluid: not until 1992 was alcohol used in a concentration 
= 10%, sufficient to hold ecdysone in solution for the time needed 
for the injection (see legends to Tables 2 and 3). 


115 


Species 

year of breeding/ 
injection 

Hybr. rydbergi 
1982/1983 


Hybr. rydbergi 
1982/1983 


H. dahlii & x 
D. elpenor ® 
1982/1984 


Hybr. rydbergi 
1983/1984 


Hybr. fromkei 
| 1983/1984 


Hybr. fromkei 
1983 / 1984 


Hybr. hippophor- 
biae 1983/1984 


Hybr. carolae 
1984/ 1985 


Hybr. fromkei 
1983/1985 


Hybr. hippophor- 
biae 1983/1985 


Hybr. epilobii 
1983/1985 


Hybr. fromkei 
1983/1985 


Hybr. carolae 
1984/1985 


Hybr. fromkei 
1986/1987 


H. centralasiae 
siehei 1986/1987 


116 


Table 2 


Experiments with 20-hydroxyecdysone 


3 
3 


BE Hi di: 
Nn 
© 


shipped 


3 shipped 


shipped 


4 non-shipped 
18 shipped 


4 non-shipped 
4 shipped 


2 non-shipped 
8 shipped 
shipped 
shipped 
non-shipped 

1 non-shipped 
4 shipped 


5 non-shipped 
4 shipped 


2 —;3 — 
| FRERE 


Present procedure for preparation of the injection fluid 


To obtain a 0.1% hormone solution in 12% alcohol, first 0.12 ml alcohol 
are mixed with | mg of the hormone crystals which rapidly dissolve. 
To this mixture 0.08 ml water is added in order to enhance the sterilizing 
capacity of alcohol (65% is optimal). Ten minutes later 0.8 ml water 
are added resulting in the final concentrations. 


Injection technique 


The apex of the head of the pupa is iodinized about 10 minutes before 
puncture. Anaesthesia is achieved by exposing the pupa for 20-40 
minutes to CO, evaporating from dry ice. To protect the pupa against 
cold, it is placed on a 2-3 cm layer of linen covering the ice. 


The anaesthetized pupa is held in the horizontal position while the 
needle is inserted through the iodinized apex of the head deep through 
the thorax into the upper abdomen, along the sagittal axis. The injection 
is slow. The needle remains in situ for about 10 seconds, to allow 
diffusion of the injected fluid. After removal of the needle, the puncture 
site is immediately covered with collodium. Twenty-four hours later, 
the pupa can be handled normally. 


Results 


Tables 2 and 3 represent the results in chronological order. The synopsis 
given in Table 4 is an attempt to identify preconditions for success- 
ful eclosion of adults. It appears that during the first six years (20- 
hydroxyecdysone experiments ; Table 2), a much smaller number of 
pupae was treated under optimum conditions than during the next 
three years (the ecdysone experiments ; Table 3). Therefore, the eclosion 


Table 2: Chronological assessment of the results of 20-hydroxyecdysone injections, 
from 1983 to 1989. The injection fluid was consistently filtered. The composition varied 
considerably : 1) 0.1% beta-ecdysone“S” in 2.5% ethanol and 0.81% NaCl ; 2) 0.01%*R” 
further idem ; 3) 0.05%“S” further idem ; 4) same solution after storage at -20°C for 
3 months ; 5) 0.04%“S” in 4% ethanol and 0.54% NaCl; 6) same solution as under 
3) after storage at -20°C for 4 years ; 7) same solution as under 5), after storage at 
-20°C for 26 months ; 8) 0.04%“S” in 4% ethanol. 
Under “Results” the quality of metamorphosis is indicated by : + = perfect eclosion ; 
+ = almost perfect eclosion ; + = crippled imago ; - = signs of metamorphosis but 
no eclosion ; —- = no signs ‘of metamorphosis. From hybr. hippophorbiae treated 
in 1985 through H. centralasiae siehei treated in 1987, the injection site was sealed 
with cyanoacryl glue. 


11307) 


Species 

year of breeding/ 
injection 

Hybr. fromkei 
1988 / 1989 


Hybr. fromkei 
1988 / 1989 


Hybr. paranicaea 
cretica, 1988/89 


Hybr. paranicaea 
conspicua, 
1988/89 

idem, inj. 1990 


Hybr. fromkei 
1988/1990 


G. isabellae & x 


A. luna 9, 1989/90 


Hybr. fromkei 
1988 / 1990 


Hybr. fromkei x 
fromkei, 1989/90 


H. centralasiae 
siehei 1986/1991 


Hybr. fromkei 
1990/91 


E. imperialis & x 
E. magnifica ®, 
1989/91 


Hybr. gschwand- 
neri 1991 


Hybr. gallyarensis 
1991/1992 


118 


Table 3 


Experiments with ecdyson 


ug/g Handling 
ber of | live 
pupae | weight 


3 non-shipped 
3 shipped 


1 non-shipped 
1 shipped 


shipped 
shipped 


shipped 


2 non-shipped 
2 shipped 
2 non-shipped 
2 shipped 
2 non-shipped 
2 shipped 


Result 


2++,1--,9 


i++;1+ 


3 ++ ; ovaria unripe 
Saar 


——, reinj. 1990 


H+++ 
~~ 


all 4 ++, ») 


no eclosion 
remained alive 


——, remained alive 


Species 

year of breeding/ 
injection 

Hybr. nicertilio 
1991 / 1992 


Hybr. gschwand- 
neri 1991/92 


Hybr. harmuthi 
1991 / 1992 


Hybr. nicertilio 
1991/1992 


Hybr. gallyarensis 
1991 / 1992 


Hybr. fromkei 
1991041992 


Hybr. paranicaea 
Gecki, 1991 / 1992 


B. europaea, 
1990 / 1992 


Hybr. gschwand- 
neri 1991/92 


Hybr. gallyarensis 
1991/1992 

Hybr. rydbergi 
1992 


Hybr. nicertilio 
1991/1992 


Hybr. rydbergi & 
1992 


Table 3 (continued) 


Experiments with ecdyson 


ug/g Handling Result 
ber of | live 
pupae | weight 


2 non-shipped 
2 shipped 
2 non-shipped 
1 shipped 
2 non-shipped 
2 shipped 


7 non-shipped 
10 shipped 


2 non-shipped 
3 shipped 
2 non-shipped 
4 shipped 


oS ol eats 
3 non-shipped | 3 ++ 
1 shipped - 


shipped no eclosions, 
survived 


7 non-shipped | 2 ++ ;2+;3- 
3 shipped Rea = 


Boe 


shipped 


109 


Table 3 (continued) 


Experiments with ecdyson 


Species | Injec-| Num- | ug/g 
year of breeding/ | tion | ber of | live 
injection fluid | pupae | weight 


H. euphorbiae 
mauretanica ®, 
1992 


Handling Result 


H. euphorbiae 1 + 


conspicua 66, 
1992 


D. elpenor &, 2 RE 


1992 


2++; 4—-, surviving, 


D. elpenor 3 64 
4 eclosing 1993 


and 3 9Q, 1992 


--, surviving and 


P. proserpina 
eclosing 1993 


3 66 and 3 99, 
1992 


Table 3 : Chronological assessment of the results of the ecdysone injections, from 1989 
through 1993. After treatment of hybr. paranicaea cretica, the injection fluid was no 
longer filtered ; the composition varied : 1) 0.04%“S” in 4% ethanol ; 2) same solution 
after storage at -20°C for 6 weeks; overt turbidity ; 3) 0.05%“F” in 9% ethanol ; 
4) same, after storage at -20°C for 1 year ; overt turbidity ; 5) addition through same 
needle of 4 mg/g live weight beta-ecdysone “S” which had been stored at -20°C for 
1 year, displaying no turbidity ; 6) as under 3), but freshly prepared ; 7) 0,035%“R” 
in 9% ethanol ; 8) 0.05“F” in 5% ethanol, displaying slight turbidity, further diluted 
with water ; 9) 0.1“F” in 13% ethanol; 10) same solution after storage at -20°C for 
8 days ; overt turbidity. 

Flawlessly eclosed imagines behaved normally: a) one specimen mated with a spon- 
taneously eclosed male hybr. fromkei, began oviposition 5 days later (5 eggs), died 
4 weeks after eclosion, with about 180 ripe eggs in abdomine ; b) one specimen mated 
with H. gallii and started oviposition on E. angustifolium 5 days later ; of the 82 eggs 
76 were fertile ; c) one mated with D. elpenor, but died 8 days later without having 
laid eggs ; it contained about 50 ripe eggs ; the other specimen began oviposition 30 days 
after eclosion (49 sterile eggs); it lived for 67 days; d) this pupa, injected 4 days 
after ecdysis and kept at 35°C, eclosed 15 days later, containing about 100 ripe eggs ; 
e) mated with a spontaneously eclosed female which produced 383 fertile eggs. 
Qualification of metamorphosis as for Table 2. 


rates calculated for the two periods should be assessed carefully. How- 
ever, the higher eclosion rates for ecdysone are overly clear. Irrefutable 
is the deleterious effect of shipment on injected pupae, particularly on 
those of hybrid nicertilio : only one of the 14 shipped pupae underwent 
eclosion and produced a more or less flawless imago. 


120 


Sealing the puncture with cyanoacryl glue turned out to be toxic ; 
the puncture sites sometimes turned black, especially among Hyles 
centralasiae siehei pupae injected in 1987. This sealing procedure was 
abandoned after 33 pupae had produced only one acceptable imago. 


Figure | illustrates the interrelationship between the duration of meta- 
morphosis and the temperatures to which the injected pupae were 
exposed. At temperatures of 30°C to 35°C less than a fortnight was 
needed for procreation ; in contrast, at temperatures < 20°C meta- 
morphosis took more than five weeks. The data presented in Figure 2 
demonstrate the species-specificity of the interrelationship, hybrid 
nicertilio requiring about ten percent more time than hybrid gallyarensis 
to complete the process of metamorphosis. Moreover, the higher the 
dose, the faster the process. 


Table 4 


Eclosion rates after ecdysteroid injections 


Pupae Flawless eclosions 
injected 
number | Number | percentage 


NS ET ff 


optimal conditions (sphingidae hybrids only) 

< 3 mg/g live weight (suitable hybrid pupae only) 
shipped pupae (suitable hybrid pupae only) 
unsuitable conditions 


optimal conditions sphingidae-hybrids 
non-hybrids 

< 3 mg/g live weight (suitable hybrid pupae only) 

shipped pupae (suitable hybrid pupae only) 

unsuitable conditions 


1) one pupa is counted a second time under “unsuitable conditions” 


Table 4 : Grouping of the results according to the type and dosage of the ecdysteroid, 
the quality of the pupae and treatment procedure. “Optimal conditions” indicate a 
good pupa, a sufficient amount of hormone injected, a flawless injection technique, 
and no shipment. “Unsuitable conditions” mean a deep-frozen ecdysone solution, 
imperfect pupa, pupa older than one year, insufficient dosage, substantial loss of 
haemolymph after the injection, and shipment. 


121 


LO 


30 


fe) 
a 
i 
© 
+ 
® 
a 
e 


TEMPERATURE PUPAE ARE EXPOSED TO (°C) 


158) 
oO 


20 30 40 


DURATION OF METAMORPHOSIS (DAYS) 


Fig. 1. Relationship between the duration of metamorphosis and the temperature of 
storage of the injected pupae. The season-dependent temperatures indicated are rough 
assessments. Temperature was not controlled except for eight of the hybrid fromkei 
specimens injected in 1990 (26°C), and the hybrid rydbergi specimen injected in 1992 
EI) 


Doubling the amount appears to shorten the period, also by about 
ten percent. 


When we switched from 20-hydroxyecdysone to ecdysone (June 1, 1989) 
at a dose of four mg per gram live weight, the duration of the process 
of metamorphosis was found to differ between the two hormones: 
among hybrid fromkei pupae kept at room temperature, the appearance, 
on translucence, of the eyes occurred three days later in those treated 
with ecdysone than in those treated with 20-hydroxyecdysone under 
the same conditions. The delays in the appearance of the wing pattern 


122 


AMOUNT OF ECDYSONE INJECTED ( #9/g LIVE WEIGHT) 


20 25 30 


DURATION OF METAMORPHOSIS (DAYS) 


Fig. 2. Relationship between the amount of ecdysone injected and the duration of 
metamorphosis. The mean duration for hybrid nicertilio injected on July 4, 1992 (©) 
and July 22nd, 1992 (e), was 26.2 days, whereas that for hybrid gallyarensis injected 
on July 4 (©) and July 22™4 (&) was 23.1 days, approximating a difference of ten 
per cent. The broken lines roughly indicate the relationship between the dose applied 
and the duration of metamorphosis. 


and the eclosion of the imago were similar. In contrast, there was no 
difference in the duration of metamorphosis of the two hybrid para- 
nicaea conspicua specimens, one of which was treated with a com- 
bination of the two ecdysteroids (see Table 3). 


Discussion 


When we started this series of experiments in 1983, nothing was 
known about the induction by the injection of ecdysteroids of the 


123 


metamorphosis of females of certain hybrid species of the family 
Sphingidae, which do not eclose spontaneously or only in exceptional 
cases. By trial and error, the goal of our endeavor to terminate diapause 
in these females as well as in normal pupae was achieved. 


In the beginning, special attention was directed toward the prevention 
of infection of the pupae, isotonicity, and non-toxicity of the injection 
fluid. The injection site was iodinized about ten minutes before the 
injection. Filtration of the injection fluid to free it from possible patho- 
genic microorganisms, a routine procedure in the early stages of our 
experiments, was abandoned after the switch from 20-hydroxyecdysone 
to ecdysone. Instead, ecdysone was exposed to diluted alcohol, 65 per 
cent in water, to enhance its sterilizing effect. With these precautions, 
microbial infections were never observed. 


Isotonicity of the fluid seemed to be of little importance, since dis- 
continuation of the addition of physiological saline in 1985 did not 
cause a change in the results. 


The apex of the head of the pupa as the preferred injection site in 
Sphingidae pupae was adopted from experienced investigators (MEYER, 
1953). It obviously does not cause prohibitive damage of the brain 
nor the prothoracic region. Abdominal injection sites may give raise 
to untoward blood loss once pupae start to move again after awakening 
from narcosis. 


Signs of alcohol toxicity were not apparent. The ethanol or isopropanol 
concentration was increased step by step, up to the 12 per cent needed 
to keep ecdysone in solution long enough to complete successful in- 
jection of a series of pupae. In storage, at low temperatures in particular, 
flocculation of ecdysone takes place if the solution contains one mg 
per ml ; after flocculation, ecdysone seems to loose much of its activity. 


With this ecdysone solution — one per mille in 12 per cent alcohol/ 
water — the volume injected remains below one per cent of the pupal 
volume and the intrapupal pressure barely increases, so that the 
procedure does not induce undesirable loss of haemolymph as long 
as anaesthesia is effective and the injection site is sealed. 


Results with 20-hydroxyecdysone were unsatisfactory in spite of a 
meticulous injection procedure. For non-shipped pupae treated with 
sufficient amounts of hormone, the success rate remained below 50 per 
cent. Three to five mg per g live weight did not always induce complete 
metamorphosis. Six mg per g might be the optimum amount. Our 
results obtained with 20-hydroxyecdysone correspond closely to those 


124 


reported earlier for other hybrids: in 34 female pupae of a Bombyx 
species, the eclosion rate obtained at a dosage averaging 3.9 mg per 
g live weight was almost 50 per cent, the percentage of flawless imagines 
being approximately 15 per cent (ADEs et al., 1989). 


It was by pure chance that ecdysone was introduced in our series of 
experiments. The first application, in female hybrid fromkei pupae, 
in June 1989, resulted in a complete success : one of the imagines mated 
normally, started oviposition soon afterwards, and displayed normal 
longevity. The vitality of this specimen was sufficient reason to continue 
our study with only ecdysone. 


The minimum effective dose of ecdysone was assessed at three mg per g 
live weight, = 4 mg per g never failing to induce the process of meta- 
morphosis. Six to 12 mg per g regularly produced flawless adults. In 
one instance (hybrid paranicaea conspicua, 1990) a dose as high as 
18 mg per g was just as successful, indicating a wide range from the 
optimum dosage of ecdysone for hybrids, and confirming results re- 
ported for non-hybrids (WiLLIAMs, 1968 ; OHTAKI & WırLıams, 1970 ; 
SLAMA et al., 1973). it is interesting that the addition of 20-hydroxy- 
ecdysone (four mg per g live weight) to ecdysone (18 mg per g) did 
not appreciably accelerate the process of metamorphosis: the two 
hybrid paranicaea conspicua specimens, one treated with the combi- 
nation and the other with ten mg per g only, eclosed simultaneously. 


We suggest that six mg ecdysone per g live weight will be sufficient 
for induction of the metamorphosis of female hybrids of Sphingidae 
remaining in diapause. 


Using this dosage, we recently were successful in breeding an F, of the 
hybrid vespertilioides Boisduval (LOELIGER & KARRER, in press). 


In hybrids of Papilionidae, the optimum dose is probably much lower 
(CLARKE & WILLIG, 1977). For non-hybrids, the optimum dosage has 
not yet been assessed. Depending on the stage of diapause, it might 
be considerably higher, although procedural failures and/or flocculation 
of the ecdysone solution might have been the reason for non-induction 
of metamorphosis in D. elpenor and P proserpina in 1992. 


From observations of hybrid gallyarensis, nicertilio, and rydbergi, we 
conclude that the process of metamorphosis is about halfway when 
the eyes become visible on translucency ; moreover, the period between 
the appearance of the wing pattern and eclosion equals 15 to 20 per cent 
of the total duration of metamorphosis. These assessments are in good 
agreement with observations of Samia cynthia (WILLIAMS, 1968). 


125 


It is quite clear that the process of metamorphosis is highly dependent 
on the ambient temperature for the injected pupae (Figure 1). Our 
results agree with those reported for spontaneously developing hybrids 
(HARBICH, 1982) and non-hybrids (HARBICH, 1976 and 1977). Figure 2 
demonstrates that the process is also dose-dependent, though to a much 
lesser degree. Moreover, the overall values presented indicate a species 
specificity, the difference amounting to ten to 15 per cent of the total 
duration. 


After the injection of ecdysone it takes about ten per cent more time 
to obtain the adult than after injection of 20-hydroxyecdysone. A 
similar difference was observed when hybrid fromkei pupae received 
the two hormones simultaneously in 1989. It reflects the time needed 
for the biotransformation of the inactive ecdysone into the active 20- 
hydroxyecdysone well-known in slow release reactions in endocrinology. 


Our investigations of the feeding and mating behaviors of hormone 
procreated imagines is limited to only a few specimens receiving 
ecdysone (see Legend to Table 3). Feeding was not always easy because 
the two parts of the proboscis were insufficiently adjusted to each other 
or did not curl, mainly in hybrid nicertilio. But females of hybrid 
gschwandneri, gallyarensis, and vespertilioides F, (LOELIGER & KARRER, 
in press) could be fed and remained in good health for up to two 
months. The ovaries of gschwandneri were atrophic, whereas gallyarensis 
produced eggs from the 30 day after eclosion on. Another gallyarensis 
mated within 24 hours after eclosion with D. elpenor, but died nine 
days later without having deposited any of the eggs it contained. Hybrid 
fromkei, which eclosed in June 1990, mated with A. gallii and remained 
alive for three weeks. The offspring of this couple were flawless adults, 
very much resembling gallü. Fertility of the F, of the hybrid vesperti- 
lioides was surprisingly high (LOELIGER & KARRER, in press). 


Among the injected non-hybrids, the H. euphorbiae conspicua male 
adult, eclosing in September 1992, mated with a spontaneously eclosed 
female of the same species that produced more than 350 eggs, the 
vast majority of which were fertile. Finally, four female H. euphorbiae 
conspicua specimens, eclosing in late December 1993, appeared on 
section to contain numerous ripe eggs. 


The low eclosion rate observed for shipped pupae was disappointing. 
Only one of the 14 shipped hybrid nicertilio displayed satisfactory 
eclosion, compared to ten of the 13 non-shipped specimens. For hybrid 
gallyarensis, the deleterious effect of shipping was somewhat less 


126 


impressive, the respective figures amouting to 4/11 and 9/14. Considered 
together, successful eclosion of 70 per cent of the non-shipped pupae 
occurred, compared to only 20 per cent of those shipped. Bumping 
and rocking during shipment must account for the disturbance of meta- 
morphosis. 


We may conclude with the assumption that ecdysone, due to its 
continuous slow release biotransformation — the biological t% as 
assessed in Samia cynthia was approximately six hours (OHTAKI & 
WILLIAMS, 1970) — produces sufficient amounts of physiologically 
active 20-hydroxyecdysone to induce metamorphosis if doses of six 
mg or more are injected per g live weight of a dormant pupa. 


Acknowledgements 


We are indebted to all those who contributed healthy livestock material and/ 
or carefully reported results about shipped pupae : 


A. Baumgartner, Säckingen (D), H. Czipka, Fürth-Erlenbach (D), H. Harbich, 
Salz (D), F. Hohl, Soultz (F), A. Pelzer, Wennigsen (D), F. Renner, Erbach- 
Ringingen (D), B. Surholt, Dülmen (D), and F. Weber, Riehen (CH). 


Dry ice was kindly supplied by the immunochemistry laboratory of the depart- 
ment of nephrology, director M.R. Daha, of the University Hospital, Leiden. 


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LOELIGER, E. A. & KARRER, F., in press. Breeding of an F, of Hyles hybr. 
vespertilioides (BOISDUVAL, 1827) after induction with ecdysone of 
metamorphosis in diapausal pupae. Nota lepid. 19. 

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128 


Nota lepid. 19 (1/2) : 129-140 ; 21.X1.1996 ISSN 0342-7536 


The migration of Danaus plexippus (Linnaeus, 1758) 
during October 1995 in the UK 
(Lepidoptera : Nymphalidae, Danainae) 


Bart VANHOLDER 


Droeskouter 33, B-9450 Haaltert, Belgium. e-mail : bvholder@innet.be 


Summary 


Sightings of Danaus plexippus (Linnaeus, 1758) are reported from the UK, 
Ireland, Holland, Belgium and France in October 1995. We strongly suggest 
that these migrated across the Atlantic Ocean, just as most likely has been the 
case with past migrations. Evidence for this theory is supported by coinciding 
reports of another North American butterfly in England, Vanessa virginiensis 
(Drury, [1773]) and many North American birds. The meteorological conditions 
during September and October 1995 don’t exclude transatlantic migration. 


Résumé 


Des observations de Danaus plexippus (Linnaeus, 1758) sont rapportées en 
provenance du Royaume-Uni, d’Irlande, des Pays-Bas, de Belgique et de France 
en octobre 1995. II est suggéré, arguments a l’appui, que ces exemplaires sont 
des migrateurs originaires de l’autre côté de l’océan Atlantique, comme il en 
a été plus que probablement pour les migrations antérieures. Cette théorie 
est corroborée par les mentions simultanées d’un autre papillon nord-américain 
en Angleterre, Vanessa virginiensis (Drury, [1773]) et de nombreux oiseaux 
nord-americains. Les conditions météorologiques pendant les mois de septembre 
et d'octobre 1995 n’excluent pas une migration transatlantique. 


Introduction 


Danaus plexippus (Linnaeus, 1758), the Monarch butterfly, has since 
long been suspected to cross the Atlantic Ocean. Most likely trans- 
atlantic migration was the way it established colonies on the Azores, 
Madeira and the Canary Islands. From the UK about 450 records 
of Danaus plexippus were known between the first British observation 
on 6th September 1876 (Walker, 1914) and 1988 (Bretherton, 1989). 
Some large and sharply defined immigrations during September and 


129 


October were observed in the past : an invasion of 38 butterflies in 
1933 was described by Williams et al. (1942). In 1968 the main invasion 
started on 30th October with 63 reported butterflies, in 1981 the 
invasion started on 24th September and 135 butterflies were reported 
(Bretherton & Chalmers-Hunt, 1982). In 1983 still a dozen specimens 
were reported (Bretherton, 1984). The largest immigration ever noticed 
in the UK was between 29th September and 22nd October 1995 ; more 
than 150 butterflies were reported. They were mainly observed at the 
south coast, just as was the case with previous immigrations (Bretherton, 
1989). Again, coinciding reports of many American birds during the 
same period strongly suggest an American origin. Contacts via Internet 
with entomological, ornithological and meteorological scientists have 
lead to an acceptable hypothesis about this migration. 


UK-sightings of Monarchs in 1995 


The first sighting of a Monarch butterfly in England in 1995 was on 
6th August with further singletons on 10th and 23rd August. A 
probable explanation for these singletons in August could be found 
in the escape of about 30 Monarchs during July from the greenhouse 
of Dr. M. Rothschild in Ashton, Northants (P. Batty, pers. comm.). 
Indeed, the first sighting was at Winchester (Hants), about 120 km 
from Ashton. On 29 th September a Monarch was reported at Liver- 
pool, about 50 km from Ashton. The presumed early October im- 
migration however, large and sharply defined, cannot be explained by 
these escapees. Almost all observations were done at the south coast 
of the UK, in the counties of Cornwall, Devon and Dorset. 


A map with all sightings of Monarchs during this period in the UK 
was compiled together with Simon Coombes (pers.e-mail) (Fig. 1). If 
we superpose the local weather information we can give a fairly good 
explanation for the histogram (Fig. 4). 


“From 28th September to Ist October strong westerly winds blew, 
having culminated in a west-southwest gale on 30th Sept. The westerly 
winds were followed from 2nd to 5th October by four days of south- 
westerly winds, after which continuous southerly winds blew until 
14th October. Unsettled weather, with up to 95% cloud cover persisted 
from 24th September to 3rd October. This almost certainly prevented 
recording and reduced flight-time of the Monarchs. On 4th and 
5th October 75% cloud cover was reported followed by two days of 
total cloud cover, immediately followed by Sunday 8th October, with 
100% sunshine and 20°C. This day was also ‘International Birdwatch 


130 


Danaus plexippus 


(Linnaeus, 1758) 


July - 5 October 


al D 


N 


29 September 
a ) Ce + + TN 
e TA 30 Monarchs escape : July, vd 
FA re tl S N 6A = 


ca fi ] J > re 
Sy e RS ' 
FERRÉ 6 August; Re À 
ara ci) Se. [À 


1 € rae 
ar ef NaS sq OK 


Key: 
e 1-2 records LEER 


+ probable escapees 


Danaus plexippus 


(Linnaeus, 1758) 


6 - 14 October 


Wind: South-east 


Key: 
@ 1-2 records 


® 3-9 records 


® 10+ records 


Fig. 2 


Fig. 1. Sightings of Danaus plexippus (Linnaeus, 1758) in the UK (July - 5!" October 


1995). 


Fig. 2. Sightings of Danaus plexippus (Linnaeus, 1758) in the UK, Ireland, Holland 


and France (6! - 14th October 1995). 


Day’ and “many birders were out and about to record the Monarchs” 
(R. Lane, pers. comm.). 


As discussed later we presume that the main arrival of the Monarchs 
in the UK should have been around 3rd October. During the following 
days 14 Monarchs were reported until 7th October despite the very 
poor Weather conditions. This proves that they were already present 
before 8th October, the day on which most of the Monarchs were 
seen (Fig. 2). | 


The numbers reported immediately declined and 7 days later, on the 
14th, most of them seemed to have disappeared ; many ‘Monarch- 
seekers’ were successful during the morning of the 14th and reported 
Monarchs being restless and flying strongly until midday. But many 
were frustrated after midday, with no afternoon sightings anywhere 
in Britain. Also the next day, 15th October, was a bright sunny Sunday 
but again almost no sightings anymore. The only positive evidence 
of change was a 90 degree shift in wind direction from south-east on 
14th October to south-west on 15th October. This was the first change 
back to south-westerlies since 5th October. It seems that they dis- 
persed in different directions. Remarkably coinciding with the disappea- 
ring of the butterflies at the south coast of England, one was reported 
in Ireland on 12th October from Belfast and one in Co. Clare and 
the only one from Cockermouth in Cumbria, UK on 13th October. 
In Holland at Oostvoorne in the Meuse valley on 14th October a 
singleton was reported as well as a record from Downpatrick, Co. 
Down in Ireland. It was the first Monarch-sighting since 1969 in 
Holland. On 15th October the last report from Ireland was in co. Down 
and from the same day was the only report from Belgium near the 
coast at Raversijde, Oostende (province West-Flanders) (Vanholder, 
1996). It was first seen by R. Dejaegher through a binocular, flying 
high but afterwards coming down out of the sea. After 14th October 
only 12 specimens were reported from the UK at the south coast. The 
Monarchs in Holland and Belgium probably ended up there with the 
westerly winds blowing until 14th October. In France (Dept. Vendée 
and Dept. Finistère) only 4 sightings were reported and 3 of them 
were seen after 20th October (Dubois, 1996). Although this sample 
is rather small, it suggests that some of the Monarchs might have been 
migrating south, exactly like they should have done in the USA on 
their southbound migration. The last sighting was a butterfly at Les 
Sables d’Olonne (Dept. Vendée) seen flying strongly southwards on 
29th October (Figs. 3 & 4). 


132 


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France 
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25 Ireland 
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Fig. 4 


Fig. 3. Sightings of Danaus plexippus (Linnaeus, 1758) in the UK, Belgium, Holland, 
Ireland and France (15th - 29t October 1995). 

Fig. 4. Histogram of sightings of Danaus plexippus (Linnaeus, 1758) in the autumn 
of 1995. Histogram compiled from 162 records. 


Exactly analogous with the previous migrations in 1968 and 1981 to 
the UK, very few were seen after about ten days of arrival. One female, 
caught in Darwish, Devon, laid eggs in November 1995. The cater- 
pillars were reared successfully by Nigel South in his greenhouse. It 
is a pity that the female was not kept as a voucher specimen. By so- 
called cardenolide-fingerprinting the origin of the specimen could 
have been detected. Only an abdomen, fore- and hind-leg would have 
been sufficient. Cardenolides are the cardiac glycosids present in the 
adult Monarchs and these are responsible for their unpalatability to 
birds. These cardenolides are concentrated in the caterpillars by feeding 
on Asclepias sp. and depending on the foodplant used, a different carde- 
nolide fingerprint is found. It is possible to predict accurately the 
foodplant used by the caterpillar in this way (Brower, 1984 ; 1995). 
This knowledge could have given an indication about the origin of 
the Monarch : e.g. if it had been an Asclepias syriaca-type — the main 
foodplant used by the southbound migrant Monarchs in the USA 
(Brower, 1995) — or an Asclepias curassavica-type, which is the food- 
plant on the Canary Islands. 


The fact of capturing a fertile female at Darwish cannot give much 
additional information about its origin. In Minnesota, USA the first 
non-reproductive Monarchs occurred from 15th August. In the more 
southern states like Kansas even on 18th September 1995 Monarchs 
were reported to be reproductive, because mating pairs were reported. 
In the south, however, the situation is less clear, normally the front 
of the remigration reaches Kansas around 10th September. In the 
extreme South, like Texas, eggs were still found on 15th November. 
It seems safe to conclude that in the northern and middle states 
Monarchs seen after 10th September are usually non-reproductive. 
Nevertheless, fertility cannot be an indication for the origin of the 
female. Migrant females need only 6 days for the conversion of the 
non-reproductive stage into initiation of egg-laying and this can be 
induced by increasing the photoperiod to 14 hours and raising 
temperatures to 25 °C (C. Taylor, pers. e-mail). This is most likely 
what happened in Nigel Souths’ greenhouse, thus explaining why the 
female laid eggs during the first days of November. 


Hypothesis about the way Monarchs travelled across the Atlantic 

In the past wind-assisted transatlantic migration was suggested many 
times. Felt (1928) indicated Danaus plexippus to be a prime candidate 
for crossing the Atlantic on its own power. In Britain most of all past 


observations were done at the south Coast. Lempke (1972) interpreted 


134 


sightings at the coast as an indication that the Monarchs travelled across 
the Atlantic by ship. Probably ship-assistance occurs from time to time, 
when singletons are reported in Europe. But this would mean that 
in the case of a large migration like the present one, high numbers should 
have been travelling together on the same ship. Indeed, to have reports 
of at least 150 Monarchs in the UK, several thousands should have been 
on the wing. On some occasions (Brewer, 1967 ; Varey in Urquhart, 
1987) swarms of Danaus plexippus were seen flying along ships and 
sometimes landing on ships in the Atlantic, suggesting migration. The 
colonization of the Bermuda Islands more than 1000 miles offshore 
proves that Monarchs fly across the Atlantic. Observations of butterflies 
flying over the Atlantic were in many cases reported during hurricanes 
or stormy weather conditions (Scudder, 1876 ; Varey in Urquhart, 1987 ; 
Brewer, 1967). 


For this reason one hypothesis suggested that Monarchs in 1995 could 
have been ejected in the lower jet-stream by September tropical 
hurricanes in the USA and in this way have crossed the Atlantic Ocean. 
During September 1995 two tropical storms, Luis and Marilyn, were 
tracked on the USA east coasts, the supposed area of origin of the 
migrating Monarchs. Luis was tracked from Ist Sept. to 14th Sept. 
when a breakaway low from its remnants was located 300 nautical | 
miles from W. Belfast (Ireland). Mike Campbell (pers. e-mail) analysed 
the jet to be between 200 and 250 mb (millibar) with average tem- 
peratures of -44°C. These temperatures are almost 100% lethal for 
the butterflies. The jet was characterized by a velocity of 125 kts. This 
means the distance from the American East Coast to Ireland would 
have been travelled in 24 hours. These data don’t seem to fit with 
our sightings in the UK. A theoretical possibility for Monarchs to travel 
across the Atlantic with hurricane Luis could only have been as follows : 
an instability existed for about 200 nautical miles radius from Luis’s 
center. Monarchs could not have been incorporated into the flow of 
Luis and ejected in the jet stream unless there were low level jet streams 
or thunderstorms near the United States’ coast wich could have ingested 
the Monarchs and fed them to Luis beyond its 200 nm radius. A second 
tropical September storm, Marilyn, was tracked from 21st September 
to 26th September. The dates seem to fit with the sightings in the UK 
but... Marilyn did never come close to any land. 


A reasonable explanation however for the migration can be found in 
the existence of a very unusual zonal flow. On 29th September there 
was a low pressure centre over St. Anthony, Newfoundland (Canada). 
It intensified to a rather deep 961 mb low by 30th September and 


185 


deepened further to 951 mb to lie about 62 degrees North 33 degrees 
West. At the same time a high pressure centre of 1033 mb was located 
200 nautical miles South of Cape Race. The high was moving along 
a ridge wich extended from Nova Scotia to the Azores. On this day 
westerly surfaces with a velocity of 30-35 kts were reported by stations 
in Newfoundland and ships across the Atlantic to Ireland. Such a wind 
pattern is considered a zonal flow, a condition whereby the winds are 
nearly parallel to the lines of latitude. Winds generally follow isobars 
and in this zonal flow around Ist October the isobars were stretching 
from North America to Europe. These lows of such depth are con- 
sidered an abnormal situation at that time of the year (they are more 
common during January through March). Also the position of the 
ridge of high pressure was unusual. This made the zonal flow across 
the Atlantic indeed very unusual. On 2nd October the zonal flow was 
breaking down. In the zonal flow the air temperatures would have 
been much warmer and hospitable to the Monarchs. Indeed the tem- 
peratures between 1500 and 600m level were between 10 and 15 degrees 
Celsius. As the air moved across the ocean, it was being warmed by 
the Gulf Stream. This caused the airmass to become increasingly stable, 
hence non-turbulent. This layer was likely to be warmer by several 
degrees at the time it reached Europe. Concluding from this we can 
argue that if a bunch of Monarchs were to take flight above 950 mb — 
wich represents about 600 m height- around 29 th September (when 
the zonal flow was to be established), they could have been conceivably 
carried across the Atlantic at speeds around 30 kts and have ended 
up in Europe on 3rd October. Indeed, in the past, Monarchs have 
been reported (by pilots) on height levels of about 1500 m. The ‘weak’ 
point remains the question where they have been at night. During the 
yearly southerly remigration of Monarchs from Canada to Mexico, 
Monarchs normally don’t migrate during the night, but roost together 
in trees. However, Williams (1958) and Ross (1993) suggested that under 
certain conditions Monarchs are also able to fly at night, especially 
during warm nights when temperatures do not urge the butterfly to 
become inactive. Brower (1995) suggests that migration across large 
water masses like oceans is merely possible thanks to wind-assistance 
and hitch-hiking on ships, e.g. during the non-active periods. Although 
several occasions are described when Monarchs have been seen landing 
on the water-surface and after a while taking off again, Brower (1995) 
states that resting on the watersurface for longer than a few minutes 
would be lethal to the butterfly and is probably not the explanation 
for what Monarchs do at night when crossing oceans. 


136 


Previous to the period of suspected migration across the Atlantic the 
following observations at the East Coast of the USA were obtained 
from Dick Walton (pers.e-mail). At Cape May, New Jersey, his research 
includes 3 daily Pollard-type transects of approximately 5 miles each. 
The seasonal average for Monarchs per hour of censusing in 1995 was 
about 26. Daily averages during the suspected migration period were 
(Danaus plexippus per hour) : 


Sept. 26 - 15.79 
Sept. 27 - 26.00 
Sept. 28 - 69.31 
Sept. 29 - 32.54 
Sept. 30 - 36.21 


From these observations it can be concluded that there was indeed 
a build-up of Monarchs along the East Coast during the time period 
at which we suggest that migration should have started. Although 28th 
September was not the ‘biggest day of the year’, it was significantly above 
average for the season and may indicate a source for the transatlantic 
move of the Monarchs. 


We may conclude that at least the potential concentrations of Monarchs 
on the East Coast of the USA were present at the time at which the 
suitable weather conditions for migration across the Atlantic were 
establishing. 


Coinciding American bird observations 


Together with the Monarchs in the UK, many American bird species 
were reported. This strongly supports the American origin of the 
Monarchs. October 1995 is likely to be remembered as the best month 
ever for N. American warblers (fam. Parulidae) in Britain (Millington, 
1995). American warblers are such small, delicate birds, hardly larger 
than Monarchs! They normally migrate across the Gulf of Mexico. 
The fact that some of them were seen in Britain is as remarkable as 
Monarchs to cross the Atlantic. Some of these have only occurred 
once or twice in the UK. Different species of N. American warblers 


were reported between Ist and 29th October: Dendroica coronata, 
D. petechia, D. castanea, D. pensylvanica, D. striata, Parula ameri- 
cana (Millington, 1995). All of these are extremely rare migrants, with 
D. coronata being the most regular. Just like the Monarchs they were 
noticed mostly on Scilly Islands and in Cornwall. Blackpoll warblers 
(D. striata) were noticed for the first time in the UK coinciding with 
a previous Monarch migration in 1968 (Burton & French, 1969). In 


137 


1995 again 2 specimens of the same bird species, wich is extremely 
rare in the UK, were noticed in localities where also Monarchs were 
found (Cornwall and Isles of Scilly). 


Immigration of American passerine birds, small to medium land birds, 
was observed during the same period in the UK ; Zonotrichia leucophrys 
(fam. Emberizidae), Catharus ustulatus, C. guttatus, C. fuscescens (fam. 
Turdidae), Dolichonyx oryzivorus (fam. Icteridae) and Pheucticus 
ludovicianus (fam Cardinalidae). Other northern American birds were 
Charadrius vociferus, Pluvialis dominica (fam. Charadriidae), Limno- 
dromus griseus, L. scolopaceus, (fam. Scolopacidae), Coccyzus ame- 
ricanus (fam. Cuculidae), Anas discors (fam. Anatidae), Petrochelidon 
pyrrhonota (fam. Hirudinidae). Only the members of the Scolopacidae 
and Anas discors are more or less yearly visitors in the UK, but never 
common. All others belong to the ‘extremely rare’ category (1.e. far 
less than annual occurrence). Coccyzus americanus, the yellow-billed 
Cuckoo, has turned up only a dozen times, mostly on the Isles of 
Scilly (P. Doyle, pers. e-mail). The observation of 22 specimens of the 
red-eyed vireo (Vireo olivaceus) was remarkable, although this is an 
annual immigrant, esp. on Scilly Isles. Both birds and Monarchs were 
apparently displaced by hurricane conditions and both could have been 
at the mercy of strong westerly winds wich persisted across the Atlantic 
thereafter. 


Coinciding North American Lepidoptera 


Another sighting that supports the American origin of the Monarchs 
is that of the North American Vanessa virginiensis (Drury, [1773]) 
(Lepidoptera, Nymphalidae) on 13th October at Prawle Point, Devon, 
seen together with a Monarch on the same day. I would strongly 
recommend to examine the genitalia of all Nomophila Hubner, [1825] 
(Lepidoptera : Pyralidae) caught during the same period at the south 
coast. It seems almost predictable that the N. American Nomophila 
nearctica (Munroe, 1973) was also present as a migrant species. In 
the past at least one Nomophila nearctica was found on the British 
Isles on 22nd August 1919 (Pelham-Clinton, 1984). 


Conclusion 

In the past (1968), simultaneous migrations of D. plexippus and V. 
virginiensis have occurred, but for both species the Canary Islands 
(Spain) could have been a source of migration. It has always been 


a matter of dispute whether the migration did start in the Canary 


138 


Islands or in N. America. The present paper strongly supports evidence 
for a Northern American origin of all involved migrating species of 
Lepidoptera and birds in 1995, correlated with suitable meteorological 
conditions. If the Monarchs did not come from America, the timing 
is a surprising coincidence, as it then has also been in several other 
years (1968, 1981), given the potential displacement by the meteoro- 
logical conditions. 


Acknowledgements 


First of all I wish to underline that without the facilities of Internet, collecting 
so much information would have been impossible in such a short time. I 
have to agree that Internet, the way by which information can be shared 
so fast, is fantastic and opens very promising possibilities. It was really 
fascinating to receive so much valuable information from ’specialists ’ all over 
the world. Information cannot travel any faster ! I’d like to thank especially 
Roger Lane, UK for his very valuable suggestions, his very detailed local 
weather reports from Cornwall and for sending me a list with all the sightings 
of Monarchs and birds in the UK ; Nick Bowles, UK for compiling this list 
of sightings ; Simon Coombes <simon@captain.zynet.co.uk> for plotting, 
scanning and e-mailing of distribution maps of the Monarchs and mailing 
me the reference about the migration of Monarchs in 1968 in the UK ; Dick 
Walton, NJ, USA <rwaltonxxx@aol.com> for the kind communication of 
local reports of Monarch censusing and for his useful thoughts on fertility 
in Monarchs during migration; Mike Campbell, Newfoundland, Canada 
<campbellm@aesbed.am.doe.ca> for the meteorological analysis of the 
American weather conditions. Further I’d like to thank Mark Dixon, UK 
<gc77@dial.pipex.com> for the communication of the escapees of Monarchs 
in the UK ; Willem Pier Vellinga, Holland <w.p.vellinga@phys.rug.nl> for 
mailing me the observation of a Monarch in Holland ; Dirk Maes, Brussels 
<instnat@isl.vub.ac.be> for mailing me the observation in Belgium ; Dr. J. 
Dubois, France, for the kind communication of 4 sightings in France ; Paul 
Batty, UK for the facilities to publish requests about Monarch sightings in 
the ELG-list (Entomological Lifestock Group, UK); Patrick Doyle, UK 
<P.Doyle@aberdeen.ac.uk> for suggestions concerning bird migration to the 
UK. Last but not least I’d like to thank S. Coombes for plotting the very 
nice maps added to this paper. Especially thanks to A. Olivier for his revision 
and for supplying me more Monarch references than I could ever dream of. 


References 


BRETHERTON, R. F., 1984. Monarchs on the move-Danaus plexippus (L.) 
and D. chrysippus (L.). Proc. Trans. Br. ent. nat. Hist. Soc. 17 : 65-66. 
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139 


Hesperiüdae - Nymphalidae. The Butterflies. - pp. 283-285. Harley Books, 
Martins, Great Horkesley, Colchester, Essex. 

BRETHERTON, R. F. & CHALMERS-HUNT, J. M., 1982. The immigration of 
Lepidoptera to the British Isles in 1981 including that of the Monarch 
butterfly : Danaus plexippus L. Entomologist’s Rec. J. Var. 94 : 81-87, 
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Brewer, J. 1967. Wings in the meadow. Houghton Mifflin Company. Boston. 

Brower, L. P., 1984. Chemical defence in butterflies. Jn Vane-Wright, R. I. 
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Brower, L. P., 1995. Understanding and Misunderstanding the migration 
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J. Lepid. Soc. 49 (4) : 304-385. 

Burton, J. F. & FRENCH, R. A., 1969. Monarch butterflies coinciding with 
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Dusois, P. J., 1996. Observations de Monarques Danaus plexippus en France 
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Fett, E. P., 1928. Dispersal of insects by air-currents. Bull. New York State 
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LEMPKE, B. J., 1972. De Nederlandse Trekvlinders. (Tweede druk). Thieme 
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MILLINGTON, R., 1995. Bird-Watching Magazine UK, December 1995 : 83-85. 

PELHAM-CLINTON, E. C., 1984. A British specimen of Nomophila nearctica 
Munroe (Lepidoptera: Pyralidae). Entomologist’s Gaz. 35: 155-156. 

Ross, G. N., 1993. Butterflies round trips. Nat. Hist. 102 : 3. 

SCUDDER, S. H., 1876. A cosmopolitan butterfly. 1. Its history. Am. Nat. 
10 : 602-611. 

URQUHART, F. A., 1987. The Monarch Butterfly : international traveller. 
Neslon-Hall, Chicago. 

VANHOLDER, B., 1996. Trekvlinders in 1995, twaalfde jaarverslag (Lepidoptera). 
Phegea 24 (2) : 49-68. 

WALKER, J. J., 1914. The geographical distribution of Danaida plexippus L. 
(Danais archippus, F.) with especial reference to its recent migrations. 
Entomologist’s mon. Mag. 50 : 181-193, 224-237. 

WILLIAMS, C. B., 1958. Insect Migration. Collins, London. 

WILLIAMS, C. B, CockBiILL, G. F., Gisss, M. E. & Downes, J. A., 1942. 
Studies in the migration of Lepidoptera. Trans. R. ent. Soc. Lond. 
92 (1) : 101-283. 


140 


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Nota lepidopterologica 


Vol. 19 No. 3/4 Basel, 21.X11.1996 ISSN 0342-7536 


Editor : Alain Olivier, Luitenant Lippenslaan 43 B14, B-2140 Antwerpen- 
Borgerhout, Belgium. 


Assistant Editors : Dr. Roger Dennis (Wilmslow, GB) 
. PD Dr. Andreas Erhardt (Binningen, CH) 
Dr. Enrique Garcia-Barros (Madrid, E) 
Dr. Christoph Häuser (Stuttgart, D) 
Dr. Alexander Pelzer (Wennigsen, D) 
Dr. Erik J. van Nieukerken (Leiden, NL) 


Contents — Inhalt — Sommaire 


Wüsrt, P. : New and rare species of Heterocera (Lepidoptera) and some 
notes on their distribution in different habitats in the Nestos Delta, 
Mieedontau@reecet tr. Men ara tA D. ey ui. seen... à sons 142 

GARCIA PEREIRA, P. & GARCIA-BARROS, E. : Spot distribution in Ma- 
niola jurtina (Linnaeus, 1758) (Lepidoptera : Nymphalidae, Satyrinae) 


MCE nalSpama nun MARNE CERTA LE niit. 152 
STEFANESCU, C., LOCKWOOD, M. & CEBOLLA, M. : The butterfly assem- 

blages of northeastern Spanish wetlands .......................................... 165 
OLIVIER, A. & DE Prins, R. : The butterflies of the Greek island of Kos : 

a synthesis (Lepidoptera : Hesperioidea & Papilionoidea .................. 185 


BURGHARDT, F. & FIEDLER, K. : Myrmecophilous behaviours in cater- 
pillars of the butterfly, Polyommatus icarus (Rottemburg, 1775) : tem- 
poral patterns and age dependency (Lepidoptera : Lycaenidae) ......... 212 

PELZER, A. : Illustrierter Bestimmungsschlüssel für die Präimaginalsta- 
dien der Schwärmer Europas und Nordafrikas (Lepidoptera : Sphin- 
Ida) ee HeLa eo: AR) Delle. noise Ae 81s ein 230 

LOELIGER, E. A. & KARRER, F. : Breeding of an F, of Hyles hybrid vesper- 
tilioides (Boisduval, 1827) after induction with ecdysone of metamor- 
Phosismmdiapausalpupaed. HN cw Teams. lawianma.... 243 


Short communications — Kurze Mitteilungen — En bref 


SAMRAOUI, B. : Breeding status and range expansion of Danaus chry- 
sippus (Linnaeus, 1758) in the Algerian Sahara (Lepidoptera : Nym- 


jolnalbalae MI) Ata ANTS) tees RE N 261 
Book reviews — Buchbesprechungen — Analyses ................................ 264 
Vol. 19 — 1996 
Dates of publication — Publikationsdaten — Dates de publication 268 
COMMENTS TAILLES OINAITE eee c cote sens eses eme carenresere cones 268 
New taxa described in Vol. 19 — Neue Taxa in Band 19 beschrieben — 

Nouveausetaxardeenits’dansdlevol.tl9 N... nee 270 


Nota lepid. 19 (3/4) : 142-151 ; 21.XIL.1996 ISSN 0342-7536 


New and rare species of Heterocera (Lepidoptera) 
and some notes on their distribution in different habitats 
in the Nestos-Delta, Macedonia, Greece 


Peter WUsT 


Universitat GHS Essen, Fachbereich 9 - Allgemeine Zoologie, Universitätsstr. 5, 45117 Essen, 
Germany 


Summary 


During the present survey 1554 Heterocera of 190 species were caught in 
different habitats between the years 1988 and 1993 in the western part of 
the Nestos-Delta, Greece (Wiist, 1997). 13 species were newly recorded for 
Greece: Pelosia muscerda (Hufnagel, 1766), Dysauxes ancilla (Linnaeus, 
1767), Sideridis albicolon (Hiibner, [1813]), Hadena rivularis (Fabricius, 1775), 
Hemithea aestivaria (Hübner, 1799), Lycia hirtaria (Clerck, 1759), Cyclophora 
quercimontaria (Bastelberger, 1897), Cyclophora punctaria (Linnaeus, 1758), 
Cyclophora linearia (Hübner, 1799), Melanthia procellata ([Denis & Schiffer- 
miller], 1775), Apeira syringaria (Linnaeus, 1758), Chariaspilates formosaria 
(Eversmann, 1842) and Sclerocona acutellus (Eversmann, 1842). 14 species 
were recorded for the first time in the Nestos-Delta apart from being rare 
in other Greek regions : Eilema muscula (Staudinger, 1899), Diaphora mendica 
(Clerck, 1759), Ptilodon capucina (Linnaeus, 1758), Cerastis rubricosa ([ Denis 
& Schiffermüller], 1775), Mythimna turca (Linnaeus, 1761), Trachea atriplicis 
(Linnaeus, 1758), Ipimorpha subtusa ([Denis & Schiffermüller], 1775), Colo- 
bochyla salicalis ([Denis & Schiffermüller|, 1775), Herminia tarsicrinalis 
(Knoch, 1782), Macaria notata (Linnaeus, 1758), Chloroclystis v-ata (Haworth, 
1809), Lomographa dilectaria (Hübner, 1799), Ourapteryx sambucaria (Lin- 
naeus, 1758) and Epione repandaria (Hufnagel, 1767). Some of these reach 
their distribution limit in the Nestos-Delta. 


Zusammenfassung 


Im Nestos-Delta wurden in den Jahren 1988 bis 1993 bis jetzt 1554 Nachtfalter 
aus 190 Arten nachgewiesen (Wiist, 1997). Dabei wurden 13 Arten erstmals 
fiir die Fauna von Griechenland nachgewiesen : Pelosia muscerda (Hufnagel, 
1766), Dysauxes ancilla (Linnaeus, 1767), Sideridis albicolon (Hübner, [1813]), 
Hadena rivularis (Fabricius, 1775), Hemithea aestivaria (Hübner, 1799), Lycia 
hirtaria (Clerck, 1759), Cyclophora quercimontaria (Bastelberger, 1897), 
Cyclophora punctaria (Linnaeus, 1758), Cyclophora linearia (Hübner, 1799), 


142 


Melanthia procellata ([Denis & Schiffermiiller], 1775), Apeira syringaria 
(Linnaeus, 1758), Chariaspilates formosaria (Eversmann, 1842) und Sclerocona 
acutellus (Eversmann, 1842). Es wurden 14 weitere Arten nachgewiesen die 
für das Nestos-Delta neu und in anderen Regionen Griechenlands sehr selten 
sind : Eilema muscula (Staudinger, 1899), Diaphora mendica (Clerck, 1759), 
Ptilodon capucina (Linnaeus, 1758), Cerastis rubricosa ([Denis & Schiffer- 
müller], 1775), Mythimna turca (Linnaeus, 1761), Trachea atriplicis (Linnaeus, 
1758), Ipimorpha subtusa ([Denis & Schiffermüller], 1775), Colobochyla 
salicalis ([ Denis & Schiffermüller], 1775), Herminia tarsicrinalis (Knoch, 1782), 
Macaria notata (Linnaeus, 1758), Chloroclystis v-ata (Haworth, 1809), Lo- 
mographa dilectaria (Hübner, 1799), Ourapteryx sambucaria (Linnaeus, 1758) 
und Epione repandaria (Hufnagel, 1767). Einige dieser Arten leben im Nestos- 
Delta an der Randzone ihrer Verbreitung. 


Introduction 


Heterocera were caught in different habitats in the years 1988 to 1993 
in the western part of the Nestos-Delta in the North-east of Greece 
(40° 53’ N and 24° 43’ E) (Map 1). The investigation was part of 
a research programme conducted by the University of Essen (Szijj et 
al., 1996). 


Map 1. Position of survey area. 


Materials and methods 
The Macroheterocera were caught by 15 W superactinic light traps 


of the Minnesota type. The traps were used at nine different sites (Wüst, 
1994) (Map 2): 


143 


— (A) on a costal salt marsh 

— (B) on the beach dune of a peninsula at the west end of Keramoti, 
near Phragmitea 

— (C) on the roof of a house in Keramoti 

— (D) on a beach dune near Keramoti adjacent to the dune forest 

— (E) in a dune forest of a riparian forest character 

— (F + G) on an inland dune | 

— (H) in the centre of a native, fenced-in river forest (hardwood 
riparian forest) 

— (J) at the edge of the river forest (H), opposite to the poplar forest 

— (K) in the middle of a cultivated poplar forest consisting of Populus 
canadensis trees (hybrid) 


Results 


Pelosia muscerda (Hufnagel, 1766) was recorded in Greece for the first 
time. The southern distribution area of this species reaches its maximum 
extent in the Nestos-Delta. This species is endangered as swamps have 
become more and more scarce in Greece and particularly in the Nestos- 
Delta. In all, 13 individuals of this species were caught : three animals 
at the fringe of the river forest, two inside this forest, five in the poplar 
forest opposite and three in the dune forest. The first specimen was 
caught on the 12.5.1989 in the river forest. The observed flight season 
extended from the middle of May (12.5.) to the middle of July (14.7). 


Eilema muscula (Staudinger, 1899) was caught in different habitats. 
Twenty animals were recorded altogether, the first one on the 11.5.89 
at the edge of the river forest. Eight specimens were caught within 
the river forest, six animals in the dune forest, four animals at the 
coastal dune and one on the inland dune. There is a possibility that 
this species displays three generations, as the animals were found in 
the months of May, July and September. It is also possible that this 
species distribution further is not just restricted to the island of Rhodos 
and the Turkish mediterranean coast (de Freina, 1987) but may also 
be present in coastal areas in Greece. 


Five specimens of Diaphora mendica (Clerck, 1759) were caught in 
the Nestos-Delta. The first individual was found on the 10.5.1988 at 
the fringe of the hardwood river forest. This shows that D. mendica 
is present at the Greek coast. Three more specimens were caught on 
the 9.5.89 in the same habitat and one more was found on the 25.4.92 
on an inland dune. 


144 


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145 


On the 17.5.89 Dysauxes ancilla (Linnaeus, 1767) was recorded in 
Greece for the first time and was caught at the fringe of the hardwood 
river forest (WüsT 1994). A total of 9 animals was found in three 
different habitats : on the 21.5.89 three animals were recorded in the 
bushy fringe of the hardwood river forest and on the 29.5.89 and the 
5.7.91 one animal each time was caught at the same site. On the 8.9.93 
two specimens were caught in the dune forest with a dense vegetation. 
On the 30.5.89 D. ancilla was found 150m from the water line on the 
beach dune dominated by Hypericum perforatum. As that night wind 
was low and the dune forest was as near as 300 m, this animal could 
as well have been a visitor from the dune forest. In Greece D. ancilla 
displays two generations. The first generation is active from May to 
July and the second from September to October. Two individuals of 
this species were also recorded by Peter Kuhna on the 15.10.1990 near 
Volakas (province Drama). Keeping that in mind, it is possible that 
D. ancilla has a wider distribution in the shady forests in Greece and 
also that this species would more often be found if more work was 
done on the distribution of Lepidoptera in Greece. 


Ptilodon capucina (Linnaeus, 1758) reaches the southernmost limit of 
its distribution in the Nestos Delta and was found newly in this coastal 
area. One specimen was recorded on the 9.8.93 in the hardwood river 
forest. 


As Mentzer’s work on the splitting of Noctua janthina ([Denis & 
Schiffermüller], 1775) in three species was published in 1991 there have 
been only a few reliable data (von Mentzer et al., 1991) about the 
distribution of Noctua tertia (von Mentzer, Moberg & Fibiger, 1991) 
in Greece since then. In the Nestos-Delta only N. janthina has been 
found until now. On the 5.7.91 four N. janthina were caught for the 
first time in the poplar forest. Two more individuals were recorded 
on the 11.07.91 on the costal salt marsh and on the 14.7.91 in the 
dune forest. 


Apart from Kozani the Nestos-Delta is the second place where Ce- 
rastis rubricosa ([ Denis & Schiffermüller], 1775) has been found in 
Greece. This confirms Hackers supposition that this species has a wider 
distribution in Northern Greece (Hacker, 1989), where it reaches the 
limit of its southern distribution. Four animals were found altogether 
during the study, on the 9.4.93 (3) and 10.4.93 (1) in the river forest. 


As regards Sideridis albicolon (Hübner, [1813]) Hackers supposition 
is confirmed that this species should be found in northeast Greece 
(Hacker, 1989). According to its ecology, S. albicolon prefers sandy 


146 


and ruderal soil. One specimen was caught on the 7.5.88 for the first 
time in Greece on a peninsula-like coastal dune near Keramoti. 


Hadena rivularis (Fabricus, 1775) was found new in Greece on the 
13.5.88 in its characteristic habitat at the fringe of the river forest. 
Eleven animals were caught altogether, at the fringe of the river 
forest (7), in the centre of the river forest (2), as a visitor in the poplar 
forest (1) and in Keramoti (1). This proves Hackers presumption (1989) 
that this species should be present in Northern Greece. 


Mythimna turca (Linnaeus, 1761) has been recorded so far only in 
Leptokaria near the Olymp (Hacker, 1989). In the Nestos-Delta one 
animal of this species was caught in the poplar forest on the 17.5.89. 
Regarding the biology of this species it might have been a visitor from 
the river forest, the latter being opposite to the poplar forest. This 
species may also be found in low numbers in other habitats due to 
its specific habitat requirements. 


Contrary to Hacker’s presumption (1989), Trachea atriplicis (Linnaeus, 
1758) could also be found in the northeast of Greece apart from the 
few specimens found in the Olymp area, the west and the northwest 
of Greece. On the 5.7.91 one individual was caught in the poplar forest. 


Ipimorpha subtusa ({Denis & Schiffermüller], 1775) has been recorded 
for the second time in Greece 1.e. after it was found in Kavisos (East- 
Thracia) in 1984 (Hacker, 1989). It was recorded in the poplar forest 
on the 17.5.89 and on the 5.7.91. I. subtusa might display an even 
wider distribution in northern Greece. 


So far Colobochyla salicalis ({Denis & Schiffermüller], 1775) has only 
been found in the Disron Mountains near Terpilos in 1982 (Behounek, 
1984). During the present survey 20 specimens were found in the Nestos- 
Delta, at the fringe of the hardwood river forest (12), in the river 
forest (3), in the polar forest (2) and in the dune forest (3). On the 
9.5.88 this species was found for the first time in the Nestos-Delta 
at the fringe of the river forest. It should be quite common in suitable 
habitats in northern Greece. 


Since 1981 Herminia tarsicrinalis (Knoch, 1782) has only been reported 
once from Greece, in the Olymp area near Platamon (Hacker, 1989). 
In the Nestos-Delta this species is one of the most common. It was 
caught for the first time on the 11.5.89 at the fringe of the hardwood 
river forest. 19 animals were caught altogether, at the fringe of the 
river forest (17), in this forest (1) and in the poplar forest opposite 
to the river forest (1). This species should therefore also be common 


147 


in northern Greece especially in mixed forests consisting mainly of 
Beech-trees. 


Macaria notata (Linnaeus, 1758) has so far been published only by 
Staudinger 1870 (leg. Krüper) as present in Greece. It probably was 
present in the Parnassos-region (Gozmany, pers. comm.). This eurasiatic 
species could also be found in the Nestos-Delta. Two animals were 
caught in the hardwood river forest and one animal in the poplar forest 
opposite to the river forest. The first newly found animals of this species 
were caught on the 17.5.89 in both habitats. 


Hemithea aestivaria (Hübner, 1799) is an eurasiatic species, its dis- 
tribution extends in the south from Spain to Japan. It was found for 
the first time in Greece on the 9.5.89 at the fringe of the hardwood 
river forest. In the course of this survey six animals were recorded 
there. 


Lycia hirtaria (Clerck, 1759) was the third-most-frequently recorded 
species in the Nestos-Delta. However, more individuals were seen 
around the light-traps than were caught. This eurasiatic species is widely 
distributed but it was unknown for the fauna of Greece so far. This 
does not only depend on its early flight period but also on the scarcity 
of lepidopterological activities in Greece. Most of the foreign ento- 
mologists start their research in June. 74 animals were caught altogether 
by light-traps, at the fringe of the hardwood river forest (21), in this 
river forest (20), in the poplar forest opposite to the river forest (12) 
and on the inland dune (21). On the 24.5.89 this species was caught 
on the inland dune, and this for the first time in Greece. In the Nestos- 
Delta L. hirtaria was flying in one generation from early April until 
late May. The maximum of flight activity was recorded in late April. 


The eurasiatic species Cyclophora quercimontaria (Bastelberger, 1897) 
is locally common in the Balkan Peninsula (Gozmany, pers. comm.). 
It was caught in Greece for the first time on the 10.8.93 in the river 
forest of the Nestos-Delta. The specimen caught was fresh, which leads 
to the assumption that C. quercimontaria has two generations in the 
Nestos-Delta. 


One individual of the eurasiatic species Cyclophora punctaria (Linnaeus, 
1758) was caught for the first time in Greece on the 23.4.92 in the 
hardwood river forest. 


On the 12.5.89 the eurasiatic species Cyclophora linearia (Hübner, 1799) 
was caught for the first time in Greece. Eight specimens were recorded 
altogether, at the fringe of the hardwood river forest (4) and inside 
this forest (4). 


148 


One individual of the eurasiatic species Melanthia procellata ([Denis 
& Schiffermiiller], 1775) was caught for the first time in Greece on 
the 14.7.91 at the fringe of the hardwood river forest. 


The eurasiatic species Chloroclystis v-ata (Haworth, 1809) has been 
recorded so far only in Olympia (Hacker & Lukasch, 1984) and in 
Gonia (Reisser, 1974). In the Nestos-Delta one individual was caught 
on the 20.4.92 in its natural habitat at the edge of the hardwood riparian 
forest. 


Lomographa dilectaria (Hübner, 1799) is a southeastern species known 
from Bulgaria and Macedonia and common in the whole of Turkey 
(Kuhna, pers. comm.). According to Kuhna this species should also 
be widely distributed throughout the whole of Greece. So far it has 
been recorded in Taygetos (Thurner, 1967) and Asprovalta (Hacker & 
Lukasch, 1984). 12 animals were caught in the Nestos-Delta altogether. 
Here it was found near to its hostplant Populus. Four animals were 
found at the fringe of the hardwood river forest and five animals were 
recorded in the poplar forest. The first recording of L. dilectaria was 
on the 9.5.88 at the fringe of the river forest. 


Two individuals of Apeira syringaria (Linnaeus, 1758), an eurasiatic 
species were caught in the Nestos-Delta. One was recorded at the fringe 
of the river forest and one in the forest. A. syringaria was first recorded 
in Greece on the 13.5.89 in the river forest. 


Ourapteryx sambucaria (Linnaeus, 1758) was known to be present in 
two places in Greece. In 1988 Hassler, Schmidt and Feil caught this 
species in Neraidorachi (mount Chelmos, 1800 m, 16.7.) and in Xiro- 
kambi (Taygetos, 1000 m, 27.7.) (Gozmany, pers. comm.). The third 
record for Greece was on the 5.7.91 in the poplar forest. Therefore 
the distribution of O. sambucaria is extended in Greece from the coast 
to mountaneous regions. 


The eurasiatic species Epione repandaria (Hufnagel, 1767) has so 
far been recorded and published by Rebel (1905) from Kalavryta, 
Peloponnese. This species was found in two habitats with moist soil : 
on the 21.5.89 at the fringe of the river forest and on the 28.5.89 in 
the dune forest. 


The distribution area of Chariaspilates formosaria (Eversmann, 1842) 
is postglacially disjunct. C. formosaria lives in warm swamps of south- 
eastern Europe, e.g. in Hungary and it should thus be present in the 
Romanian swamps of the Danube (Gozmany, pers. comm.). In the 
Far East it can be found in the Amur region. In Greece, C. formosaria 


149 


was recorded on the 26.5.89 for the first time. It was caught on an 
inland dune of the Nestos-Delta. There it lives in damp dune valleys 
with Phragmitetea (Szyj et al., 1996). This patchily distributed, rare 
species is more and more endangered in Greece by farming activities 
in the Nestos-Delta. 


Sclerocona acutellus (Eversmann, 1842) was recorded on the 19.5.89 
for the first time in Greece in the river forest. This species is rare 
everywhere in Europe because it is characteristically found in swamps 
(Gozmany, pers. comm.). It is especially endangered in the Nestos- 
Delta as damp biotopes become more and more scarce. 


Acknowledgements 


I would like to thank Mr. P. Kuhna, Wipperfürth, for reviewing and re- 
determinating my collection of Greek moths. My special thanks to Dr. L. 
Gozmany, Budapest, who contributed his data on the distribution of the 
Geometridae to this piece of work. Thanks are also due to C. Ackermann, 
Hattingen, and Dr. P. King, Galway, for reviewing my manuscript. 


References 


BEHOUNEK, G., 1984. Ergebnisse einer Sammelreise durch Nordgriechenland 
und Anatolien (Lepidoptera: Noctuidae). Nachr. ent. Ver. Apollo 4: 
71-80. 

FORSTER, W. & WoHLFARTH,T. A., 1955-1981. Die Schmetterlinge Mittel- 
europas. 5 Bande, Franckhsche Verlagshandlung, Stuttgart. 

DE FREINA, J. & Wirt, T., 1987. Die Bombyces und Sphinges der West- 
palaearktis. Band 1. Edition Forschung & Wissenschaft Verlag, Miinchen. 

Hacker, H., 1989. Die Noctuidae Griechenlands. Herbipoliana 2: 1-589, 


I-XIL. 
HACKER, H. & LuKascu, J., 1984. Geometriden und Noctuiden aus der 
Sammlung J. Lukasch — ein Beitrag zur Fauna von Griechenland. 


Spixiana 7 : 51-61. 

REBEL, H., 1905. Lepidopteren aus Morea, II. Nachtrag. Berl. Ent. Z. 50: 
291-314. 

REISSER, H., 1974. Zur Lepidopterenfauna Kretas : neue Zugänge zum Arten- 
bestand. Ann. Mus. Goulandris 2 : 137-140. 

Sz1JJ, J., DETSIS, V., Wust, P. & WAGNER, H.J., 1996. Das Nestosdelta. For- 
schungen zur Okologie und naturschiitzerischen Wertanalyse. Berichte 
aus dem Arbeitsgebiet Entwicklungsforschung. Heft 26. Geographisches 
Institut der Universitat Miinster. 

THURNER, J., 1967. Lepidopteren aus Morea. Ein weiterer Beitrag zur Fauna 
des Peloponnes (Griechenland). Z. wien. ent. Ges. 52 : 50-58. 


150 


VON MENTZER, E., MOBERG, A. & FIBIGER, M., 1991. Noctua janthina ([ Denis 
& Schiffermüller]) sensu auctorum a complex of three species (Lepi- 
doptera : Noctuidae). Nota lepid. 14 (1) : 25-40. 

Wust, P., 1994. Faunistic characteristics and ecology of the Macroheterocera 
in different structured areas of Nestos Delta. Bios (Macedonia, Greece) 
2 : 263-269. 

Wüsrt, P., 1997.: A first contribution to the Macroheterocera fauna (Lepi- 
doptera) in the Nestos-Delta, Macedonia, Greece. Hellenic Zoological 
Archives (in print). 


151 


Nota lepid. 19 (3/4) : 152-164 ; 21.XII.1996 ISSN 0342-7536 


Spot distribution in Maniola jurtina (Linnaeus, 1758) 
(Lepidoptera : Nymphalidae, Satyrinae) in Central Spain 


Patricia GARCIA PEREIRA* & Enrique GARCIA-BARROS** 


* Rua Carlos Mardel, 121, 1Dto.,1900 Lisboa, Portugal 
** Departamento de Biologia (Zool.),Universidad Autonoma de Madrid, Cantoblanco, E-28049 
Madrid, Spain 


Summary 


The pattern of wing spotting in the satyrine Maniola jurtina (Linnaeus, 1758) 
was investigated in a sample of over 900 collection specimens from Central 
Spain. The frequency distributions of spots, and their relationships with other 
variables (date of collection, adult size, site elevation), were determined and 
compared with published data concerning Northern and Southern Iberian 
populations. 

The general spotting pattern fits well to those described from South Iberian 
populations (especially from the Portuguese Algarve), showing no transition 
towards North Iberian phenotypes. The number of spots decreases towards 
the end of the summer season, a trend that is shared by Mediterranean popu- 
lations that are phenotypically distant (e.g., Iberian and Italian). This may 
be taken as evidence for a relationship between female aestivation behaviour 
and some kind of selection against high-spotted morphs. However, the fact 
that the same trend appeared in both sexes (while males are not known to 
aestivate) indicates that other underlying factors probably exist, and that these 
may operate on both sexes. This idea is reinforced by the fact that spotting 
was significantly correlated to adult size. 


Resumo 


O padräo de pontos existentes nas asas de Maniola jurtina (Linnaeus, 1758) 
fol observado numa amostra com mais de 900 espécimes pertencentes a 
colecçôes entomolögicas, recolhidos na zona Central de Espanha. Analisou- 
se a distribuicäo de frequéncias dos pontos, a relacäo com outras variäveis 
quantificadas (tamanho dos individuos, data e altitude) e procedeu-se à com- 
paracäo destes dados com outros trabalhos publicados sobre populaçôes da 
Peninsula Ibérica. 

A distribuicäo de pontos é semelhante ao descrito para populaçôes do Sul 
(especialmente à zona do Algarve, Portugal). As populaçôes do Sul e Centro 
da Peninsula Ibérica säo homogéneas quanto a esta caracteristica, nao tendo 


152 


sido detectada uma zona de transicäo fenotipica ou a fronteira com o Norte 
da Peninsula Ibérica. 

Verificou-se a diminuiçäo do numero de pontos ao longo do Veräo, parti- 
cularmente nas fêmeas. E um fenömeno que ocorre igualmente noutras popu- 
lacöes mediterrâneas e que € independente da distribuicäo dos pontos. Tem 
sido explicado pela accäo da selecçäo natural, que elimina os fenötipos com 
maior numero de pontos, associada a um comportamento de estivacäo das 
femeas. No entanto, o facto de também ocorrer diminuiçäo nos machos (que 
näo estivam) e existir uma correlagäo significativa entre o padräo de pontos 
e o tamanho dos adultos, parece indicar que existem outros factores que 
actuam em ambos os sexos. 


Introduction 


The distribution of spots on the underside of the wings of the mea- 
dow brown butterfly, Maniola jurtina (Linnaeus, 1758), has been the 
subject of numerous studies over more than 40 years. Dowdeswell and 
McWhirter (1967) analysed samples from the whole geographic range 
of the species in order to evaluate the variability of this feature. They 
found a highly uniform spotting pattern over most of continental Europe 
and Great Britain which they termed “General European Area”. In 
addition, smaller zones (“peripheric areas”) showed particular spot- 
ting patterns. It seems that the spot distribution 1s stable over long 
periods of time, as well as over wide geographic areas, in the two 
sexes (Dowdeswell & McWhirter, 1967 ; Ford, 1975 ; Brakefield, 1984 ; 
Brockie, 1972 ; Shreeve, Dennis & Williams, 1996). In general, the 
females display a remarkably constant spotting pattern while the males, 
although unimodal at 2 spots, are far more variable (Ford, 1975). The 
General European Area was explained as a result of the powerful action 
of natural selection, which would produce a uniform pattern in spite 
of important environmental variation. Because of their small size and 
isolated character, the peripheric areas might have achieved different 
patterns by adjustment to local conditions (Brockie, 1972). Field data 
indicate that intra-seasonal shifts in spot distribution are restricted to 
these peripheric areas. While natural selection is generally accepted as 
the ultimate cause of these patterns, there is no agreement about the 
precise factors that are responsible for such variation. Proposed ex- 
planations have invoked climate (e.g. by Bengston, 1978 ; Scali, 1972), 
behavioural adaptation (Brakefield, 1984), habitat heterogeneity (Mc 
Whirter, 1957) and predation (Brakefield, 1984 ; Owen & Smith, 1990). 


The situation on the Iberian Peninsula (one of the peripheric areas) 
is not well studied. There is some evidence indicating sharp differences 


153 


between the Northern and the Southern Iberian populations, the latter 
being characterized by a very low spot average in females (large modes 
at 0 spots) and a high spot average in males (mode at 2-3 spots), as 
well as by particular costality values, high in females and low in males 
(see Owen & Smith, 1993). The work made on South European popu- 
lations (Italy : Scali & Masetti, 1975 ; Algarve : Owen & Smith, 1993) 
revealed that a marked decrease in female spotting occurs along the 
summer (i.e., intra-seasonal shift). Since the long summer drought has 
important effects on the behaviour and phenology of the species in its 
Mediterranean range (Masetti & Scali, 1972 ; Garcia-Barros, 1987), it 
is likely that some of the peculiarities of the South European peninsular 
populations could be related to climatic factors, and more specifically 
to the presence of a long, hot and dry summer season. 


The present work relies on data gathered from populations in the 
central area of the Iberian Peninsula, which are presumably adapted to 
a long, adverse summer (Garcia-Barros, 1987). On this basis, the aims 
of this study are : First, to make available updated data from Central 
Spain, in order to contribute to the delimitation of the boundary 
between the North and South spotting pattern of the species within 
Iberia. Second, and related to the former, to check whether or not the 
spotting pattern of Central Spanish M. jurtina is consistent with that 
of South Iberian populations. Third, to assess the relationship between 
the presence of a dry summer period (and female aestivation behaviour) 
and spotting, by trying to discover whether a seasonal shift occurs in 
this area. Finally, to investigate the relationships between spotting and 
two variables (adult size, elevation) that might be related to spotting, 
and might help to explain part of the variation found as a result of 
phenotypic plasticity. 


Material and methods 


The sample consisted of 925 set butterflies from the collections at the 
Universidad Autonoma de Madrid (661 specimens), Natural History 
Museum at Madrid (167), and the Universidad Complutense de Madrid 
(97). Collection dates cover a period of 80 years (1912 to 1991), and 
the selected area spreads over about 200 km around Madrid city, with 
individuals collected in the provinces of Madrid, Segovia, Guadalajara, 
Cuenca, Avila and Teruel. This area is characterized by an important 
elevational gradient (400 to 1500 m in our samples). To cope with this 
heterogeneity, we compared every result from the pooled sample with 


154 


a)CENTRAL IBERIA 


c)SOUTH IBERIA 


50 


[æ) 
EN 
N 
& 
> 
on 


d)NORTH IBERIA 


) 1 2 3 4 5 
e)CENTRAL ITALY 
% 80 
60 - 
40 - 


[@) 
_—_ 
N 


Number of spots 


LEFT: FEMALES; RIGHT: MALES 


0 1 2 3 4 5 
50 | 
25 
0 
0 1 2 3 4 5 
50 | 
25 + 
| 
0 
0 1 2 3 4 5 


Fig. 1. Spot distribution in Southern populations (x-axis = number of spots ; y-axis = %). 


a) data from this study 
b) data from Owen & Smith (1993) 


c), d) data from Dowdeswell & McWhirter 


e) data from Scali & Masetti (1975) 


155 


those derived from a sub-sample (443 individuals) collected by A.Pino 
in a single site (Casa de Campo, close to the city of Madrid, 1980-1981). 


All measurements and spot counts were made on the left wing, and 
by the same person, to ensure consistency. The spots on the underside 
of forewing and hindwing were counted. Wing length was measured 
from base to apex of the forewing, to the nearest 1.0 mm. For each 
specimen, the collection site elevation (meters a.s..) and date were 
recorded. The spots were labelled following Owen & Smith (1993), 
that is 1 to 5 on the hindwing, and 1 to 4 on the forewing (anal to 
costal). The Splay, Median and Costality index were calculated after 
Owen & Smith (1993). 


The data were compared with available information on other Iberian 
areas (data after Owen & Smith, 1993 ; Dowdeswell & McWhirter, 
1967). Published data from Italy, Wales and the Swedish Archipelago 
were included as a contrast (using the spot frequencies given by Scali 
& Masetti, 1975 ; Shreeve, Dennis & Williams, 1996 ; Bengtson, 1978). 
The frequencies of each hindwing spot in males and females were used 
as variables, and a correlation matrix generated. This was used to 
construct a dendrogram (UPGMA procedure, with Pearson correlation 
as distance estimate) showing the affinities between the different areas. 


For regression and correlation analyses, wing length was logarithmically 
transformed, and spotting (number of spots) was squared-root trans- 
formed. 


0,35 — 
0,3 Spot 2 
= Gas À 
cS @. © Spot 1 Ei 
3 02 m “ 
= ae | 
EH 015L 
— 
© 0,1 + 
©. 
A 005 «Xe Spot 3 


Spots 


6 7 Month 8 d 


Fig. 2. Seasonal variation of spot average in the Central Iberian sample [bars : + stan- 
dard deviation]. 


156 


Males Females 
2 
4 
3 1 
24 
0 
Dome 9 GT ES" 9 


Fig. 3. Seasonal variation in the frequencies of individual spots (spot 1 = costal, to 
spot 5 = anal) in Maniola jurtina (Linnaeus, 1758) from Central Iberia. 


Table 1 


Number of spots on the hind- and forewing of Maniola jurtina (Linnaeus, 1758) 
from Central Spain (pooled sample), and a single site (Casa de Campo). 
The figures given summarize the number of individuals with 0 to 5 spots, 

sample size (n), and mean number of spots (x) 


N° hindwing spots N° forewing spots 
3 5 


Males 
Casa de Campo 


Total 

Females 

Casa de Campo 
Total 


Results 


The frequencies of hindwing and forewing spot numbers are given in 
Table 1. Male and female spot distributions were markedly different 
in both forewing and hindwing patterns. Both sexes showed little 
variation in forewing spotting, which was particularly low in the males. 
Hindwing spot average was of 3.033 in males and 0.581 in females. 
A comparison of the spot distributions of the pooled sample and the 
collection from Casa de Campo presented no evidence for a statistically 
significant difference (X?s, = 3.361 ; P > 0.05, for males ; X73) = 0.747 ; 


1157 


P > 0.05, for females). The females are strongly unimodal at 0 spots, 
while males have a more “flat-topped” distribution, with high propor- 
tions of 2, 3 and 4 spotted individuals (Fig. la). Table 2 shows the 
percentage of hindwing phenotypes in our sample, as well as the overall 
Costality, Splay and Median indices. Twenty-four different phenotypes 
were found, of which five are restricted to the female sex and six to 
the males. 


The comparison between our data and those from other areas indicated 
close affinity between Central Spain and Algarve (the clustering 
hierarchy is shown in Fig. 4). 


Table 2 


Hindwing phenotypes found in female Maniola jurtina (Linnaeus, 1758) 
from Central Iberia, and their relative frequencies and values for three descriptive indices, 
that are often used in research on this species (see text) 


Phenotypes Females 


Total 359 
Costality Index 0.457 
Splay Index 0.56 
Median Index 0.159 


158 


Table 3 


Relationships between the number of spots, size, date of capture and elevation, 
as measured by Pearson correlation (n = sample size ; p = significance level) ; 
[females-upper right half of the table ; males- lower left half)] 


Elevation Date Forewing N hindwing N° forewing 
length spots spots 


Elevation 0.166 — 0.0854 0.0121 0.0191 


ne A n= 560 n= 570 n = 580 

p = 0.0003 p = 0.0434 p = 0.7734 p = 0.6457 
0.0612 — 0.0496 — 0.2096 — 0.073 
n= 280 n= 450 n = 460 n = 469 


p= 0.3072 p=0.0000 p=0.1144 


— 0.1248 0.1536 — 0.0058 
n= 347 n = 274 n= 552 n = 560 
p= 0.0201 p = 0.0003 p = 0.8906 


~ 0.0151 0.1627 
n= 355 n= 281 n= 350 n Son 
p = 0.7775 p = 0.0001 
0.16 
n= 356 
p = 0.7636 


p = 0.2940 


Forewing length 


N° hindwing spots 


[N° forewing spots 


Algarve 

Central Iberia 

South Iberia 

North Iberia 
——. À Central Italy 

North Wales 

Southern Sweden 


0.80 OCT) 


Fig. 4. Dendrogram showing the affinities between Central and other Iberian popu- 
lations of Maniola jurtina (Linnaeus, 1758). Data from three non-Iberian populations 
are included as a contrast. Affinities are based on the frequencies of spots on the hind- 
wings of females and males, using Pearson correlations as a measure of similarity and 
the unweighted arithmetic average clustering method (UPGMA). 

[Algarve = data from Owen & Smith (1993) ; North and South Iberian = Dowdeswell 
& McWhirter (1967) ; North Wales = Shreeve et al. (1996) ; Southern Sweden (island 
populations) = Bengtson (1978)]. 


159 


The correlations between spot number, date of capture, wing size and 
collection site elevation are shown in Table 3. Results for both sexes 
were consistent, the number of hindwing spots being significantly cor- 
related with date and forewing length. In other words, larger individuals 
tend to present a higher number of spots, while spotting decreases 
during the season (see also Fig. 2 and Table 4). Elevation was negatively 
correlated with forewing length in both sexes. Since variation in male 
forewing spotting was small, and female fore- and hindwing spotting 
were positively correlated (Table 3), all subsequent analyses were 
restricted to the number of hindwing spots. 


Table 4 


Monthly variation in spotting patterns (hindwing underside) 
n = sample size ; x = mean values) 


N° hindwing spots 


Males 
May 
June 
July 
August 
Females 
May 
June 
July 
August 
September 


Stepwise multiple regression analyses were used to discover the relative 
importance of each variable in the number of hindwing spots, and 
to control for any effects of elevation (since elevation and date were 
correlated in females : Table 3). Several runs were performed regressing 
spot number on date, wing length and elevation. The last three variables 
were entered in different orders, and both separately for each sex or 
for all the data pooled. All analyses were consistent, and thus the results 
are not given in full. The variable ,date” accounted for the largest 
proportion of the variance in spot numbers (R2 = 0.033 for males ; 
R? = 0.043 for females). Adding the variable „forewing length” to the 
regression raised the explanatory power (R2 = 0.064 for males ; R2 = 
0.063 for females) ; the effect of „elevation”, once the other two variables 
had been entered, was negligible (R? = 0.061 for males ; R? = 0.062 
for females). We may thus conclude that date is the main source of 
variation related to spot number. 


160 


Fig. 2 represents monthly mean spot number. The trend towards low 
spotted patterns as the summer progresses is more evident in the 
females, and seems to operate in the same way for each of the individual 
spots (especially for spots 1, 2 and 4, since spots 3 and 5 are relatively 
rare ; see Fig. 3). Chi-square comparisons revealed significant differen- 
ces between June and September females Ka) = 15.396 ; 0.001 << P 
< 0.01), as well as between July and September females (X73) = 12.540 ; 
0.001 << P < 0.01). As far as our material allowed for more detailed 
comparisons, the results are generally true for the samples from Casa 
de Campo (see Table 5). 


Table 5 


Relationship between the number of hindwing spots and date and wing size, 
measured in butterflies from a single population (Casa de Campo, Madrid). 
Values are Pearson correlation coefficients (n = sample size ; P = significance level) 


n Date Wing length 


Males 


1980 95)" 01142 ~ 0.013 
(P = 0.171) (P = 0.09) 
1981 19, 210.392 — 0.085 
(P < 0.001) (P = 0.455) 
1980/1981 KLAR, 0250 ~ 0.015 


(P < 0.001) (P = 0.844) 


Females 


1980 163 —0.268 0.016 

(P = 0.016) (P = 0.016) 
1981 63 0.294 0.201 

(P = 0.02) (P = 0.115) 
1980/1981 226 0.273 0.175 


(P <0.001) (P = 0.008) 


Discussion 


The comparison between the spot distributions, as well as the Costality, 
Median and Splay index values, reveals high similarity between Central 
and South Iberian populations (e.g. data from Owen & Smith, 1993 and 
this study), especially in females. In agreement with previous studies, 
males are more variable than females (in fact, males might be used 
to characterize Central Iberian populations relative to e.g. those from 
Algarve). Our results show no South to North transition, and thus 
support the contention that most of the Iberian Peninsula is occupied 
by a stable phenotypic distribution, which probably coincides with the 
so-called subspecies M. jurtina hispulla (Esper, [1805]) (Thomson, 1973). 


161 


Whether or not a transition zone exists, not far away to the North, 
remains unknown. 


Our finding of a seasonal trend towards reduction in spot number 
in females (and less markedly in males) fits the similar intra-seasonal 
shifts described from other Mediterranean areas, like Algarve (Owen 
& Smith, 1993) and Central Italy (Scali, 1975), where the females are 
known to aestivate (Scali & Masetti, 1975 ; Brakefield, 1979 ; Garcia- 
Barros, 1987). This suggests a relationship between female aestivation 
and the seasonal change in spot frequencies. A selective elimination 
of the high-spotted phenotypes may account for this pattern and would 
explain the homogeneous, low-spotted, female phenotypes found in 
September (as formerly proposed, e.g. Scali, 1972). It is thus interesting 
that a similar trend was discovered in the males, because no evidence 
for male aestivation has been published. The negative correlation be- 
tween spot number in males and date is not much lower than the 
one determined for the females. The male flight period is shorter than 
that of females (few are found in September), and this may contribute 
to obscure the relationship. On this basis, variables that may act in the 
two sexes — including seasonally determined phenotypic variation — 
might help in explaining seasonal spot variation in M. jurtina. 


None of the quantitative variables tested in this study was highly corre- 
lated with the spot number, even when the correlations were statistical- 
ly significant. This may be due, in part, to the heterogeneous condition 
of our sample, but also suggests an important amount of variation due 
to other factors. Assuming a relationship between aestivation and spot 
selection, some relationship between spotting and elevation might be 
expected (as observed in Italy, Scali & Masetti, 1975). The cooler 
mountain climate (and subsequently shorter aestivation period) should 
lead to higher spot frequencies. This is not supported by our data: 
elevation is correlated to adult size, but has no obvious relationship 
with spot patterns. 


Intra-seasonal spotting shifts and the female aestivating behaviour seem 
to be generalized features of the Mediterranean area, independently 
of the spot distribution. If the relationship between these two factors 
can be demonstrated, the evidence would suggest either convergent 
phenological adaptation at the population level, or fast genetic diver- 
gence after the stabilization of a “Mediterranean adapted” population. 
Further studies on the behaviour and phenology of Maniola jurtina 
in this area would probably contribute to our understanding of the 
precise mechanisms related to spot selection. 


162 


Acknowledgements 


We wish to thank the curators of the Departments of Entomology of the 
Madrid Natural History Museum (Dr. I. Izquierdo) and the Universidad 
Complutense (R. Outerelo) for kind assistance while working with their 
collections. 


References 


BENGTSON, S.-A., 1978. Spot distribution in Maniola jurtina (Lepidoptera) 
on small islands in southern Sweden. Holarct. Ecol. 1 : 54-61. 

BRAKEFIELD, P. M., 1979. An experimental study of the maintenance of 
variation in spot pattern in Maniola jurtina. Unpublished PH. D. Thesis, 
University of Liverpool. 

BRAKEFIELD, P. M., 1984. The ecological genetics of quantitative characters 
of Maniola jurtina and other butterflies. Jn Vane-Wright, R. I. & Ackery, 
P. R. (Eds.) : The Biology of Butterflies, Symp. R. Ent. Soc. No. 11: 
167-190. Academic Press, London. 

BROCKIE, R. E., 1972. Evolutionary studies on Maniola jurtina in Sicily. 
Heredity 28 : 337-345. 

DowDEsWwELL, W. H. & McWhirter, K. G., 1967. Stability of spot-distri- 
bution in Maniola jurtina throughout its range. Heredity 22 : 187-210. 

Forp, E. B., 1975. Ecological genetics. 3rd edition. London: Chapman & 
Hall. 

GARCIA-BARRoS, E., 1987. Observaciones sobre la biologia de Maniola jurtina 
(L., 1758) en el centro de la Peninsula Ibérica : fenologia general del 
ciclo biolögico, duraciön del periodo de prepuesta y fecundidad potencial 
de las hembras (Lep. Nymphalidae). Boletin Asoc.esp Entom. 11 : 235- 
247. 

MAsETTI, M. & Scauı, V., 1972. Ecological adjustments of the reproductive 
biology in Maniola jurtina from Tuscany. Rend. Accad. Naz. Lincei 
Rc., serie VII, vol. LIII : 460-470. 

MCWHIRTER, K. G., 1957. A further analysis of variability in Maniola jurtina 
L.. Heredity 11 :359-371. 

Owen, D. F, & SMITH, D. A. S., 1990. Interpopulation variation and selective 
predation in the meadow brown butterfly, Maniola jurtina (L.) (Lepi- 
doptera : Satyridae) in the Canary Islands. Biol. J. Linn. Soc. Lond. 
38) DS 26T. 

Owen, D. F. & SmitH, D. A. S., 1993. Spot variation in Maniola jurtina 
(L.) (Lepidoptera : Satyridae) in southern Portugal and comparison with 
the Canary Islands. Biol. J. Linn. Soc. Lond. 49 : 355-365. 

SCALI, V., 1972. Spot-distribution in Maniola jurtina : Tuscan Archipelago, 
1968-1970. Heredity 29 : 25-36. 


163 


SCALI, V. & Masetti, M., 1975. Variazioni intrastagionali dello spotting e 
selezione in Maniola jurtina L. (Lepidoptera, Satyridae). Rend. Accad. 
Naz. Lincei Rc., Serie VIII, vol. LVIII : 244-257. 

SHREEVE, T. G., Dennis, R. L. H. & WiirraMs, W. R., 1996. Uniformity 
of wing spotting of Maniola jurtina (L.) in relation to environmental 
heterogeneity (Lepidoptera : Satyrinae). Nota lepid. 18 (1) : 77-92. 

TuHomson, G., 1973. Geographical variation of Maniola jurtina (L.) (Lepi- 
doptera, Satyridae). Tijdschr. Ent. 116 : 185-227. 


164 


Nota lepid. 19 (3/4) : 165-184 ; 21.X11.1996 ISSN 0342-7536 


The butterfly assemblages of northeastern 
Spanish wetlands 


Constanti STEFANESCU*, Michael Lock woop** and Mariano CEBOLLA*** 


* Parc Natural dels Aiguamolls de ’Empordä, El Cortalet, E-17486 Castello d’Empuries, Spain. 
** Parc Natural del Delta del Llobregat, Gran Via, 612-614, 2n, E-08007 Barcelona, Spain. 
*** Parc Natural del Delta de l’Ebre, Pca 20 de maig, 2, E-43580 Deltebre, Spain. 


Summary 


An ecological assessment of the butterfly assemblages of the last three relatively 
large wetlands existing in the northeastern Iberian Peninsula was carried out. 
Data were collected in 1988-1995 using the Butterfly Monitoring Scheme 
methodology. Four monitoring transect routes were considered, one in the 
Emporda Marshes Natural Park, two in Llobregat Delta Nature Reserves 
and one in the Ebro Delta Natural Park. Nearly 45,000 butterflies belonging 
to 59 species were recorded, of which only 35 breed regularly in the study 
areas. Despite the different number of regular breeders in the four transect 
routes, there was a very close coincidence in the number of species which 
form open populations. Some of these species (e.g., Cynthia cardui, Vanessa 
atalanta, Pieris rapae, Pontia daplidice, Colias crocea) are common migrants 
in the region, and northward migration was observed in the spring months, 
especially in May and June. The difference in the species’ richness of the 
four areas was related to the different number of species with closed 
populations, which in the case of the Ebro Delta was exceedingly low. In 
comparison with other surrounding areas, the sites surveyed in this study are 
characterized by very poor butterfly faunas. Moreover, the extinction of at 
least four species was recorded in an eight years’ period in the Emporda 
Marshes Natural Park, the richest of the four areas. Habitat fragmentation 
following the adoption of modern agricultural practices is identified as the 
main factor leading to the progressive impoverishment in the butterfly 
populations. 


Resumen 


El presente estudio analiza, desde una perspectiva ecolögica, las comunidades 
de ropalöceros de las tres ultimas grandes marismas que se conservan en el 
noreste de la Peninsula Ibérica. Los datos se obtuvieron durante el periodo 
1988-95, aplicando la metodologia del Butterfly Monitoring Scheme en cuatro 
transectos distintos : uno en el Parque Natural de los Aiguamolls de l’Em- 


165 


pordà, dos en las Reservas Naturales del Delta del Llobregat y uno en el 
Parque Natural del Delta del Ebro. En total se contabilizaron unas 45,000 
mariposas pertenecientes a 59 especies, de las cuales 35 pueden considerarse 
reproductoras regulares. A pesar de que entre las cuatro areas existen marcadas 
diferencias en cuanto al numero de especies que se reproducen regularmente, 
se observa una gran coincidencia respecto al numero de mariposas con 
poblaciones de tipo abierto. Algunas de estas especies (e.g., Cynthia cardui, 
Vanessa atalanta, Pieris rapae, Pontia daplidice, Colias crocea) son migradoras 
comunes en el NE de la Peninsula Ibérica, siendo frecuentes las migraciones 
hacia el norte durante los meses de primavera, especialmente en mayo y junio. 
Las diferencias en la riqueza especifica se relacionan, por tanto, con el distinto 
numero de especies con poblaciones cerradas, que en el caso del Delta del 
Ebro es extraordinariamente bajo. En comparacion con otras areas cercanas, 
las localidades estudiadas presentan una fauna lepidopterolögica muy pobre. 
Hay que destacar, ademas, la extincidn de un minimo de cuatro especies 
en un periodo de ocho años en el Parque Natural de los Aiguamolls de ’Em- 
porda, el area mas diversa de las cuatro estudiadas. Se sugiere que este pro- 
gresivo empobrecimiento de las comunidades de mariposas es una consecuencia 
directa de la fragmentacion de los habitats, resultante de la implantaciön de 
las técnicas agricolas modernas. 


Introduction 


Largely as a result of the construction of infrastructure for the tourist 
industry, the Mediterranean coastline of the Iberian Peninsula has been 
dramatically transformed over the last few decades. In Catalonia (NE 
Spain) this process has been especially notable (Folch I Guillén, 1976) 
and today only a few areas of the Catalan coastline possess well- 
preserved natural ecosystems (Orta et al., 1992). Three of the most 
important of these remaining areas are the last relatively large wet- 
lands left in the northeastern Iberian Peninsula: the Ebro Delta, the 
Llobregat Delta and the Emporda marshes (Fig. 1). These three areas 
were declared Natural Parks in the 1980s due to their importance as 
sites for breeding and migrating birds. Recent studies have shown, 
moreover, that these areas are also of high interest for plants as well 
as for other animal groups than birds. 


The terrestrial invertebrate fauna of these wetlands has very special 
characteristics and a very limited range within Catalonia and, in general, 
within the Iberian Peninsula. This is the case for the distribution of 
some moth species (one of the best-studied groups) restricted to marsh 
and brackish environments (Pérez De-Gregorio, 1990 ; Stefanescu & 
Miralles, 1994). 


166 


The butterflies, on the other hand, are dominated by a small number 
of opportunistic species found throughout the Iberian Peninsula (e.g., 
Miralles & Stefanescu, 1994). Most of these species have been favoured 
by the transformation of the countryside and the replacement of natural 
ecosystems by cultivated and degraded areas. However, the study of 
these rather limited communities does provide the opportunity to 
analyze interesting and little-known aspects of the ecology of Medi- 
terranean butterflies. It is also important to take into account the fact 
that some species migrate along the Mediterranean coast and a long- 
term study would provide much information on the nature of these 
migrations. 


Furthermore, the different degrees of conservation of these three wet- 
lands furnishes an excellent opportunity to study the responses of 
butterfly populations to the destruction and transformation of the 
countryside, a process which has severely affected butterflies over the 
last few decades. 


The aim of this paper is to employ data collected in 1988-1995 using 
the British Butterfly Monitoring Scheme methodology, a standard 
technique for monitoring butterfly populations (Pollard & Yates, 1993), 
to analyze some of these issues. 


Material and methods 


Study sites 


Three distinct geographical areas were covered by this study : Emporda 
Marshes Natural Park (EMNP), Llobregat Delta Nature Reserves 
(LLDNR) and Ebro Delta Natural Park (EDNP) (Fig. 1). 


From a phytogeographic point of view, the three areas represent coastal 
wetlands (Riba er al., 1980; Orta et al., 1992) in which halophytic 
(Phragmition) and saltmarsh communities dominate. However, all of 
the areas are highly populated and contain large areas of agricultural 
land. 


The EMNP covers 4824 ha to the north and south of the Muga river. 
The northern section corresponds almost exactly to the area formerly 
occupied by the Castellö d’Empüries lagoon and consists of extensive 
reedbeds and well-preserved halophytic vegetation, as well as the re- 
mains of the fluvial woodland which once surrounded the lagoon. To 
the south there is a series of saline coastal lagoons lying between the 
mouths of the Fluvia and Muga rivers, the neighbouring beach and 
the interior Cortalet lagoon. The vegetation mainly consists of saltmarsh 


167 


BARCELONA 


Fig. 1. Map of the study area. 1: Empordà Marshes Natural Park ; 2: Llobregat 
Delta Nature Reserves ; 3 : Ebro Delta Natural Park. 


168 


“steppes” and reedbeds depending on their distance from the coast 
and the salinity levels of the soil. Pastures liable to flood (known locally 
as “closes”) and cultivated fields (principally maize, sunflowers and 
alfalfa) are common in both areas. 


The LLDNR covers 500 ha to the south of the Llobregat river. Its 
proximity to Barcelona and the Barcelona/ Baix Llobregat conurbation 
has meant that the original landscape has been radically altered. Today, 
only 5% of the area is covered by semi-natural vegetation, the remaining 
area being occupied by market gardening and orchards, built-up areas 
and industrial complexes. The fluvial woodland which once must have 
occupied a large area has been all but destroyed and only a very few 
elms and poplars still stand along the banks of the river and the 
irrigation channels. 


The EDNP covers a total of 7736 ha on both sides of the Ebro river 
and penetrates 22 km into the Mediterranean Sea. The transformation 
of the Delta as a result of demographic growth goes back as far as 
the 14th Century, although the Delta underwent greatest change in 
the 19th Century. During this century rice cultivation expanded in the 
marsh areas which until then occupied a large part of the Delta and 
which currently only occupy 20% of the total area. As a result of the 
development of market gardening and orchards, fluvial woodland is 
non-existent. 


Sampling methodology 


The sampling procedure was based on the British Butterfly Monitoring 
Scheme (BMS) methodology (Pollard, 1977), a scheme used in Britain 
since 1976 to monitor butterfly abundance. The monitoring method 
has been previously well described (e.g., Hall, 1981 ; Pollard & Yates, 
1993), and therefore only a brief description is given here. 


The BMS consists of visual counts of butterflies along a fixed route. 
The transect is walked once a week and only those butterflies which 
come within 5 m in front of the recorder are counted. At the end 
of the season, an annual index of abundance is calculated for each 
species. This index is the sum of the weekly counts, including the 
estimated figures for missing values. Although the annual index does 
not provide the absolute population size of a species in a site, a close 
correlation between this measure and population estimates obtained 
with MRR methods has been shown on many occasions (e.g., Pollard, 
1977 ; Thomas, 1983). 


169 


As most Spanish multi-brooded butterflies have overlapping flying 
periods, we have preferred to calculate a single annual index of 
abundance instead of a separate index for each generation, as is usually 
done in the British Butterfly Monitoring Scheme. 


In this study, sampling started on March Ist and ended on Sep- 
tember 26th, thus comprising a total of 30 weeks. Recording time was 
always restricted to the time between 10.00 am and 2.00 pm. When 
the temperature was lower than 15°C, transects were walked only if 
sunshine was higher than 75%. 


Apart from a global analysis of the areas comprised within the transect 
routes, there is also a section dedicated to the migration of some 
butterfly species in 1995. Most of the data have been taken from the 
results of the transects, although observations by the authors and other 
people that work in the field are also included. This additional 
information has been very valuable in the case of Danaus chrysippus, 
a species which was recorded mainly in the EDNP on the “Illa de 
Buda” a few kilometres from the transect area, and once in the LLDNR 
and EMNP within the transect route. 


Transect routes 


In total four transect routes have been considered : one each in EMNP 
and EDNP and two in LLDNR. All of them are included in the Butter- 
fly Monitoring Scheme which has been running in Catalonia since 1994, 
comprising 18 sites in 1995 (Stefanescu, 1994). Table 1 summarizes some 
characteristics of these transects. 


Table 1 


Some characteristics of the transect routes used in the present study. 
EMNP : Emporda Marshes Natural Park ; 
LLDNR : Llobregat Delta Nature Reserves ; EDNP : Ebro Delta Natural Park 


Transect Location Recording | Number Transect Distance 
route period of sections | length to the sea 
El Cortalet 16 =]. 

El Remolar 


Cal Tet 
La Marquesa 


In EMNP, the transect route (here designated as “el Cortalet”) is located 
around the Park information center, and mainly consists of a network 
of paths surrounded by fluvial woodland (elms, ash and oaks), hay 
meadows used for grazing, cultivated fields of sweet corn and sunflower, 
and ruderal habitats (see Miralles & Stefanescu (1994) for more details). 


170 


Two different transect routes were defined in LLDNR. The first route 
(designated as “el Remolar”) is located inside the Nature Reserve of 
“el Remolar-Filipines”. While reed beds and ruderal vegetation are 
extensive along all the transect, some other habitats (e.g., patches of 
pines mixed with dry meadows and some remnants of dune vegetation) 
are also well represented. The second transect (designated as “cal Tet”) 
is located inside the Nature Reserve of “la Ricarda-ca l’Arana” a small 
remnant of the original vegetation completely surrounded by irrigated 
fields. Ruderal vegetation is extensive along the entire transect, but 
meadows and paths through a patch of pine wood are also included. 


The transect route in EDNP (designated as “la Marquesa”) is located 
along the beach and is divided in two distinct areas, both dominated 
by salt-tolerant and ruderal plant species. In the first one (“platja de 
la Marquesa”) there are still good remnants of the dune system, while 
in the second one (“bassa de les Olles”) some patches of long turf 
grass mixed with Medicago marina are also sampled. 


The four transects are representative of the dominant environments 
found in each area. Only in the EMNP, the most heterogeneous area 
from a phytogeographical point of view, do patches of typically Medi- 
terranean vegetation exist, each with characteristic butterfly populations 
that are not represented in the transect routes. In broad terms, therefore, 
and with the exception mentioned above, the butterfly composition 
found in each site is very similar to that found in the whole of the 
protected areas. 


Results 
Butterfly assemblages at the different sites 


In the period 1988-1995, 44,848 butterflies were recorded belonging 
to 59 different species (Table 2). Comparisons of species richness are 
difficult given the different sampling sizes of each area, a factor directly 
related to the number of recording years and the length of each transect 
(Table 1). Logically, a larger sample size leads to more observations 
of vagrant species which, due to their greater mobility, disperse far 
from their breeding areas. The relatively large number of vagrant species 
which have been recorded in “el Cortalet” once or only a very few 
times during the recording period (e.g., Pieris mannii, Colias alfaca- 
riensis, Anthocharis cardamines, Charaxes jasius, Aglais urticae, Issoria 
lathonia, Hipparchia semele, etc.) is worth remarking on. Presumably 
in years to come the number of such species recorded in the other 
areas will also increase considerably. 


171 


A more balanced analysis can be carried out if only the species which 
breed regularly in each area are considered. In Table 2 these species 
are listed in accordance with the observations accumulated over the 
recording periods. “El Cortalet” (EMNP, 29 spp.) has the greatest 
diversity, followed by “el Remolar” (LLDNR, 27 spp.), “cal Tet” 
(LLDNR, 22 spp.) and “la Marquesa” (EDNP, 14 spp.). In the EMNP 
and the LLDNR these totals are probably accurate, whereas in the 
EDNP where only one year’s records are available, the total may be 
expected to change. 


In broad terms, the breeding species can be divided in two distinct 
groups according to their different population structures: species 
having closed populations (species that form well-defined colonies 
within discrete areas) and species having open populations (species that 
range more widely or regularly migrate from one breeding area to 
another) (Thomas, 1984 ; Warren, 1992). The species belonging to each 
of these groups are indicated in Table 2, according to Thomas (1984) 
and Warren (1992) and, for the butterflies not considered by these 
authors, according to distributional data of the species within the 
transect routes and unpublished data of their movement patterns. 


The proportion of the species within these two main groups is re- 
presented in Figure 2. Despite the different number of species breeding 


HO 7003cZ 


NO D—.0 DTA 


EI Cortalet El Remolar Cal Tet La Marquesa 


BE open INN closed 


Fig. 2. Number of regular breeding species with open and closed populations in the 
four study areas. 


172 


regularly (resident) in the four areas, there is a very close agreement 
in the number of species with open populations. The difference in 
number of resident species is due, therefore, to the number of species 
with closed populations, which in the case of the Ebro Delta is 
exceedingly low (cf. Table 2). 


Table 2 


Butterflies present at the four studied areas during the period 1988-95. 
(1): El Cortalet, EMNP (1988-95) ; (2): El Remolar-Filipines, LLDNR (1994-95) ; 
(3) : Cal Tet, LLDNR (1994-95) ; (4) : La Marquesa-Ses Olles, EDNP (1995). Symbols 
used : + : regular breeding species ; * : vagrant species ; e : extinct species during the 
recording period ; —: species not detected at the study site; 0: species with open 
populations ; c: species with closed populations. Systematic order and nomenclature 
follow Higgins & Riley (1980). 


(1) (2) (3) (4) population 
structure 


PAPILIONIDAE 
Papilio machaon + 
Iphiclides feisthamelii - 


PIERIDAE 

Pieris brassicae 

P. rapae 

P mannii 

P. napi 

Pontia daplidice 
Euchloe crameri 
Anthocharis cardamines 
A. euphenoides 
Colias crocea 

C. alfacariensis 
Gonepteryx rhamni 
G. cleopatra 
Leptidea sinapis 


LYCAENIDAE 
Strymonidia spini 

S. w-album 
Callophrys rubi 
Lycaena phlaeas 
Lampides boeticus 
Syntarucus pirithous 
Cacyreus marshalli 
Everes argiades 
Celastrina argiolus 
Plebejus argus 
Aricia cramera 
Lysandra hispana 
Polyommatus icarus 


“+ 
Mi 
IL 
© 


| 
| 
fo) 


"+++ ett 
+++ ++ 
+++ ++ 


ME les cn or 
+ | 
+ | 


* | 
| 
| 


GoOO00009000C000 


OR x +] 


o 
~ 
| 
| 
| 


| pet Tee 
oes À 
| + * | 


+*+++o1+4++1 + 


173 


NYMPHALIDAE 
(Nymphalinae) 
Charaxes jasius 
Nymphalis antiopa 
Inachis io 

Vanessa atalanta 
Cynthia cardui 
Aglais urticae 
Polygonia c-album 
Issoria lathonia 
Melitaea phoebe 
M. didyma 


(Satyrinae) 

Melanargia lachesis 
Hipparchia semele 
Neohipparchia statilinus 
Brintesia circe 

Maniola jurtina 
Pyronia tithonus 

P cecilia 

P bathseba 
Coenonympha pamphilus 
Pararge aegeria 
Lasiommata megera 


(Danainae) 
Danaus chrysippus 


HESPERIIDAE 
Pyrgus malvoides 

P armoricanus 
Spialia sertorius 
Carcharodus alceae 
C. boeticus 
Thymelicus acteon 

T. lineola 

Ochlodes venatus 
Gegenes nostrodamus 


Total individuals 
Total species 


Resident species 
Vagrant species 
Extinct species 


TN 
* — 
Nr 


** «+ 44 x) 


x + 


# +++ *+4+00% 


bt ettt ett 


ne nai deuil 


++i +1 + ei + 


(3) 


(4) 


population 
structure 


Qa 0 
er 210 OO Oo © Clore 


© @ GEGEOZOROTOERZOEAHT 


OD OR Oi OMe © i © Pk @) 


Logically, the most abundant species in all of the sites are also resident 
butterflies. Although these experience annual fluctuations in numbers 
and, therefore, changes in the dominance between the recording years 
(Miralles & Stefanescu, 1994), each site can be characterized by a subset 
of its most common butterfly species. In Table 3 a rank list of the five 
butterfly species with the highest annual indices in 1995 is given. These 


174 


Table 3 


The most abundant species in each of the sample sites, 
listed according to their annual indices in 1995. 
The percentage of the species’ annual index with respect to the sum 
of the annual index of all species is given in brackets 


El Cortalet La Marquesa 


P. aegeria 832.5 (20.6) P. rapae 335.0 (29.0) 
P. rapae 686.5 (17.0) C. cardui 287.5 (24.9) 
P. napi 678.5 (16.8) P. daplidice 212.0 (18.4) 
P. icarus 604.0 (14.9) C. crocea 115.0 (10.0) 
M. lachesis 181.5 (4.5) P. icarus 64.5 (5.6) 


El Remolar Cal Tet 


P. rapae 1456.0 (37.2) P. rapae 1627:.02@19) 
P. cecilia Te) (ID) P. napi 1196.0 (23.5) 
C. alceae 358.0 (9.1) P. aegeria 413.0 (8.1) 
P. napi 336.5 (8.6) L. megera 355.0 (7.0) 
P. icarus 160.0 (4.1) C. cardui 345.0 (6.8) 


species have the highest population sizes at each site and their annual 
indices account for nearly 75% or even more of the sum of the annual 
indices of all species (73.8% in EMNP, 77% in both transect routes 
in LLDNR, and 87.8% in EDNP). 


Extinctions 


In “el Cortalet”, where data have been gathered for over eight years, 
the extinction of some species has been recorded. We consider that 
a population has become extinct when no individuals were seen in 
two successive recording years, after a period with two or more years 
with regular records. Although the criteria established by Pollard & 
Yates (1992) are more conservative, the patterns of annual fluctuations 
of the species detailed below undoubtedly indicate that local populations 
have disappeared. 


During the recording period, the breeding populations of at least four 
species are assumed to have become extinct (Table 2) : Leptidea sinapis, 
Everes argiades, Brintesia circe and Maniola jurtina. Two other species, 
Nordmannia spini and Neohipparchia statilinus may also have become 
extinct during this period, although the presence of breeding populations 
when they were recorded in the transect has not been proved. For 
all these butterflies populations showed declining numbers that even- 
tually led to their extinction. This is especially true for B. circe and 
M. jurtina, two species which became gradually scarcer after an increase 
in 1989, and disappeared completely from the area in 1994 (Fig. 3). 


175 


Leptidea sinapis Everes argiades 


15 
10 à : 
6 see 
0 ? — — 
88 89 90 91 92 93 9 


4 95 


X®0Q3— —nc33» 
X92Q3— —-VC33> 


Brintesia circe Maniola jurtina 


X0Q3— —-»c.33> 
X0Q3— -»E233> 


Fig. 3. Extinctions of butterfly species recorded in “el Cortalet” (EMNP) during the 
period 1988-95. No data were available in 1990. 


Everes argiades colonized in small numbers one section of the transect 
in 1991, but after an increase in 1993 it disappeared in the following 
year. Leptidea sinapis was recorded in the first two years in small 
numbers, and afterwards disappeared from the area. 


Migration 


An important proportion of the species that regularly breed in the 
four areas are common migrants over their entire distribution range. 
As defined by Baker (1969) and Shreeve (1992), a species can be 
considered as a migrant when a part of its population is involved in 
predictable seasonal and directional movements. This type of behaviour 
is well exemplified by multi-brooded species such as Cynthia cardui, 
Vanessa atalanta, Pieris rapae, Colias crocea and Pontia daplidice, 
which show directional flights northward in spring and summer, and 
southward in late summer and autumn (Williams, 1951 ; Baker, 1969). 


In 1995 the migration of some species was detected in the four sites. 
Table 4 summarizes the dates of the main migrations observed during 
the recording period (March Ist-September 26th). Other minor mi- 
grations were observed more or less regularly throughout the spring 
months. 


176 


Unlike the normal pattern of weekly counts observed for the generation 
of a butterfly with a typically closed population (a steady rise to a 
peak during the brood’s emergence, followed by a decline as individuals 
die), the data of Table 4 refer to abrupt changes in abundance that 
can only be explained by the mass arrival of migrants. In most cases 
the butterflies were recorded with a clear directional flight towards the 
north, indicating that a migratory process was actually taking place. 


Migration events were observed for all the species mentioned above, 
and were especially important in the EDNP. In this area the arrival 
of migrants is not only much more frequent, but also more intense 
than in the rest of the sites. Moreover, in some cases there were mixed 
butterfly migrations, with several species involved. The most evident 
was observed on 16th June, when high counts of P rapae, C. cardui, 
C. crocea and, to a lesser extent, V. atalanta, were recorded (Fig. 4). 
Other mixed migrations were detected in LLDNR on 27th May and 
in EMNP on 2nd June (Table 4). 


Unfortunately the BMS recording period ends on 26th September, and 
therefore most autumn migrations such as those described by Lack 
& Lack (1951), Williams (1951), Baker (1969) and Templado (1977) 
are lacking from our study. However, it is interesting to note one 


Pieris rapae Colias crocea 


+ <i 


œ 
Q 


e-3c.00 <-zee$ 
a 
[=] 
.-23c.90 <-zoe$ 


LS) 
Le] 


6 ; 
0 || N BR 0 „hl LE 
March April May June July Aug Bept — March April May June July Aug Bept 


Vanessa atalanta Cynthia cardui 


g— 


60 3 


30 : 


Lan t : ill | 
May June July Aug Bep March April May June July Aug Bept 


Fig. 4. Weekly counts of four migrants in “la Marquesa” (EDNP), in 1995. A dot 
indicates no count in that week. Note the mixed migration that took place on 16th June 
(shown by an arrow). 


u-3c00 <-zee$ 
a>3c00n <—-xe0F 


March April t 


177 


Table 4 


Dates of the main migrations detected in 1995 at the four study areas 
during the recording period (March Ist-September 26th). 
In brackets, the number of butterflies recorded during the transect walks. 
ND : No data available on transect counts 


El Cortalet El Remolar Cal Tet La Marquesa 


Pieris rapae 16.VI (73) 
Pontia daplidice 2.VI (4) 6.VII (44) 
Colias crocea ZEN (IV) 30.IV (12) 

16.VI (29) 


Vanessa atalanta 22.1V (ND) 
16.VI (6) 

Cynthia cardui ZNO) 13.V (11) 27.V (124) 22.1V (ND) 
14.V (28) 
8.VI (47) 
16.VI (65) 


migration that was observed at “el Remolar” on 28th September, when 
34 Pieris brassicae and small numbers of C. crocea and V. atalanta 
were seen flying southward in a straight line above the beach for a 
period of one hour. 


The EDNP was the only area where Danaus chrysippus, an African 
butterfly that periodically reaches the Iberian coast (e.g. Masö, 1992), 
was observed during the monitoring in 1995. That year, however, the 
migration was not as large as recorded in previous years, and almost 
all of the sightings were restricted to the area known as “Illa de Buda” 
a few kilometres from the monitored areas. 


The first example was seen on 25th June, but it was not until 25th July 
that three more butterflies were recorded. On this latter occasion egg- 
laying was observed on Cynanchum acutum, one of the previously 
recorded feeding plants of the larvae in the northeast of the Iberian 
Peninsula (Maso, 1992). Afterwards, many eggs, larvae and pupae were 
found on C. acutum, and adults were seen regularly until the end of 
October. One adult was observed in “el Remolar” on 16th October, 
possibly a migrant from the Ebro Delta population. Another one was 
seen in “el Cortalet” on 30th October. 


Discussion 


The Mediterranean region is characterized by a high biodiversity in 
comparison with other areas in central and northern Europe (e.g., 
Mooney, 1988). This pattern of diversity is also very pronounced in 
the case of butterflies, as has recently been shown by Dennis & Williams 
(1995) and Munguira (1995). Within the Mediterranean region, the 


178 


Iberian Peninsula, with more than 200 species (Gömez-Bustillo & 
Fernandez-Rubio, 1974 ; Fernandez-Rubio, 1991), harbours one of the 
most diverse butterfly faunas. In this area, Martin & Gurrea (1990) 
locate the highest levels of species richness in the northeastern region, 
and explain this pattern as a consequence of a peninsular effect affecting 
the distribution of butterflies, as well as other organisms. 


The sites surveyed in this study are within the richest region of the 
Iberian Peninsula (cf. Martin & Gurrea, 1990). However, they are 
characterized by a very poor butterfly fauna (Table 2), contrasting with 
that present in surrounding areas (Stefanescu, 1994). This low species 
richness can be explained, in part, by the location of the study sites 
at sea-level, taking into account that altitude has been suggested as 
the second most important factor affecting the distribution of Iberian 
butterflies, being positively correlated with the number of species found 
in an area (Martin & Gurrea, 1990). 


Two other main reasons for this abnormally low species richness can 
also be advocated. First, the low observed butterfly diversity is a 
response to the reduced diversity of the vegetation communities of 
wetlands (reed beds and saltmarshes which still cover large parts of 
the areas), as has been previously noted in the case of moths (Stefanescu 
& Miralles, 1994). Moreover, the impoverishment of the butterfly fauna 
is highlighted by the fact that there are actually very few Iberian 
butterflies adapted to exploit true wetland plants. 


Secondly, modern agricultural practices have resulted in the destruction 
of a great part of the formerly widespread and richest butterfly habitats 
(cf. Feber & Smith, 1995). A good example is provided by the events 
that have taken place in the EMNP (Vaqué et al., 1989). Nearly all of 
the present protected area was occupied by extensive marshlands a 
few centuries ago. In the 17th and 18th Centuries an intensive process 
of drainage and desiccation of the wetlands began, and vast portions 
of the land were converted into grazing meadows and cultivated fields. 
The new conditions created suitable habitats for butterflies, in particular 
the locally named “closes” that is, highly diversified meadows enclosed 
by tracts of riverine forest (mainly ash, oaks, elms and, in saline environ- 
ments, tamarisks) used for hay and cattle grazing. Not only are these 
meadows rich in both larval foodplants and abundant nectar sources 
(e.g., Gramineae, Leguminosae and Compositae) for typical grassland 
species, but they also provide a system of glades and rides that favour 
woodland butterflies. 


The present century (and especially over recent decades) has resulted 
in a further transformation of the landscape, leading to important 


199 


changes in land-use and the loss of major butterfly habitats. An in- 
creasing number of “closes” have been ploughed and transformed into 
cultivated fields of corn, lucerne and sunflower, while others have been 
abandoned. Consequently only a few good butterfly habitats remain 
today inside the park, and these sites are becoming more and more 
isolated. Most of the best conserved “closes” have become virtual is- 
lands within vast extensions of arable land, intensively “improved” with 
the use of herbicides and fertilizers. | 


In the EDNP and LLDNR the situation is similar to that described 
above for the EMNP, and historical events have also led to a significant 
impoverishment in the butterfly fauna. In both areas, however, the 
present situation is even worse and is the consequence of extreme 
habitat loss. No riverine woodland remains in either area and the 
intensification of agriculture has been more severe (employing very high 
levels of herbicides, insecticides and fertilizers). 


Many authors have recently argued that habitat fragmentation and 
isolation is acting as one of the major causes of the decline of European 
butterflies in the second half of the century (e.g., Kudrna, 1986; 
Thomas, 1984, 1991 ; Warren, 1992). Moreover, the way in which this 
process is taking place has been satisfactorily explained on theoretical 
grounds for an increasing number of species following the development 
of metapopulation dynamics theory (Thomas er al., 1992 ; Hanski & 
Thomas, 1994). 


It is to be expected in degraded areas and in areas with good but 
highly fragmented habitats that butterfly communities consist principally 
of very mobile species, and that the proportion of sedentary species 
with closed populations is increasingly smaller. This is the current 
situation in the remaining three wetlands in the northeastern Iberian 
Peninsula, where there has been a recent and relatively rapid decline 
in several species with closed populations (Fig. 3). The more extreme 
example is provided by the EDNP, where the loss of habitats has been 
so severe that it has led to the disappearance of almost all the sedentary 
species with closed populations (Fig. 2). The few which still resist hang 
on in small numbers, in no way comparable to the numbers of species 
with open populations (Table 3). 


Moreover, it is important to note that the majority of resident species 
with closed populations (Table 2) are capable of exploiting resources 
available in disturbed and early seral habitats. This is the case of ubi- 
quitous species such as Polyommatus icarus, Lycaena phlaeas, Pararge 
aegeria, Lasiommata megera and Carcharodus alceae (the last mainly 


180 


restricted to the Mediterranean region), which can breed in almost all 
habitat types (Dennis, 1992) and in the area have a multivoltine strategy 
with at least three complete broods per year. Although a closed popu- 
lation structure 1s currently accepted for these species, there is no doubt 
that they have good dispersal abilities, which enable them to colonize 
other areas easily (e.g., Shreeve, 1995). 


In fact, the distinction of open and closed populations is an oversim- 
plification of the reality, as noted by Thomas (1995) who argues that 
these two categories could be replaced with a continuum of mobility. 
For instance, the “closed” category may represent those species showing 
a Levin’s type metapopulation structure (Hanski & Kuussaari, 1995), 
while the “open” category would be applicable to those species with 
large “patchy population” structures. Even the placement of a species 
within one of these two groups may be viewed as something variable 
in time, as factors affecting mobility (e.g., those associated with the 
habitat, with population parameters and with environmental factors 
which influence individual activity) are in permanent change (Shreeve, 
1995). However, the dichotomy of open vs. closed populations can 
still be useful as it exemplifies two contrasting structures in the popu- 
lations of most butterflies (Thomas, 1995) : those species in which local 
abundance is mainly determined by immigration and emigration, and 
those species in which local birth and death processes are the major 
determinants of the population dynamics. 


The butterfly communities today of the three last wetlands remaining 
in the northeastern Iberian Peninsula are composed, thus, of a few 
number of species characterized by their great adaptability in a changing 
environment that has lost many of its former suitable habitats for these 
insects. Although different ecological characteristics (e.g., whether 
populations are “closed” or “open”/ migratory, the stage at which hiber- 
nation occurs, the type of larval food plants, etc.) can be found in 
these most successful species, all of them conform to the concept of 
“butterflies of the future” expressed by Bink (1992). As observed in 
other European countries, the distribution of these species has not 
declined in the second half of this century and some have even expanded 
their ranges and become more abundant (van Swaay, 1990; Pollard 
et al., 1995). 


In spite of the absence of almost any rare butterfly in these areas, 
continuous monitoring will provide abundant and interesting informa- 
tion on different aspects of the ecology of many species (e.g., phenology, 
migration, population trends, etc.), an important achievement given 
the lack of reliable data being collected in the Mediterranean region. 


181 


Acknowledgements 


The Butterfly Monitoring Scheme in Catalonia is funded by the “Departament 
de Medi Ambient de la Generalitat de Catalunya”. Empordà Marshes Natural 
Park and Llobregat Delta Natural Reserves (“Departament d’Agricultura, 
Ramaderia 1 Pesca de la Generalitat de Catalunya”) have also given financial 
support to this project. Thanks are due to Marta Miralles, Bernat Garrigés, 
Ricard Gutiérrez, Javier Santaeufemia, Paco Cerda, Xesca Palmer and Jordi 
Figuerola, for their help in the field work. An early draft of the manuscript 
was kindly read and criticised by Dr. M. Nieminen and Dr. E. Pollard. Drs 
R.L.H. Dennis and A. Erhardt also provided helpful comments on the manu- 
script. Susan Watt carefully revised the English version. 


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184 


Nota lepid. 19 (3/4) : 185-211 ; 21.XII.1996 ISSN 0342-7536 


The butterflies of the Greek island of Kos : a synthesis 
(Lepidoptera : Hesperioidea & Papilionoidea) 


Alain OLIviER* & Roland DE Prıns** 


* Luitenant Lippenslaan 43 B 14, B-2140 Antwerpen, Belgium 
** Liersesteenweg 437, B-2800 Mechelen, Belgium 


Summary 


The geography and vegetation of the Greek island of Kös are briefly described, 
followed by a review on the history of its lepidopterological investigation. 

In the Systematic part, all previous literature records as well as numerous new 
field data gathered by both authors are dealt with. 46 species are known at 
present from the island, one of which — Gonepteryx farinosa farinosa (Zeller, 
1847) — is entirely new. Danaus chrysippus chrysippus (Linnaeus, 1758) is 
again reported from Kös: this is the first record for Kos and indeed for 
the Eastern Aegean Islands in over 65 years. Since, the species has been found 
on Rodos (Rhodes) as well. A bilateral gynandromorph of Pontia (daplidice) 
edusa (Fabricius, 1777) is also mentioned. 

The authors believe that at present over 90% of the actual butterfly fauna of 
Kos is known. No single species has differentiated significantly, not even sub- 
specifically, pointing to a very recent isolation of the island’s fauna (<< 9.000 B.P.). 
Vicariance of a biota that was still continuous in the recent past, Kos being 
part of the Anatolian mainland, largely explains the present configuration 
of its butterfly fauna, while present-day dispersal of migratory taxa is of more 
marginal significance to its make-up. 


Résumé 


L’article commence par un bref exposé sur la géographie et la végétation de l’île 
grecque de Kös, suivi d’un aperçu de l’histoire de son exploration lépidopté- 
rique. 

Dans la partie systématique sont traitées toutes les données précédemment 
citées dans la littérature, ainsi que les nombreuses nouvelles mentions résultant 
des activités sur le terrain des deux présents auteurs. Actuellement, on recense 
46 espèces sur l’île, dont une — Gonepteryx farinosa farinosa (Zeller, 1847) — 
est entièrement nouvelle. Danaus chrysippus chrysippus (Linnaeus, 1758) a 
également été retrouvée récemment a Kos: il s’agit de la première mention 
depuis plus de 65 ans pour Kös, et même pour les îles égéennes orientales. 
Depuis, l'espèce a également été retrouvée à Rodos (Rhodes). Un gynandro- 


185 


morphe bilatéral de Pontia (daplidice) edusa (Fabricius, 1777) est également 
mentionné. 

Les auteurs pensent que plus de 90% de la faune des papillons diurnes de Kos 
est connue à l’heure actuelle. Aucune espèce ne montre une différenciation 
significative, même au niveau subspécifique, probablement à cause d’un isole- 
ment très récent de la faune de l’île (< 9.000 ans). 

Une fragmentation récente d’un biota qui était continu dans un passé proche, 
lorsque Kos faisait toujours partie de la masse continentale anatolienne, 
explique en grande partie la présente configuration de sa faune lépidoptérique, 
alors que l’apport d’espèces migratrices contemporaines n’influe celle-ci que 
marginalement. 


Introduction 


The Greek island of Kos has an area of 286 km?, is about 43 km 
long and 8-10 km wide, stretching from NE to SW. It lies only at 
about 3,5 km off the SW Turkish coast, facing the Bodrum Peninsula 
(province of Mugla). The nearest neighbouring islands are Psérimos 
and Kälimnos to the north, and Giali and Nissiros to the south (Fig. 1). 
The total length of the coast of Kös amounts to approximately 112 km. 


The island can roughly be divided into four areas as far as the landscape 
is concerned : 


1) the lowland along the northeastern and northern coast, from Akron 
Fokas until about Marmari. 

2) the Dikeos mountain chain, stretching over about 17 km, with the 
highest summit at 846 m. To the south it falls abruptly to the 
southern coast of the island. 

3) the plateau, reaching an altitude of up to 160 m around the town 
of Andimähia, from Pili in the NE to the small isthmus of Agios 
Stéfanos in the SW. 

4) the mountain chain on the Kéfalos Peninsula, culminating at an 
altitude of about 428 m on Oros Lätra. 


The lowland consists mainly of fertile cultivated land, though there 
are several botanically interesting localities, principally the extensive 
sandy plains along the northern coast, especially near Akron Psalidi, 
Akron Skandari, Lambi and Tingäki. Further the coastal meadows 
and lagunes, especially near Bellecitte and Tingaki and a truly wet, 
though strongly cultivated area to the west of Kös Town, with many 
meadows, ditches, rivulets, etc., that hauses a series of marsh and 
aquatic plants, that otherwise are quite rarely present on the Eastern 
Aegean Islands. This area is, moreover, strongly grown with trees and 


186 


bushes in field areas and, seen from a distance, it sometimes seemingly 
approaches a true forest area. Due to a steady rise of tourism and 
intensified irrigation for agricultural purposes, large areas of lowland 


dry out, with detrimental effects on the botanically interesting parts. 


The Dikeos mountain chain is, on its northern side, only grown 
moderately with trees and bushes in the higher, steep parts (Olea, 
Quercus, Cupressus, Pinus halepensis, Pistacia), but mostly cultivated 


Fig. 1. Geographic situation of the island of Kos in the SE Aegean. 


Bodrum Peninsula 
Resadiye Yarımadası 
Daraçya Yarimadasi 
Kalimnos 

Télendos 

Psérimos 

Astipalea 

Giali 

Nissiros 


Anhwn-OnAw 


Tilos 

Simi 
Rodos 
Alimnia 
Halki 
Saria 
Karpathos 
Kassos 


187 


or covered with meadowland. The geological structure of its ridge is 
quite varied, with several sorts of limestone, white marble and areas 
of volcanic stone. 


The plateau is composed of marl and seasand, and over wide stretches 
it is overlaid by volcanic pumice tuffs, being much deteriorated in many 
places by repeated erosion caused by rains. The newly built roads and 
the construction of the airport, that have devastated the vegetation, 
show these phenomena very clearly. Large areas in this part are not 
cultivated anymore, just being grazed at best. At the northern coast 
near Mastihäri, there is a considerable area of drift sand with imposing 
dunes, that are often grown with old Juniperus oxycedrus ssp. macro- 
carpa bushes. 


The Kéfalos Peninsula is of volcanic origin. It is very dry and mostly 
covered with a typical phrygana-vegetation. The old volcanic cone of 
Oros Lätra lies in the south and is sparsely covered with Pinus hale- 
pensis forest. Nearly all the inhabitants of the peninsula live in the 
villages of Kéfalos and Kamari. Detailed accounts on the geology and 
flora of the island can be found in resp. Jacobshagen (1986) and Hansen 
(1980). 


The climate of the island can be described as arid Mediterranean, with 
a relatively short, cool, wet winter, followed by a long, hot and dry 
summer. 


History of the lepidopterological investigation of Kos 


The island of Kös was under Italian administration from 1912 to 1945: 
therefore, it is not surprising that the first-ever butterfly records for 
this island were established by an Italian. Professor Alessandro Ghigi 
did some collecting on the islands of the Dodekänissa, including Kös, 
in August 1926. From the latter island he recorded 13 species, that 
are treated in Turati (1929) and a few of these also by Ghigi (1929) 
himself. 


On 6-8.VI.1935, the Austrian herpetologist Dr Otto Wettstein-Westers- 
heim collected several butterflies on the island, around the town of 
Kardämena and on Oros Dikeos. 18 species were found, of which 12 
newly. These data are dealt with by Rebel (1936). 


Hartig (1940) lists several species, based on previous records in Turati 
(1929) and Rebel (1936), but adds one additional taxon, “Satyrus semele 
L. cretica Rebel” [sic ! recte Hipparchia senthes (Fruhstorfer, 1908), 
see below]. 


188 


Thomson (1985) reports 19 species from Kos, collected on 30.V.1983 
at Kös Town, 3 of which are new. A fourth species, Cupido minimus 
(Fuessly, 1775), has not been retained (see below). 


Kudrna (1986) mentions one species as new for Kos, that has not been 
listed in any previous publication. 


Olivier (1986) summarizes existing knowledge on the butterflies of the 
island of Kos, publishing a checklist and dealing with own records 
for 19 species, 8 of which are new to the island. 


Olivier (1987) lists 15 species, adding a series of new faunistic data 
for the island, but none of the species mentioned is new. 


Finally, Olivier (1993a) notes 45 species from Kos, 7 of these for the 
first time. 


Further records can be found in Rebel (1937, 1939), Bernardi (1961, 
1971), Bretherton (1966), Tauber & Tauber (1968), Koutsaftikis (1973), 
Thomson (1987, 1990), van Oorschot, van den Brink & van Oorschot 
(1987), van Oorschot & van den Brink (1992), Olivier (1990, 1991, 
1993b, 1996), Olivier & Coutsis (1993, 1995) and Hesselbarth, van 
Oorschot & Wagener (1995). 


The senior author of the present contribution visited Kos on four 
occasions, viz. 10 & 11.1V.1986, 25-29.V.1988, 1, 4, 8 & 16-20. VI.1992 
and 2-4.1V.1995. The junior author visited the island once in autumn, 
between 17 and 28.1X.1992. Together, we observed 44 species, one of 
which is entirely new to the island, while another one is reported again 
for the first time in over 65 years ! This brings the new total for Kos 
to 46 species. Only two taxa recorded in the literature have not been 
found by us. 


Systematic part 


Here we list both all previously published records about the butterflies 
of Kos that we could trace in literature (L) as well as our own new 
data (N) : together, 24 localities are involved (see Fig. 2) and coverage 
extends over each single month from early April until the very end 
of September. 


Nomenclature largely follows Olivier (1993a) and Hesselbarth, van Oor- 
schot & Wagener (1995), except when stated otherwise. Data on ecology 
and phenology can be found in those two works and these will not 
be repeated here. 


189 


Fig. 2. Map of localities listed in the Systematic part. 


1 Agios Stéfanos 13 Oros Dikeos (400-846 m) (“Mt. Dikeo”) 
2 3 km SW Andimähia (50 m) 14 1 km W Agios Dimitrios (300 m) 
3 Kardämena (“Cardamena”) 15 Agios Dimitrios (250-300 m) 
4 Mastihari (0 m) 16 2km NE Agios Dimitrios (150 m) 
5 4kmE Andimahia (0-50 m) 17 7 km W Kos (0 m) 
6 Pili (50-100 m) 18 5 km W Kös (0 m) 
7 Marmari (0 m) 19 Kos (0-50 m) (“Kos Town”) 
8 Tingaki 20 Paradisi (0-50 m) 
9 Zipari (0 m) 21 2km E Paradisi (0-50 m) 

10 2kmN Asfendiou (100 m) 22 AkmEKoös 

11 Asfendioü 23 1 km N Akron Fokäs (0-50 m) 


12 Zia (300-450 m) (“above Zia”) 24 between Akron Fokas and Empros Therma 


Thymelicus sylvestris syriacus (Tutt, [1905]) 
L — previously reported by Olivier (1993a : 198 [Table 12]) and Hes- 
selbarth, van Oorschot & Wagener (1995 : 1110 [Table 27]) 
N — 3 km SW Andimähia (50 m), 25.V.1988 
— 2 km N Asfendioü (100 m), 26.V.1988 
— Paradisi (0-50 m), 26.V.1988 
— Zipari (0 m), 27.V.1988 
— Oros Dikeos (400-800 m), 28.V.1988 (obs.) 
— Mastihari (0 m), 29.V.1988 


Thymelicus acteon acteon (Rottemburg, 1775) 
L — “Ein kleines, dunkles Pärchen von Kos, Cardamena, 6.VI. 
[1935].” (Rebel, 1936 : 27) 


190 


— “Kos Town. Very common. [30.V.1983].” (Thomson, 1985 : 157) 

— “Tigäki [Tingaki], 26.V.1979, 2 4 (C [W. Curispisn]).” (Olivier, 
1987 : 79) 

— further reported by Olivier (1986: 113; 1993a: 36, 198 [Ta- 
ble 12]) and Hesselbarth, van Oorschot & Wagener (1995 : 1110 
[Table 27]) 

N — Paradisi (0-50 m), 26.V.1988 

— 2 km N Asfendiou (100 m), 27.V.1988 

— Zipari (0 m), 27.V.1988 

— Mastihari (0 m), 29.V.1988 

— 5 km W Kos (0 m), 16.V1.1992 


Gegenes pumilio pumilio (Hoffmansegg, 1804) 
L — previously reported by Olivier (1993a : 38-39, 45, 198 [Table 12]) 
N — Zipari (0 m), 27.V.1988 

— Kos (0-50 m), 20.1X.1992 

— Marmari (0 m), 22.1X.1992, 28.1X.1992 


Carcharodus alceae alceae (Esper, | 1780]) 
L — “Kos, Cardamena, 6.VI[1935], ein sehr großes © (Vorderflügel- 
lange 16 mm)” (Rebel, 1936 : 27) 
— “west of Kos Town. Locally common. [30.V.1983].” (Thomson, 
1985 : 157) 
— “above Zia, 6.VII.1978, 1 © (K [M.C. & G. KRUSEMAN])” (Oli- 
vier, 1987 : 79) 
— further reported by Olivier (1986: 113; 1993a : 45, 198 [Ta- 
ble 12]) and Hesselbarth, van Oorschot & Wagener (1995 : 1110 
[Table 27]) 
N — Zipari (0 m), 27.V.1988, 18.VI.1992 
— Mastihäri (0 m), 29.V.1988 
— Kos (0-50 m), 20.IX.1992 
— Marmari (0 m), 17.1X.1992, 19.IX.1992, 22.1X.1992 


Carcharodus orientalis Reverdin, 1913 
L — previously reported by Olivier (1993a : 43, 192, 198 [Table 12]; 
1993b : 21) and Hesselbarth, van Oorschot & Wagener (1995 : 
1110 [Table 271) 
N — Zia (300 m), 28.V.1988 
— Zià (450 m), 4.V1.1992 
— Zipari (0 m), 1.V1.1992 
— Asfendiou (300 m), 16.V1.1992 


191 


Carcharodus stauderi ambiguus (Verity, 1925) 

L — previously reported by Olivier (1993a : 43-45, 192, 198 [Table 121) 
and Hesselbarth, van Oorschot & Wagener (1995 : 1110 [Ta- 
ble 271) 

N — Zipäri (0 m), 27.V.1988 


Spialia (sertorius) orbifer orbifer (Hübner, [ 18231) 

L — previously reported by Olivier (1993a : 198 [Table 12]) and Hes- 
selbarth, van Oorschot & Wagener (1995 : 1110 [Table 271) 

N — Mastihäri (0 m), 29.V.1988 


Zerynthia cerisy cerisy (Godart, [ 18241) 

L — “very few specimens at A [1 km W Agios Dimitrios (300 m), 
10.1V.1986, 11.1V.1986] and C [2 km N Asfendiou (100 m), 
11.1V.1986], abundant at B [7 km W Kos (0m), 11.IV.1986] 
(...)” (Olivier, 1986 : 111-113) 

— “2 km N. of Asfendiou (100 m), 11.1V.1986, 1 & (very worn) 
(O[LIVIER |) 
7 km W. of Kos, 11.1V.1986, 3 & (very worn) (O[LIvVIER |)” (Oli- 
vier, 1987 : 80) 

— further reported by Olivier (1990 : 171 ; 1991 : 73 ; 1993a : 198 
[Table 12]) and Hesselbarth, van Oorschot & Wagener (1995 : 
1110 [Table 27]) 

N — Agios Dimitrios (300 m), 2.IV.1995 
— 2km E Paradisi (0-50 m), 3.1V.1995, 4.1V.1995 
— Zià (450 m), 4.1V.1995 


Archon apollinus apollinus (Herbst, 1798) 
L — “H. PIepPer (pers. comm.) found the species in the island of Kos” 
(Kudrna, 1986 : 91) 
— further reported by Olivier (1993a : 47, 198 [Table 12]) and Hes- 
selbarth, van Oorschot & Wagener (1995 : 261, 1110 [Table 27]) 


Iphiclides podalirius podalirius (Linnaeus, 1758) 

L — “B[7 km W Kos (0 m), 11.1V.1986] and C [2 km N Asfendiou 
(100 m), 11.1V.1986]. One specimen observed in Kös Town on 
10-IV.[1986].” (Olivier, 1986 : 111, 113) 

— further reported by Olivier (1993a : 198 [Table 12]) and Hessel- 
barth, van Oorschot & Wagener (1995 : 1110 [Table 27]) 
N — Zipari (0-50 m), 27.V.1988 (obs.), 18.V1.1992 
— Oros Dikeos (400-800 m), 28.V.1988 (obs.) 
— Marmari (0 m), 22.1X.1992, 23.1X.1992, 28.1X.1992 


Papilio machaon syriacus Eller, 1936 


L 


— “3 esemplari QQ Coo (...). Gli esemplari furono raccolti tutti e 


tre a Coo. [VIII.1926].” (Turati, 1929 : 177-178) 

“Kos, Mt. Dikeo, 8.VI.[1935], (...).” (Rebel, 1936 : 21) 

“Kos Town. Common with females ovipositing. [30.V.1983].” 
(Thomson, 1985 : 154) 

“I took one fresh male at B [7 km W Kos (0 m), 11.1V.1986].” 
(Olivier, 1986 : 111, 113) 

further reported by Bernardi (1971 : 25), Olivier (1993a : 56, 198 
[Table 12]) and Hesselbarth, van Oorschot & Wagener (1995 : 
1110 [Table 27]) 

Zipäri (0 m), 27.V.1988, 1.V1.1992 

Marmari (0 m), 22.IX.1992, 28.1X.1992 

Mastihari (0 m), 26.1X.1992 

Pili (50 m), 22.IX.1992, 26.1X.1992 


Colias crocea (Fourcroy, 1785) 


| eae 


“Kos Town. Very common, including many f. helice Hubner. 
[ 30. V.1983].” (Thomson, 1985 : 155) 

“I observed one single female at B [7 km W Kos (0 m), 11.IV. 
1986].” (Olivier, 1986 : 111, 113) 

“Kardamena, 1/7.V11.1978, 1 © (worn) (K [M.C. & G. KRUSE- 
MAN]) 

above Zia, 6. VII.1978, 1 © (worn) (K [M.C. & G. KRUSEMAN|).” 
(Olivier, 1987 : 82) 

further reported by Olivier (1993a : 198 [Table 12]) and Hessel- 
barth, van Oorschot & Wagener (1995 : 1110 [Table 271) 

2 km N Asfendioü (100 m), 26.V.1988 

Paradisi (0-50 m), 26.V.1988 

Zipari (0 m), 27.V.1988 (obs.) 

Oros Dikeos (400-800 m), 28.V.1988 

Marmari (0 m), 17.1X.1992, 19.1X.1992, 22.1X.1992, 27.1X.1992, 
28.1X.1992 


Gonepteryx farinosa farinosa (Zeller, 1847) 


Ne 


Oros Dikeos (750-846 m), 4.V1.1992 (1 &, obs.) 

First-ever record for this island ! 

Dr Klaus Rose (in litt., VI.1996) collected 1 & “1 km N Ziparı” 
on 3.VI.1996. 


Euchloe (ausonia) ausonia taurica Rober, [1907] 


| eee 


“I found the species quite commonly at A [1 km W Agios 
Dimitrios (300 m), 10.IV.1986] and C [2 km N Asfendiou 


193 


(100 m), 11.IV.1986]. All specimens observed belonged to the first 
brood.” [Erroneously recorded as Euchloe simplonia (Freyer, 
1829)] (Olivier, 1986 : 111, 113) 

— further reported by Olivier (1990 : 181 ; 1993a : 198 [Table 12]) 
and Hesselbarth, van Oorschot & Wagener (1995: I111 [Ta- 
ble 27) 

N — Zipari (0 m), 27.V.1988 
— Asfendioü (300 m), 2.1V.1995 
— 2km E Paradisi (0-50 m), 3.1V.1995 


Pieris brassicae brassicae (Linnaeus, 1758) 
L — “2 esemplari & e Q (...) sono stati raccolti a Coo. [VIII.1926].” 
(Turati, 1929 : 178) 

— “Kos Town. Very common (...). [30.V.1983].” (Thomson, 1985 : 
155) 

— “I took the species at A [1 km W Agios Dimitrios (300 m), 
10.1V.1986] and C [2 km N Asfendioü (100 m), 11.1V.1986].” 
(Olivier, 1986 : 111, 113) 

— “above Zia, 6. VII. 1978, 1 Q (fresh) (K [M.C. & G. KrusEMAN]) 
between Akron Fokäs and Émpros Therma, 3.VIL.1978, 1 
(rather worn) (K [M.C. & G. KRuUSEMAN]).” (Olivier, 1987 : 82) 

— further reported by Bernardi (1971 : 25), Olivier (1993a : 76, 198 
[Table 12]) and Hesselbarth, van Oorschot & Wagener (1995 : 
111 [Table 27]) 

N — Agios Dimitrios (250 m), 26.V.1988 

— Paradisi (0-50 m), 26.V.1988 

— Zipari (0 m), 27.V.1988 

— Oros Dikeos (400-800 m), 28.V.1988 

— 2km E Paradisi (0-50 m), 3.1V.1995 


Pieris krueperi krueperi Staudinger, 1860 
L — previously reported by Olivier (1993a : 198 [Table 12]) and Hes- 
selbarth, van Oorschot & Wagener (1995 : 1111 [Table 27]) 
N — Oros Dikeos (400-800 m), 28.V.1988 
— Oros Dikeos (750-846 m), 4.VI.1992 


Pieris rapae rapae (Linnaeus, 1758) 
L — “Ein kleineres Pärchen von Kos, 6. bis 9. VI.[1935], (...).” (Rebel, 
1936: 21) 
— “Kos Town (...). Common where noted. [30.V.1983].” (Thomson, 
1985 :, 155) 
— “I found it at B [7 km W Kos (0m), 11.IV.1986].” (Olivier, 
1986 : 111, 113) 


194 


“Kardämena, 1/7.VIL.1978, 1 4 (fresh) (K [M.C. & G. KRUSE- 
MAN |)” (Olivier, 1987 : 83) 

further reported by Bernardi (1971 : 25), Olivier (1993a : 198 [Ta- 
ble 12]) and Hesselbarth, van Oorschot & Wagener (1995 : 1111 
[Table 27]) 

Zipari (0 m), 27.V.1988 

Oros Dikeos (400-800 m), 28.V.1988 

Asfendioü (300 m), 16.V1.1992 

Kos (0-50 m), 19.1X.1992 

Marmari (0 m), 27.IX.1992, 28.1X.1992 

2 km E Paradisi (0-50 m), 4.1V.1995 


Pontia (daplidice) edusa (Fabricius, 1777) 


en 


“6 esemplari dd e P® di Rodi e di Coo (...). [VHI.1926].” 
(Turati, 1929 : 179) 

“Kos, Cardamena, 6.V1.[1935].” (Rebel, 1936 : 22) 

“Kos Town. Common. [30.V.1983].” (Thomson, 1985 : 155) 
further reported by Bernardi (1971: 25), Olivier (1986 : 113; 
1993a : 198 [Table 12]) and Hesselbarth, van Oorschot & Wage- 
ner (1995 : 1111 [Table 271) 

Pili (50-100 m), 16.V1.1992, 26.1X.1992 

Kos (0-50 m), 20.1X.1992 

Marmäri (0 m), 19.1X.1992, 22.1X.1992, 28.IX.1992 

One specimen collected at Marmäri on 28.1X.1992 is a bilateral 
gynandromorph (Fig. 3). 


Fig. 3. Pontia (daplidice) edusa (Fabricius, 1777) 


Bilateral gynandromorph 
Marmari (Kos, Greece) (0 m), 28.IX.1992 (leg. et coll. R. De Prins) 


Lycaena phlaeas phlaeas (Linnaeus, 1761) 


Be 


“Kos, Cardamena, 6.VI[19351, und Mt. Dikeo, 8.V1.[1935].” 
(Rebel, 1936 : 26) 


— “Kos Town. (...). Common to the west and north of Kos Town. 


[30.V.1983].” (Thomson, 1985 : 157) 


195 


“A single female observed at A [1 km W Agios Dimitrios 
(300 m), 10.IV.1986].” (Olivier, 1986 : 111, 113) 

further reported by Bernardi (1961 : 193 [Fig. 13] ; 1971 : 24, 26 
[Map 2], Olivier (1993a : 198 [Table 12]) and Hesselbarth, van 
Oorschot & Wagener (1995 : 1111 [Table 27]) 

2 km N Asfendioü (100 m), 25.V.1988 

Agios Dimitrios (250 m), 26.V.1988 

2 km NE Agios Dimitrios (150 m), 26.V.1988 

Zipari (0 m), 27.V.1988 

Oros Dikeos (400-840 m), 28.V.1988, 4.IV.1995 (obs.) 

5 kmW Kos (0 m), 16.V1.1992, 20.VI.1992 

2 km E Paradisi (0-50 m), 3.1V.1995 


Lycaena thersamon (Esper, [1784]) 


Be 


“8 esemplari & e 9Q di questa forma estiva caudata raccolta 
a Rodi ed a Coo. [ VIII.1926].” (Turati, 1929 : 183) 
“Kardämena, 1/7.V11.1978, 1 4 (rather worn), 1 9 (rather fresh) 
(K [M.C. & G. KRUSEMAN]) 

above Zia, 6.V11.1978, 1 4 (not so fresh) (K [M.C. & G. KRUSE- 
MAN]). (Olivier, 1987 : 85) 

further reported by Bernardi (1971 : 27), Olivier (1986: 113; 
1993a : 45, 94, 198 [Table 12]) and Hesselbarth, van Oorschot 
& Wagener (1995 : 1111 [Table 27]) 

Zipari (0 m), 27.V.1988 

5 km W Kos (0 m), 16.V1.1992, 18.V1.1992, 20.VI.1992 

Kos (0-50 m), 20.1X.1992, 22.1X.1992, 26.1X.1992 

Marmari (0 m), 17.1X.1992, 19.1X.1992, 22.1X.1992, 26.1X.1992, 
28.1X.1992 


Callophrys rubi (Linnaeus, 1758) 


Pier 


Nee 


“one specimen found on a roadside at C [2 km N Asfendiou 
(100 m), 11.1V.1986], two more taken in dry garrigue flying 
around bushes of Quercus ilex at A[1 km W Agios Dimitrios 
(300 m), 11.1V.1986].” (Olivier, 1986 : 111, 113) 

further reported by Olivier (1993a : 198 [Table 12]) and Hessel- 
barth, van Oorschot & Wagener (1995 : 1111 [Table 27]) 

2 km E Paradisi (0-50 m), 3.1V.1995 


Satyrium ilicis ilicis (Esper, [1779]) 


er 


196 


“west of Kos Town (...). A few males and females in a light- 
ly wooded grass valley in Kos. [30.V.1983].” (Thomson, 1985 : 
156-157) 


— further reported by Olivier (1986 : 113 ; 1993a : 198 [Table 12]; 
1993b : 21) and Hesselbarth, van Oorschot & Wagener (1995 : 
1111 [Table 271) . 
N — Agios Dimitrios (250 m), 26.V.1988 
— 2km NE Agios Dimitrios (150 m), 26.V.1988 
— Oros Dikeos (400-800 m), 28.V.1988 
— Mastihari (0 m), 29.V.1988 


Lampides boeticus (Linnaeus, 1767) 
L — “Parecchi esemplari dei due sessi di (...) Coo (...). [VIII 1926].” 
(Turati, 1929 : 183) 
— “Kardamena, 1/7.V11.1978, 1 & (worn) (K [M.C. & G. Kruse- 
MAN |) 
between Akron Fokäs and Empros Therma, 3.VIL.1978, 13 
(worn) (K [M.C. & G. KRuUSEMAN]).” (Olivier, 1987 : 85) 
— further reported by Bernardi (1971: 24), Olivier (1986: 113; 
1993a : 198 [Table 12]) and Hesselbarth, van Oorschot & Wage- 
ner (1995 : 1111 [Table 27]) 
N — Zipäri (0 m), 27.V.1988, 1.VI.1992 
— Kös (0 m), 8.V1.1992 
— Marmari (0 m), 17.1X.1992, 19.1X.1992, 28.1X.1992 


Leptotes pirithous (Linnaeus, 1767) 
L — “Kos, Cardamena, 6.V1.[1935], drei &.” (Rebel, 1936 : 26) 
— further reported by Bernardi (1971 : 24), Olivier (1986 : 113; 
1993a : 198 [Table 12]) and Hesselbarth, van Oorschot & Wage- 
ner (1995 : 1111 [Table 27]) 
N — Marmari (0 m), 17.1X.1992, 19.1X.1992, 26.1X.1992, 28.1X.1992 


Celastrina argiolus argiolus (Linnaeus, 1758) 

L — “A [1 km W Agios Dimitrios (300 m), 10.1V.1986, 11.1V.1986] 
and C [2 km N Asfendiou (100 m), 11.1V.1986]. Observed flying 
around buckthorn (Prunus spinosa L.).” (Olivier, 1986 : 112-113) 

— further reported by Olivier (1993a : 199 [Table 12]) and Hessel- 
barth, van Oorschot & Wagener (1995 : 1111 [Table 27]) 
N — 3km SW Andimähia (50 m), 25.V.1988 
— 2 km N Asfendiou (100 m), 26.V.1988, 27.V.1988 
— Zipari (0 m), 27.V.1988 
— Marmari (0 m), 17.1X.1992 
— Pili (50 m), 26.1X.1992 


197 


Glaucopsyche alexis alexis (Poda, 1761) 

L — “found only at A [1 km W Agios Dimitrios (300 m), 10.1V.1986, 
11.1V.1986] on dry garrigue and on an open spot with many 
Asphodelia. (...).” (Olivier, 1986 : 112-113). | 

— further reported by Olivier (1993a : 102, 199 [Table 12]) and 
Hesselbarth, van Oorschot & Wagener (1995 : 1111 [Table 27]) 
N — 2kmE Paradisi (0-50 m), 3.1V.1995, 4.1V.1995 


Polyommatus loewii loewii (Zeller, 1847) 

L — “zwei 6 von Kos, Mt. Dikeo, 8.V1.[1935], (...). Die Art erreicht 
auf Kos und Kalymnos wohl ihre nordöstliche Verbreitungs- 
grenze.” (Rebel, 1936 : 26-27) 

— “west of Kos Town. Locally common. [30.V.1983].” (Thomson, 
1985 : 157) 

— “<<Kos, 23.V.1983>>, 1 & (fresh, but badly damaged) [leg. 
J. J. v. OorT].” (Olivier, 1987 : 86) 

— further reported by Bernardi (1961 : 184, 207 ; 1971 : 28), Olivier 
(1986: 113 ; 1993a: 106, 199 [Table 12]) and Hesselbarth, van 
Oorschot & Wagener (1995 : 613, 614, 1112 [Table 27]) 

N — 2km E Paradisi (0 m), 4.V1.1992, 8.V1.1992 

— Oros Dikeos (750-846 m), 17.VI.1992 
Hesselbarth, van Oorschot & Wagener (1995 : 613-617, 627-631) 
treat the present as well as the next species as belonging to the 
genus Plebeius Kluk, 1780. We prefer to include these in the 
genus Polyommatus (cf. Olivier, 1993a : 26, 106-109), the more 
as Hesselbarth, van Oorschot & Wagener (1995 : 588) admit : 
“Bisher fehlt eine umfassende Untersuchung, die eine einiger- 
maßen gesicherte, phylogenetisch begründete Aufteilung der 
Arten dieser und der folgenden Gattung Polyommatus auf die 
einzelnen Untergattungen und deren Stellung zueinander er- 
lauben würde. (...). Unsere Beurteilung bleibt daher notgedrun- 
gen subjektiv (...).” 


Polyommatus agestis ([ Denis & Schiffermüller], 1775) 
L — “one single fresh female at A [1 km W Agios Dimitrios (300 m)] 
on 10-IV.[1986].” (Olivier, 1986 : 112-113) 
— “above Zia, 6.V11.1978, 1 © (not so fresh) (K [M.C. & G. KRUSE- 
MAN]).” (Olivier, 1987 : 87) 
— further reported by Olivier (1993a: 108, 199 [Table 12]) and 
Hesselbarth, van Oorschot & Wagener (1995 : 1112 [Table 27]) 
N — 2 km E Paradisi (0 m), 8.VI.1992 
— 5 km W Kos (0 m), 16.V1.1992 


198 


Polyommatus thersites (Cantener, [1835]) 
L — “18 e299 di Coo e Rodi (...). [VIII.1926].” (Turati, 1929 : 184) 
— “Drei frische & und ein solches © von Kos, Cardamena, 6.VI. 
[1935], (...).” (Rebel, 1936 : 26) 
— “above Zia, 6. VIL.1978, 3 4 (not so fresh), 3 9 (one fresh, two 
not so fresh) (K [M.C. & G. KRUSEMAN]).” (Olivier, 1987 : 87) 
— further reported by Bernardi (1961: 184; 1971 : 28), Olivier 
(1986 : 113 ; 1993a : 110, 199 [Table 12]) and Hesselbarth, van 
Oorschot & Wagener (1995 : 1112 [Table 27]) 
N — Pili (50-100 m), 16.VI.1992 
— Zia (450 m), 17.V1.1992 
— Oros Dikeos (750-846 m), 17.V1.1992 


Polyommatus icarus (Rottemburg, 1775) 
L — “Molti esemplari di Coo, (...). [ WII.1926].” (Turati, 1929 : 184) 

— “Kos, Cardamena, 6.VI.[1935], 4 4, 1 ® (...).” (Rebel, 1936 : 26) 

— “Kos Town. Very common. [30.V.1983].” (Thomson, 1985 : 157) 

— “One single very fresh male at A [1 km W Agios Dimitrios 
(300 m), 10.IV.1986].” (Olivier, 1986 : 112-113) 

— “Agios Stéfanos, 24/29.V1.1978, 2 2 (worn) (K [M.C. & G. 
KRUSEMAN)]) 
between Akron Fokas and Empros Therma, 3.V11.1978, 1 © 
(worn) (K [M.C. & G. KRusEMAN]).” (Olivier, 1987 : 88) 

— further reported by Bernardi (1961 : 192 [Fig. 2]; 1971 : 25, 26 
[Map 2]), Olivier (1993a : 199 [Table 12]) and Hesselbarth, van 
Oorschot & Wagener (1995 : 1112 [Table 27]) 

N — 5 km W Kos (0 m), 16.V1.1992 

— Zipäri (0 m), 18.V1.1992 

— Marmari (0 m), 26.1X.1992, 28.1X.1992 

— Kos (0-50 m), 20.1X.1992, 22.1X.1992 


Danaus chrysippus chrysippus (Linnaeus, 1758) 
L — “16 esemplari raccolti a Coo, (...). [VIII.1926].” (Turati, 1929 : 
179) 
— “[Coo.] (...). Noto, fra i Lepidotteri, abbondante Danais chry- 
sippus [sic !], (...). [VIII.1926].” (Ghigi, 1929 : 309) 
— further reported by Bretherton (1966 : 19), Bernardi (1971 : 27), 
Olivier (1986 : 113 ; 1993a : 116, 199 [Table 12]) and Hesselbarth, 
van Oorschot & Wagener (1995 : 767) 
N — Marmari (0 m), 27.1X.1992 
One very fresh female (Fig. 4). The first record from this area in 
66 years ! The senior author has been informed by Mr. Finn Iversen 


199 


Fig. 4. Danaus chrysippus chrysippus (Linnaeus, 1758) 2 
Marmari (Kos, Greece) (0 m), 27.1X.1992 (leg. et coll. R. De Prins) 


(in litt., 17.1.1994) that he observed one specimen at Trianda, on the 
island of Rodos, on 10.X.1993. 


Kirinia roxelana (Cramer, [1777]) 
L — “ein Pärchen von Kos, Cardamena, 6.VI[1935], davon das © 
ganz frisch.” (Rebel, 1936 : 24) 

— “Kos Town. (...). Common west of Kos Town. [30.V.1983].” 
(Thomson, 1985 : 155) 

— further reported by Bernardi (1971: 27), Olivier (1986: 113; 
1993a : 190, 199 [Table 12]) and Hesselbarth, van Oorschot & 
Wagener (1995 : 1112 [Table 27]) 

N — 2kmN Asfendiot (100 m), 25.V.1988, 26.V.1988, 27.V.1988 

— Zipari (0 m), 27.V.1988 


Pararge aegeria aegeria (Linnaeus, 1758) 

L — “one single male was taken at C [2 km N Asfendioü (100 m), 
11.1V.1986], fluttering discretely under a scrub, near a small 
footpath. (...).” (Olivier, 1986 : 112-113) 

— further reported by Olivier (1993a: 199 [Table 12], 207) and 
Hesselbarth, van Oorschot & Wagener (1995 : 1112 [Table 27]) 


Lasiommata megera lyssa (Geyer, |1828]) 
L — “ein Pärchen von Kos, Cardamena, 6.VI.[1935], (...).” (Rebel, 
1936 : 24) 
— “Kos Town (...). Common (...). [30.V.1983].” (Thomson, 1985 : 
155) 


200 


— “Two males and one female were taken on 10-IV.[1986] at A 
[1 km W Agios Dimitrios (300 m)] between the ruines of the 
abandoned village.” (Olivier, 1986 : 112-113) 
— further reported by Bernardi (1971 : 27), Olivier (1993a : 199 [Ta- 
ble 12]) and Hesselbarth, van Oorschot & Wagener (1995 : 1112 
[Table 27]) 
N — 1 km N Akron Fokäs (0-50 m), 26.V.1988 
— Oros Dikeos (400-800 m), 28.V.1988 
— Oros Dikeos (840 m), 4.IV.1995 
— Mastihari (0-50 m), 29.V.1988 
— Marmari (0 m), 26.1X.1992, 28.1X.1992 
— 2 km E Paradisi (0-50 m), 3.1V.1995 
De Freina ([1992]) studied the geographic variation of L. megera and 
showed the material from the Balkans, Turkey and Iran to differ 
constantly from the remaining populations in both the male and female 
genitalia. Olivier (1993a), though knowing this publication, did not fully 
realize its significance and incorrectly lumped the East Mediterranean 
populations under L. megera megera (Linnaeus, 1767). Coutsis (pers. 
comm.) confirms the distinction of a western and eastern taxon after 
examination of the male genitalia. Hesselbarth, van Oorschot & Wage- 
ner (1995 : 782-783) use the name L. megera transcaspica (Staudinger, 
1901) for Turkish populations, as they consider /yssa, the type-locality 
of which is Ragusa [Dubrovnik] in Dalmatia, as part of the “south- 
western” group on external markings. The male genitalia of a specimen 
from Dalmatia, described and illustrated by de Freina ([1992]: 169 
[fig. 5], 171), clearly place it as belonging to the eastern type. Therefore, 
we consider material from the Balkans eastwards until Iran as belonging 
to L. megera lyssa (Geyer, [1828]), with transcaspica as a junior 
synonym, as already established by de Freina ([1992] : 172). 


Lasiommata maera (Linnaeus, 1758) 
L — “Ein frisches & von Kos, Cardamena, 6.V1.[1935], (...).” (Rebel, 
1936 : 24) 

— “Two males and one female taken at A[1 km W Agios Dimitrios 
(300 m)] on 11-IV.[1986] along a steep rocky slope in an area 
of dry garrigue near a small pine wood. Two specimens are 
very fresh, one is very worn.” (Olivier, 1986 : 112-113) 

— further reported by Bernardi (1961: 184; 1971: 27), Olivier 
(1993a : 188, 199 [Table 12]) and Hesselbarth, van Oorschot & 
Wagener (1995 : 1112 [Table 27]) 

N — Oros Dikeos (400-800 m), 28.V.1988 

— Kos (0-50 m), 19.1X.1992, 20.1X.1992 


201 


Maniola telmessia (Zeller, 1847) 


[ty == 


“Von Kos liegen, von Cardamena, 6.VI[1935], und dem Mt. 
Dikeo, 8. VI:[1935], €...) vor (2). (Rebel 1936 225) 

“Kos Town and valley to the west of Kos Town (...). Very 
common in a variety of habitats (...) dry lightly wooded rough 
grassland (...) west of Kos Town, gardens at Kos Town (...). 
[30.V.1983].” (Thomson, 1985 : 155) 

further reported by Hartig (1940 : 231), Bernardi (1961 : 184; 
1971: 27), Tauber & Tauber (1968 : 83 [Fig. 3], 84), Olivier 
(1986: 113; 1990: 182; 1993a: 146-147, 148 [Table 6], 150 
[Table 7], 155, 156 [Table 8], 158 [Table 9], 199 [Table 12]; 
1993b : 22), Olivier & Coutsis (1993: 114, 115 [Fig. 1], 116, 
119-121 [incl. Plates 1 & 2], 123 [Table 1], 124 [Table 2], 126 
[Table 3], 127 [Table 4]; 1995 : 10 [Fig. 2], 11, 12-15 [Table 2], 
16 [Table 3], 18, 27, 32 [Table 12], 34 [Table 13], 36, 44 [ Figs. 44- 
45], 56), Thomson (1987 : 13, 18 [Fig. 3], 19 ; 1990: 153 [Ta- 
ble 3], 155), van Oorschot & van den Brink (1992: 149, 150 
[ Fig. 1]) and Hesselbarth, van Oorschot & Wagener (1995 : 825- 
827, 1112 [Table 27]) 

2 km N Asfendiou (100 m), 25.V.1988, 26.V.1988, 27.V.1988, 
16. VI.1992, 17.VI.1992 

3 km SW Andimahia (50 m), 25.V.1988 

Agios Dimitrios (250 m), 26.V.1988 

2 km NE Agios Dimitrios (150 m), 26.V.1988 

Paradisi (0-50 m), 26.V.1988 

1 km N Akron Fokas (0-50 m), 26.V.1988 

Zipari (0 m), 27.V.1988 

Oros Dikeos (400-800 m), 28.V.1988 

Oros Dikeos (750-846 m), 17.VI.1992 

Mastihari (0 m), 29.V.1988 

Zia (450 m), 4.V1.1992 

Marmari (0 m), 19.1X.1992, 26.1X.1992 

Pili (50 m), 26.1X.1992 

Material collected in September 1992 consists exclusively of post- 
aestivation females. 


Hyponephele lupina ? intermedia (Staudinger, 1886) 
1G 


“3 esemplari PS di Coo e Fonducli [ VUI.1926].” (Turati, 1929 : 
182) 


— “Ein frisches © von Kos, Mt. Dikeo, 8.VI.[1935], stimmt voll- 


202 


kommen mit einem solchen von Kreta.” (Rebel, 1936 : 25) 


— “west of Kos Town. Four freshly emerged males only in heath 
garique’ [sic!]. [30. V.1983].” (Thomson, 1985 : 155) 

— “Kos, 10/24.V.1982, 1 © (rather worn) (P.J.H. van Bree) 

4 km E. of Kos, 8.VII.1978, 1 2 (not so fresh) (K [M.C. & G. 
KRUSEMAN)]).” (Olivier, 1987 : 168) 

— further reported by Bernardi (1971 : 28), Olivier (1986: 113; 
1993a : 177, 199 [Table 12]) and Hesselbarth, van Oorschot & 
Wagener (1995 : 1112 [Table 27]) 

N — 4km E Andimähia (0-50 m), 19.VI.1992, 20.VI.1992 

Male material from Kös is of the bright, brassy ochreous brown morph 
with long arcuate sex brand and pointed, elongated forewings, even 
more so than Rodos specimens, thus looking very much like nomi- 
notypical H. lupina lupina (Costa, [1836]) from the Greek mainland 
and from Calabria, though material of the latter population is distinctly 
smaller-sized (Olivier, pers. obs. July-August 1995). Females collected 
by the senior author on the nearby Bodrum Peninsula agree very well 
with those from Kös, but unfortunately no male material from there 
is available to us. Baldock & Bretherton (1988), however, refer their 
single male specimen to “ Hyponephele lupina Costa rhamnusia Freyer”, 
thus suggesting that it is of the same type, recalling the Sicilian sub- 
species. We unhappily treat this material from Kos as belonging to 
ssp. intermedia, mainly because of its geographic location. A thorough 
taxonomic revision of this species is much needed ! 


Hipparchia senthes (Fruhstorfer, 1908) 

L — previously reported by Hartig (1940: 226, as “Satyrus semele 
L. cretica Rol.” [sic!]), Olivier (1986 : 110, 113 ; 1993a : 199 [Ta- 
ble 12]) and Hesselbarth, van Oorschot & Wagener (1995 : 1113 
[Table 271) 

N — Oros Dikeos (400-800 m), 28.V.1988 

— Oros Dikeos (750-846 m), 4. VI.1992, 17.VI.1992 

Hipparchia senthes has traditionally been included in a group of 

allopatric taxa referred to as the A. aristaeus (Bonelli, 1826) complex 

(see e.g. Kudrna, 1977 ; Kudrna, 1984 ; Balletto er al., 1990 ; Hessel- 

barth, van Oorschot & Wagener, 1995), being considered either as a 

subspecies of H. aristaeus or as a distinct species. Recently, Cesaroni 

et al. (1994) studied a.o. allozymes and genital morphometrics of seven- 

teen populations belonging to nine presumptive species or subspecies 

of Hipparchia (Parahipparchia), including three taxa of the A. aristaeus 

complex, convincingly refuting the conspecificity of these. Current 

investigations on the morphology of taxa of this group (Olivier & 


203 


Coutsis, in prep.) support these conclusions. Therefore, we consider 
H. senthes as specifically distinct from H. aristaeus. 


Hipparchia fatua fatua Freyer, [1844] 
L — “Gli esemplari (...) furono presi a (...) Coo (...) [WHI.1926].” 
(Turati, 1929 : 180) 
— “Agios Stéfanos, 24/29.V1.1978, 1 © (fresh) (K [M.C. & G. Kru- 
SEMAN |) | 
above Zia, 6.VIL.1978, 1 © (fresh) (K [M.C. & G. KRUSEMAN]) 
4 km E of Kos, 8.VII.1978, 1 & (not so fresh) (K [M.C. & G. 
KRUSEMAN)).” (Olivier, 1987 : 167) 
— further reported by Ghigi (1929 : 309, 331), Bernardi (1961 : 
184, 194 [Fig. 4]; 1971: 26 [Map 3], 28), Bretherton (1966 : 
36), Olivier (1986 : 113 ; 1990 : 187 ; 1993a : 135-136, 199 [Ta- 
ble 12]) and Hesselbarth, van Oorschot & Wagener (1995 : 913, 
914, 1113 [Table 271) 
N — 4km E Andimähia (0-50 m), 19.V1.1992 
— Marmari (0 m), 19.1X.1992, 28.1X.1992 (only females !) 


Pseudochazara anthelea anthelea (Hübner, [1824]) 

L — “Vier & und ein frisches © von der Insel Kos, Mt. Dikeo, 8.VI. 
[1935]. (...). Sie flog nur in der Gipfelregion, dort aber häufig. 
Weiter unten wurde sie nirgends gesehen.” (Rebel, 1936 : 23-24) 

— further reported by Rebel (1939: 508), Hartig (1940: 226), 
Bernardi (1961 : 189, [190], 194 [Fig. 4], 201, 207, [209] ; 1971 : 
28), van Oorschot, van den Brink & van Oorschot (1987 : 91, 
94), Olivier (1986 : 113 ; 1993a : 140-141, 199 [Table 12] ; 1996: 
5-6, 8 [Plate 2], 9 [Tables 1 & 2], 10-12) and Hesselbarth, van 
Oorschot & Wagener (1995 : 962, 1113 [Table 27]). 

N — Oros Dikeos (400-800 m), 28.V.1988 
— Oros Dikeos (750-846 m), 4.VI.1992 (obs.), 17. VI.1992 


Limenitis reducta herculeana Stichel, [1908] 
L — “west of Kos Town. A few very worn individuals. [30.V.1983].” 
(Thomson, 1985 : 156) 
— further reported by Olivier (1986 : 113 ; 1993a : 199 [Table 12]) 
and Hesselbarth, van Oorschot & Wagener (1995: 1113 [Ta- 
ble 27]) 


Vanessa atalanta atalanta (Linnaeus, 1758) 
L — “1 esemplare di Coo (...) [VIII.1926].” (Turati, 1929 : 182) 
— Observed at C [2 km N Asfendioü (100 m), 11.1V.1986].” 
(Olivier, 1986 : 112-113) 


204 


— further reported by Ghigi (1929 : 309), Bernardi (1971 : 25), Oli- 


vier (1993a : 199 [Table 12]) and Hesselbarth, van Oorschot & 
Wagener (1995 : 1113 [Table 27]) 

2 km N Asfendioü (100 m), 25.V.1988 

Zipari (0 m), 27.V.1988 

Agios Dimitrios (300 m), 2.1V.1995 (obs.) 

Asfendioü (300 m), 2.1V.1995 (obs.) 

2 km E Paradisi (0-50 m), 3.1V.1995 (obs.), 4.1V.1995 (obs.) 
Oros Dikeos (840 m), 4.1V.1995 (obs.) 

Zia (450 m), 4.1V.1995 (obs.) 


Very common to abundant in all forementioned localities visited in 
April 1995. 


Vanessa cardui (Linnaeus, 1758) 


Lite: 


“5 esemplari di Coo, (...) [ VIII.1926].” (Turati, 1929 : 183) 
“Kos, Cardamena, 6.V1.[1935], ein frisches 3.” (Rebel, 1936 : 22) 
“Kos Town. Common. [30.V.1983].” (Thomson, 1985 : 156) 
“Observed at A [1 km W Agios Dimitrios (300 m), 10.1V.1986].” 
(Olivier, 1986 : 112-113) 

further reported by Ghigi (1929 : 309), Bernardi (1971: 27), 
Koutsaftikis (1973 : 171), Olivier (1993a : 199 [Table 12]) and 
Hesselbarth, van Oorschot & Wagener (1995 : 1113 [Table 27]) 
2 km N Asfendiou (100 m), 25.V.1988 (obs.), 26.V.1988 

Zipari (0 m), 27.V.1988 (obs.) 

Mastihäri (0 m), 29.V.1988 

Marmari (0 m), 22.1X.1992 

Agios Dimitrios (300 m), 2.1V.1995 (obs.) 

Asfendioü (300 m), 2.1V.1995 (obs.) 

2 km E Paradisi (0-50 m), 3.1V.1995 (obs.), 4.1V.1995 (obs.) 

— Oros Dikeos (840 m), 4.1V.1995 (obs.) 

Zıa (450 m), 4.1V.1995 (obs. ) 


Very common to abundant in all forementioned localities visited ın 
April 1995, even more so than V. atalanta. 


Polygonia egea (Cramer, [1775]) 
L 


— “5 esemplari di Nisiro, Coo e Scarpanto. [ VIIL.1926].” (Turatı, 
1929 : 183) 

“Kos Town. Several in gardens. [30.V.1983].” (Thomson, 1985 : 
156) 

“One male was observed at A [1 km W Agios Dimitrios (300 m)] 
on 10-IV.[1986] and again on 11-IV.[1986].” (Olivier, 1986 : 112- 
113) 


205 


— further reported by Ghigi (1929 : 309), Hartig (1940 : 232), Ber- 
nardi (1971 : 27), Olivier (1993a : 193, 199 [Table 12]) ; 1993b : 
21) and Hesselbarth, van Oorschot & Wagener (1995 : 1113 
[Table 27]) 

N — Zipari (0 m), 27.V.1988 (obs.) 

— Oros Dikeos (400-800 m), 28. V.1988 

— Oros Dikeos (750-846 m), 4. VI.1992 

— Kos (0-50 m), 24.1X.1992 


Nymphalis polychloros polychloros (Linnaeus, 1758) 

L — previously reported by Olivier (1993a : 199 [Table 12]) and Hes- 
selbarth, van Oorschot & Wagener (1995 : 1113 [Table 27]) 

N — Oros Dikeos (400-800 m), 28.V.1988 (obs.) 


Melitaea trivia trivia ([Denis & Schiffermüller], 1775) 
L — “Vier & von Kos, Cardamena, 6.V1.[1935], (...).” (Rebel, 1936 : 
23) 
— “4 km E. of Kos, 8.V11.1978, 19 (not so fresh) (K [M.C. & G. 
KRUSEMAN)]).” (Olivier, 1987 : 166) 
— further reported by Rebel (1937 : 65), Hartig (1940 : 226), Ber- 
nardi (1961 : 184, 195 [Fig. 5] ; 1971 : 26 [Map 3], 28), Koutsaf- 
tikis (1973 : 176), Olivier (1986: 113 ; 1993a : 199 [Table 12]) 
and Hesselbarth, van Oorschot & Wagener (1995 : 1113 [Ta- 
ble 27]) 
N — Oros Dikeos (400-800 m), 28.V.1988 


Doubtful and erroneous records 


Euchloe simplonia Boisduval, [1828] 

This taxon, which is very closely related to E. ausonia, ıs restricted 
to the central and western Alps, the Pyrenees and the Cantabrian 
Mountains. Olivier (1986) was wrong in ascribing his E. ausonia records 
from Kos to E. simplonia (Freyer, 1829), that was decribed after 
material from Croatia and hence is a junior subjective synonym of 
E. ausonia (Hübner, [1804}). 


Pontia daplidice (Linnaeus, 1758) 

All records from Kös previous to Olivier (1993a) refer to Pontia 
daplidice, while this nominal taxon has been shown to consist of two 
genetically distinct taxa. Kös lies within the distribution range of the 
“eastern” taxon, 1.e. P edusa. Both are currently united in one super- 
species. 


206 


Cupido minimus (Fuessly, 1775) 

Thomson (1985 : 157) recorded this butterfly from Kos, stating : “west 
of Kos Town. Locally common in grassy valley. Found in mainland 
Greece but new to the Dodecanes.” Subsequently, Olivier (1986 : 113) 
mentions this species again in his checklist, but it is not listed anymore 
in Olivier (1993a). 

Its presence on Kos is very unlikely, considering its distribution in 
Turkey (cf. Hesselbarth, van Oorschot & Wagener, 1995). 

Thomson (in litt., 8.11.1989) says : “I do not have specimens of minimus 
from Kos”, hence we prefer not to include this species among the 
butterfly fauna of this island on present evidence. 


Hipparchia cretica (Rebel, 1916) 

This species belongs to a complex of taxa that resemble each other 
closely externally. On Kos, the senior author collected a small series 
of specimens that all belong to the rather distantly related H. senthes. 
H. cretica is an insular endemic restricted to the island of Kriti (Crete). 
Hartig’s (1940) record is surely erroneous and probably refers to H. 
senthes as well. 


Argynnis pandora pandora (| Denis & Schiffermüller], 1775) 

One large nymphalid, most probably a fritillary belonging to this 
species, was observed on Oros Dikeos, at an altitude of about 800 m, 
on 17.V1.1992. As the identification is not absolutely sure, the species 
is not included in the island list until possible confirmation in the future. 


Discussion 


All 46 species positively recorded on Kos are widespread over the 
neighbouring areas : 38 of these have been traced on at least five other 
of the Eastern Aegean Islands (Olivier, 1993a ; 1993b ; 1994) and 33 
on the adjacent Bodrum Peninsula (Baldock & Bretherton, 1988 ; 
Olivier & Coutsis, 1995 ; Hesselbarth, van Oorschot & Wagener, 1995 ; 
Olivier, unpublished records). Of the 13 taxa that remain unrecorded 
from the latter area (Gegenes pumilio pumilio, Carcharodus orienta- 
lis, C. stauderi ambiguus, Archon apollinus apollinus, Pieris krueperi 
krueperi, Callophrys rubi, Leptotes pirithous, Glaucopsyche alexis 
alexis, Polyommatus thersites, Danaus chrysippus chrysippus, Pararge 
aegeria aegeria, Lasiommata maera and Polygonia egea), 11 have been 
found in the remainder of the province of Mugla. Both remaining taxa, 
C. stauderi ambiguus and P. krueperi krueperi, have been located in 
the neighbouring provinces of Izmir and Antalya, while P krueperi 


207 


krueperi is known from Aydin as well (Hesselbarth, van Oorschot & 
Wagener, 1995). C. stauderi ambiguus has been found on more than 
five other islands in the Dodekänissa (Olivier, 1993a ; 1993b ; 1994). 


We believe that at present over 90% of the actual butterfly fauna of 
Kos is already known. Only 7 taxa known from the Bodrum Peninsula 
have not yet been recorded on Kos and the very most of these are 
probably truly absent there. The fact that 13 butterfly species listed 
from Kos are not known from the Bodrum Peninsula probably reflects 
incomplete exploration of the latter area: future investigations, espe- 
cially in early spring (April) and in early June, will certainly reveal 
the presence there of most of these taxa. Interestingly, Pseudophilotes 
vicrama schiffermuelleri (Hemming, 1929) has not been found either 
on Kos or on the Bodrum Peninsula, while it has been spotted by 
the senior author on Kälimnos and Léros (April 1995) as well as on 
Nissiros (May 1992). It has possibly been overlooked in both former 
areas so far. 


No single butterfly species on Kos has differentiated significantly, not 
even subspecifically, from its counterparts in SW Turkey : this points 
to a very recent isolation of its fauna. Indeed, about 9.000 B.P., Kos 
was still part of the Anatolian mainland, being interconnected with 
the present-day Bodrum Peninsula, Kälimnos, Léros and some smaller 
islands. To the south, the islands of Nissiros and Giali were already 
as much isolated as they are now (van Andel & Shackleton, 1982). 


Vicariance of a biota that was still connected in the recent past largely 
explains the present configuration of the butterfly fauna of Kôs, while 
dispersal of migratory taxa that reach the island either occasionally 
(Danaus chrysippus chrysippus) or on a more regular base (e.g. Lepto- 
tes pirithous), 1s of more marginal significance to its make-up. 


A supposed special biogeographic affinity of the Pseudochazara anthe- 
lea population from Kos with that from Kipros (Cyprus) (Rebel, 1936) 
has recently been shown to be unsupported (Olivier, 1996). 


Acknowledgements 


We wish to thank Dr George Thomson (Lockerbie, Scotland, U.K.), Mr Finn 
Iversen (Greve, Denmark), Dr Klaus Rose (Mainz-Bretzenheim, Germany) 
and Mr John G. Coutsis (Athens) for interesting personal communications, 
as well as Mr Willy De Prins (Antwerpen) for making both photographs 
(Figs. 3 and 4). 


208 


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DES 


Nota lepid. 19 (3/4) : 212-229 ; 21.XII.1996 ISSN 0342-7536 


Myrmecophilous behaviours in caterpillars 

of the butterfly, Polyommatus icarus (Rottemburg, L7YS): 
temporal patterns and age dependency 

(Lepidoptera : Lycaenidae) 


Frank BURGHARDT* & Konrad FIEDLER** 


* Lehrstuhl Zoologie II, Theodor-Boveri-Biozentrum der Universitat, Am Hubland, D-97074 Wiirz- 
burg, Germany 

** Lehrstuhl Tierökologie I, Universität Bayreuth, Postfach 101251, D-95440 Bayreuth, Germany 
(corresponding author) 


Summary 


Mutualistic interactions between Lasius flavus (Fabricius, 1782) ants and cater- 
pillars of the Common Blue Polyommatus icarus (Rottemburg, 1775), a facul- 
tatively myrmecophilous lycaenid butterfly, were studied in a standardised 
laboratory assay. Caterpillars delivered more and larger nutritious secretion 
droplets from their dorsal nectar organ in the initial phase of an interaction 
than later on. Activity of their tentacle organs followed the same pattern. 
Myrmecophilous behaviours remained almost constant throughout the feeding 
phase of the final larval instar, whereas non-feeding prepupal larvae showed 
a more than fivefold increase in secretion rates from the nectar organ and a 
three- to fivefold increase in the frequency of tentacle eversions. The frequency 
of secretion acts as well as tentacle eversions was drastically reduced when 
larvae were raised on low-quality food. The results indicate that P icarus 
caterpillars highly flexibly respond with their mutualistic behaviours according 
to actual needs and resource availability. The close relation between the activity 
of the nectar organ and the tentacles is further evidence that the latter organs 
produce “honest signals” which advertise the capacity of a caterpillar to provide 
food rewards to attendant ants. 


Zusammenfassung 


Mutualistische Interaktionen zwischen der Ameisenart Lasius flavus (Fabricius, 
1782) und Raupen des fakultativ myrmekophilen Blaulings Polyommatus 
icarus (Rottemburg, 1775) wurden in standardisierten Laborversuchen quan- 
titativ analysiert. In den ersten Minuten einer Assoziation sezernierten die 
Raupen häufiger und größere Sekrettropfen aus ihrem Nektarorgan als in 
fest etablierten Vergesellschaftungen. Die Aktivität der Tentakelorgane nahm 
parallel zur Sekretabgaberate ab. Die Zeitmuster der myrmekophilen Ver- 


212 


haltensweisen der Raupen blieben während der fraBaktiven Phase des vierten 
Larvalstadiums trotz einer Vervierfachung ihres Gewichts nahezu konstant, 
während in der präpupalen Phase die Sekretabgaberate und die Aktivität der 
Tentakelorgane für wenige Stunden auf das Drei- bis Fünffache gesteigert 
wurden. Unter Nahrungsstreß (Aufzucht auf maturen Blättern von Medicago 
sativa) war die Sekretabgaberate wie auch die Aktivität der Tentakelorgane 
drastisch reduziert, folgte aber denselben Zeit- und Entwicklungsmustern wie 
bei Raupen auf optimaler Diät (Blütenstände von M. sativa). Diese Befunde 
zeigen, daß die Raupen von P icarus ihre myrmekophilen Verhaltensweisen 
plastisch an ihre momentane Situation (Stabilität der Ameisenassoziation, 
Entwicklungsstand, Ressourcenverfügbarkeit) anpassen. Aus der engen Kor- 
relation zwischen der Aktivität der Tentakelorgane und der Sekretabgaberate 
aus dem Nektarorgan läßt sich ferner ableiten, daß die Tentakel bei dieser 
Art als „verläßliche Signale“ wirken könnten, die den Ameisenpartnern die 
Rentabilität einer Raupe als Futterquelle anzeigen. 


Introduction 


Ants maintain a variety of mutualistic interspecific interactions with 
organisms such as plants (Beattie, 1985 ; Huxley & Cutler, 1991), sap- 
sucking homopterans or butterfly caterpillars (Hölldobler & Wilson, 
1990). An integral part of these mutualisms is the trade of two com- 
modities : ants receive nutritious substances and, in turn, protect their 
mutualists against enemies like herbivores, predators, or parasitoids 
(eg. Pierce et.al., 1987). 


Interactions with ants, termed myrmecophily, are widespread in two 
butterfly families, viz. Lycaenidae and Riodinidae (for reviews see 
Pierce, 1987 ; Fiedler, 1991). In butterflies, myrmecophily mostly occurs 
during the later larval instars and is mediated by specialized exocrine 
glands, the so-called myrmecophilous organs. Among the better stu- 
died family Lycaenidae, interactions with ants range from mutualistic 
to commensalic or parasitic. More than 50% of the 4.500 described 
lycaenid species are myrmecophilous, and in the great majority these 
interactions with ants are believed to be mutualistic (Fiedler, 1991). 
While the caterpillars of certain butterfly species always associate with 
specific host ants and can hardly survive without these (so-called 
obligate myrmecophiles), larval ant-associations are facultative and 
unspecific in most lycaenid species. This means that caterpillars can 
survive without ants, that at any given time varying fractions of a 
caterpillar population are not tended by ants, and that the caterpillars 
are unspecifically tended by a variety of ant genera as represented in 
the local ant fauna (Fiedler, 1991). 


213 


Several types of glandular epidermal organs (Malicky, 1969 ; Cottrell, 
1984 ; Ballmer & Pratt, 1992) mediate, in combination with substrate- 
borne vibratory signals (DeVries, 1990), the associations between lycae- 
nid caterpillars and ants (Fiedler er al., 1996). Among these glands, 
the dorsal nectar organ (DNO hereafter) on the seventh abdominal 
segment is the major source of energetic reward in the caterpillar-ant 
interactions (Fiedler & Maschwitz, 1988a). From their DNO the cater- 
pillars secrete discrete droplets of a secretion rich in carbohydrates 
and/or amino acids (Maschwitz er al., 1975 ; Cushman et al., 1994). 
In addition, caterpillars of many lycaenid species possess a pair of ever- 
sible tentacle organs (TOs) located on the eighth abdominal segment. 
The TOs are most often everted when caterpillars are disturbed or 
while moving, and attendant ants respond with increased activity and 
attention (Henning, 1983 ; Fiedler & Maschwitz, 1988b). The activity 
of the DNO and the TOs can easily be quantified. 


In contrast to the excretions (“honeydew”) which mediate the ant- 
homopteran mutualisms, secretions from the myrmecophilous organs 
of butterfly caterpillars are produced at some metabolic costs. Therefore, 
caterpillars should be under selection to minimise their energetic invest- 
ment into the symbiosis according to actual needs. In concordance 
with this expectation, Leimar & Axén (1993) found that caterpillars 
of Polyommatus icarus adjust their DNO secretion and TO eversion 
rates to their actual tending level and also respond to simulated at- 
tacks with increased secretory activity. Further examples for such 
plasticity of myrmecophilous behaviours in response to larval tending 
levels have been documented from other species (Fiedler & Hagemann, 
1995, Fiedler & Hummel, 1995). This “strategic behaviour” may help 
to minimise costs. In fact, caterpillars of P icarus do not suffer from 
developmental net costs when tended by ants, but even overcompen- 
sate for their investment into myrmecophily : ant-tended males grow 
larger than untended controls (Fiedler & Hölldobler, 1992). Similar 
observations have in the meantime been made on a variety of lycaenid 
species (Wagner, 1993 ; Cushman et al., 1994 ; Fiedler & Saam, 1994), 
indicating that mechanisms to minimise costs are widespread. 


We here address two questions related to this topic. Firstly, how do 
secretion rates change with time when a caterpillar-ant association 
becomes established ? Secondly, how do secretion rates change with 
larval growth? We studied both these questions under two different 
feeding regimes, viz. high quality food conditions and severe food stress. 


214 


Material and methods 


Study organisms 


Butterflies. The Palearctic Common Blue butterfly, Polyommatus icarus 
(Rottemburg, 1775) is widespread in Eurasia and can be bred for a 
number of generations per year in the laboratory. In nature, caterpillars 
of P icarus are facultatively tended by ants belonging to at least 4 genera 
(Lasius, Formica, Plagiolepis, Myrmica) in 2 subfamilies (Fiedler, 1991), 
and some caterpillars are typically found untended (Emmet & Heath, 
1989). Various aspects of ant-interactions of P icarus have been studied 
intensively in the laboratory (Fiedler, 1990, 1991 ; Fiedler & Hölldobler, 
1992 ; Lemar & Axén, 1993; Fiedler & Saam, 1994; Burghardt & 
Fiedler, 1996). Our culture originated from mated females caught in 
the field (origin : northern Bavaria). Females were allowed to oviposit in 
glass cages kept in a greenhouse (for principals of the rearing procedure 
see Schurian, 1989). Eggs were collected daily and transferred to a 
climatic chamber (15 :9 h L :D cycle, 22.5°C constant). After hatching, 
groups of caterpillars were transferred into translucent closed plastic 
vials (125 ml) lined with moist filter paper, where they were raised 
on ad libitum food until they had moulted into the third instar. With 
the beginning of the third instar, the DNO and TOs become functional 
in P icarus, and from that time onwards all experimental caterpillars 
were kept individually until pupation. The caterpillars had contact to 
ants only during the daily experimental period (15 min per individual). 


Caterpillars of P icarus usually feed on inflorescences or young foliage 
of various herbaceous Fabaceae species (Martin, 1984 ; Ebert & Renn- 
wald, 1991), and food quality is known to strongly affect myrmeco- 
phily in that species (Fiedler, 1990 ; Burghardt & Fiedler, 1996). We 
here report on caterpillars under two feeding regimes (see Burghardt 
& Fiedler, 1996 for experiments with further plant species). One group 
(n = 20) was raised on Medicago sativa L. inflorescences through- 
out the whole larval development. This is a natural high-quality food 
for the fast growing caterpillars. A second group (n = 19) was fed 
mature foliage of the same plant species (M. sativa) sampled in the 
same habitat. As expected, this food treatment caused substantial stress 
to the larvae, which pupated at much lower weights (mean + S.E. = 
59.2 = 1.8 mg, n= 19) than on the high-quality diet (91.1 + 3.2 mg, 
n= 20; U-test, p < 0.002). 


Ants. For our experiments we chose Lasius flavus (Fabricius, 1782). 
This widespread ant species of open grasslands feeds mainly on honey- 


215 


dew excreted by specific root aphids, the diet being supplemented by 
surplus aphids to obtain nitrogen (Pontin, 1958, 1978 ; Godske, 1992). 
Because of their highly developed trophobiotic behaviour, L. flavus 
ants readily associate with lycaenid caterpillars in laboratory assays. 
It must be noted, however, that due to their mostly subterranean life- 
habits L. flavus ants rarely tend lycaenid caterpillars in nature (Fiedler, 
1991). 


We kept the ants in their natural earth nests as taken from the field 
(origin : northern Bavaria). They were maintained in large plastic arenas 
(66 cm X 44 cm 12 cm) with a bottom layer (1-2 cm) of plaster of 
Paris. Ants received honey water and dead cut cockroaches as needed. 
We used three different ant colonies for our experiments. 


Experimental procedure. For experiments, we carefully transferred 15 
foraging worker ants with a fine brush from the nest arena to a small 
test arena (a plastic box of 10 cm X 10 cm X 6 cm with a moist bottom 
layer of plaster of Paris). Arena walls were coated with Fluon to prevent 
the ants from escape. The ants were left undisturbed for 5-10 min. 
After that time all signs of alarm behaviour had faded off and the 
ants walked around in the arena in search for food. Then, a single 
caterpillar was placed in the centre of the arena. 


Starting with the first contact between the caterpillar and an ant, we 
determined the number and timing of DNO secretion droplets as well 
as of TO eversions (eversion of one or both tentacle organs were equal- 
ly counted as one event) during a test period of 15 min. After each 
experiment, we noted the age of the caterpillar (in days since hatching 
from the egg) and its weight (determined to the nearest 0.1 mg on 
an electronic Sartorius Basic 61 balance). We also measured the 
diameter (to the nearest 0.05 mm) of as many DNO secretion droplets 
as possible, using a calibrated eyepiece. 


Observations were made under a stereomicroscope at 10fold magni- 
fication under natural daylight and at room temperatures (22-25°C). 
The caterpillars had no access to food during the 15 min trials, but 
were returned to their rearing vials with ad libitum food immediately 
after the end of each experiment. Each set of ants was used for a 
maximum of three subsequent experiments, resulting in a maximum 
time of 60 min being separated from their home colony. There was 
no indication that the behaviour of ants towards the larvae changed 
during this period. Then, the ants were returned to their colony and 
replaced by a new set of 15 foragers. 


216 


To study the development of myrmecophilous behaviours throughout 
the fourth (= final) larval instar, we tested the larvae at various times 
between the moult into the fourth instar and pupation. Because the 
larvae did not grow equally fast, we could not use absolute larval age 
as a reliable basis for comparisons. Under our rearing conditions, most 
larvae passed the final instar in 5.5 d on Medicago sativa inflorescences, 
and in 6.5d when feeding on M. sativa foliage, but on both diets 
some larvae took 1-2 d longer. Therefore, we divided the final instar 
larvae in 4 relative size classes according to their weight at the time 
of testing divided by their final prepupal weight. This gives a dimension- 
less quotient of “relative weight”. We scored the larvae in the following 
classes : class I (relative weight < 0.4 ; corresponding to the first day 
of the fourth instar) ; class II (relative weight 0.4-0.8 ; corresponding 
to the following 1-2 days); class III (relative weight > 0.8 ; corre- 
sponding to the final 1-2 days of the feeding phase of the fourth instar) ; 
and class IV (“non-feeding prepupae”). “Non-feeding prepupae” denotes 
larvae which have stopped feeding and which were characterised by 
the following morphological and behavioural traits : these caterpillars 
are light-greenish, the whitish color markings of the feeding phase have 
vanished, the caterpillars’ cuticle looks glossy, they have left the food- 
plant and hide within the filter paper, and they have excreted their last 
frass pellet, which looks red due to the excretions of the Malpighian 
tubules. Prepupae were only tested as long as they were still able to 
crawl and had not yet spun a silk girdle for pupation. Experimental data 
were analysed using two-tailed non-parametric test procedures (U-tests, 
H-tests, Spearman’s rank correlation : Sachs, 1992), except where stated 
otherwise. 


Results 


Temporal patterns of myrmecophilous interactions 


We noted no significant differences with regard to DNO secretion rates, 
TO eversion rates, or actual levels of ant-attendance between larvae 
of the relative weight classes I-III, 1.e. within the feeding phase of the 
fourth larval instar (see below). Therefore, we present data on the time 
course of caterpillar-ant interactions only for fully-fed mature larvae 
(weight class III) and non-feeding prepupae (class IV). Results for the 
relative size classes I and II are entirely analogous (Burghardt, 1994). 
In both age classes and on both larval diets we observed similar 
temporal patterns in the secretory behaviour of P icarus larvae. DNO 
secretion rates were always much higher in the initial 3 min of an 


217 


interaction than later on (Fig. 1). In mature larvae (class III) reared 
on Medicago sativa inflorescences, the DNO secretion rates dropped 
from 3 droplets/3 min to a constant low level of 0.6 droplets/3 min 
within 3 min. In prepupae the decrease in secretory activity from 7 
droplets/3 min was less steep and reached a constant low level of 2.5 
droplets/3 min only after 9 min. On M. sativa foliage, the temporal 
pattern was identical, but the secretion rates were overall much lower 
(Fig. 1). : 

The frequency of TO eversions followed roughly the same temporal 
pattern as the DNO secretion rates (Fig. 2). TO eversions occurred more 
often in the initial 3 min of an interaction than later on. In addition, 
the proportion of larvae which did not evert their TOs at all increased 
from 24% (flowers) or 33% (foliage) in the initial 3 min interval to 
57% (flowers) or 90% (foliage) later on. As with the DNO secretions, 
the drop in TO activity was pronounced after 3 min in mature larvae, 
but less steep and abrupt in non-feeding prepupae (Fig. 2). 


In our experimental assay with a surplus of 15 ant mutualists available 
in a small arena, actual tending levels of caterpillars showed almost 
no change through experimental time (data not shown; Burghardt, 
1994). Within less than | min after the first encounter, each caterpillar 
was constantly visited by 4-10 ants, and there was neither an increase 
nor a decrease in the tending levels within the 15 min 1 h experiments. 


Age and food effects on secretory behaviour 


DNO secretion rates (Fig. 3) were statistically not different between 
the relative weight classes I-III within the fourth larval instar (Kruskal- 
Wallis test, H = 0.223, p > 0.8 for caterpillars reared on flowers ; H = 
0.886, p > 0.6 for larvae on foliage). In the prepupal phase, however, 
DNO secretion rates increased distinctly in comparison to mature 
feeding larvae (Mann-Whitney U-test, Z = 4.564, p< 0.0001 for 
animals fed flowers ; Z = 5.321, p< 0.0001 for caterpillars reared on 
foliage). 


Largely the same age pattern occurred with TO eversion rates (Fig. 4). 
These were practically constant throughout the feeding phase of the 
fourth instar (Kruskal-Wallis test, H = 1.179, p > 0.5 for caterpillars 
reared on flowers ; H = 0.065, p > 0.9 for caterpillars reared on foliage). 
In flower-fed caterpillars, there was a highly significant increase in TO 
eversion rates from mature feeding to prepupal non-feeding larvae 
(Mann-Whitney U-test, Z = 3.404, p< 0.001). For foliage-fed cater- 
pillars, this same difference was only observed as a statistical trend 


218 


Secretion droplets from DNO 


4 
® Med. sativa leaves Med. sativa flowers 
3 
a 

2 

\ 3 

SST 

IS 
1 SER 

N 

SEN 

RS RSS 

\ Se 

RRS NRG RSR 

‘ Re SS 

| oo 

\ \ SS 
0 SR ESS RS 

0-3 min 3-6 min 6-9 min 9-12 min 


Secretion droplets from DNO 
8 


® Med. sativa leaves Med. sativa flowers 


CLP LP SP ie SALE PEL 


> 


0-3 min 3-6 min 6-9 min 9-12 min 12-15 min 


Fig. 1. Temporal pattern of the delivery of secretion droplets from the dorsal nectar 
organ (DNO) by Polyommatus icarus caterpillars (fourth instars) to Lasius flavus ants 
(means + S.E.). Experiments lasted 15 min. White bars : caterpillars raised on Medi- 
cago sativa foliage (food stress treatment) ; cross-hatched bars : caterpillars fed M. 
sativa flowers (high-quality food treatment). A) mature feeding caterpillars, weight 
class III, relative weight > 0.8 (n = 21 for both food treatments). B) non-feeding 
prepupal larvae, weight class IV (n = 20 for both food treatments). 


TO eversions 


5 4 
[ Med. sativa leaves Med. sativa flowers 
a 
N a 
3 SON 
III 
N 
ISSN 
N 
2 
| 
0 


0-3 min 3-6 min 6-9 min 9-12 min 12-15 min 


TO eversions 


ee Med. sativa leaves Med. sativa flowers 


0-3 min 3-6 min 6-9 min 9-12 min 12-15 min 


Fig. 2. Temporal pattern of eversions of the tentacle organs (TO) by LP icarus cater- 
pillars (means + S.E.). White bars: caterpillars raised on Medicago sativa foliage 
(food stress treatment) ; cross-hatched bars : caterpillars fed M. sativa flowers (high- 
quality food treatment). A) mature feeding caterpillars, weight class III (n = 21 for 
both food treatments). B) non-feeding prepupal larvae, weight class IV (n = 20 for 
both food treatments). 


DNO droplets/ 12 min. 
20 


ey} Med. sativa leaves Med. sativa flowers 


<0,4 < 0,8 > 0,8 non-feeding larvae 


rel. weight category 


Fig. 3. Number of DNO secretion droplets (means + S.E.) delivered by P icarus 
caterpillars (fourth instars) of four relative weight classes during the final 12 min of 
experimental interactions with L. flavus ants. Relative weights are calculated as the 
quotient of actual larval weight at the end of an experiment, divided by the final 
prepupal weight reached by the same individual. The square bracket above the first 
three categories denotes that, on each food treatment, no significant differences were 
observed between the weight classes. Empty bars : caterpillars were fed M. sativa leaves 
(class I: n = 22, class II: n = 22, class III: n = 21, non-feeding prepupae : n = 20). 
Cross-hatched bars : caterpillars raised on M. sativa flowers (class I: n = 12; class II: 
n = 20; class III: n= 21 ; non-feeding larvae : n = 20). 


(Mann-Whitney U-test, Z = 1.434, one-tailed p < 0.08) due to the high 
variance. Nevertheless, overall TO eversion rates were on average twice 
as high in prepupal larvae in this latter series of experiments. 


Size of secretion droplets 


We measured the diameter of 107 DNO secretion droplets delivered 
by fourth instars of P icarus. 74 of these occurred in the initial 3 min 
of an experiment, 33 droplets in the final 12 min. Droplet diameters 
(©) did not differ significantly among larvae in the relative weight classes 
II-IV (mean diameters + S.E.; class II: © = 0.239 + 0.019 mm, 


221 


TO eversions/ 12 min. 


[| Med. sativa leaves Med. sativa flowers 


ee 
2 

3 

2 

x 

2 

3 

2 

x 
2 
= 
ete 
2 

x 
2 
I 
3 

3 

sore 

< 
= 
SE 
de 
Se 
I 
IR 
3 
SE 
= 


RER 


OR 


RSS LÉ 


< 0,4 < 0,8 > 0,8 non-feeding larvae 


rel. weight category 


Fig. 4. Number of tentacle (TO) eversions (means + S.E.) of P icarus caterpillars 
of four relative weight classes during the final 12 min of experimental interactions 
with L. flavus ants. The square bracket above the first three categories denotes that, 
on each food treatment, no significant differences were observed between the weight 
classes. Empty bars : caterpillars were fed M. sativa leaves (class I: n = 22, class II: 
n = 22, class III: n = 21, non-feeding prepupae : n = 20). Cross-hatched bars : cater- 
pillars raised on M. sativa flowers (class I: n = 12; class II: n = 20; class II: 
n = 21 ; non-feeding larvae : n = 20). 


n= 14; class HI: © = 0.258 + 0.012 mm, n= 13 ; class IV : © = 0.252 
+ 0.011, n= 32; Kruskal-Wallis test, H = 0.986, p > 0.6). The com- 
bined average droplet diameter for these three weight classes was 
hence ® = 0.250 + 0.004 mm during the initial 3 min of an inter- 
action, corresponding to a spherical volume of 0.0082 ul. Young 
fourth instars (class I), however, produced significantly smaller droplets 
(© =0.170 mm + 0.008 mm, n = 15 ; Mann-Whitney test, Uj4. 15 = 42, 
p < 0.01), with a spherical volume of 0.0026 ul. 


DNO secretion droplets delivered during the final 12 min of an ex- 
periment were significantly smaller (@ = 0.197 + 0.008 mm, volume 
0.004 ul, n = 33 [4 in class III, 29 in class IV]) than those in the 


222 


initial phase of an interaction (Mann-Whitney U-test, Z = 3.899, 
p < 0.001). These measurements show that larvae of P icarus not only 
reduce the number of DNO secretion droplets, but also droplet size 
as soon as a stable ant-association has been established. 


Discussion 


Our experimental results revealed three major patterns: 1) P icarus 
caterpillars quickly reduce the activity of their myrmecophilous organs 
as soon as a stable association With ants has been established ; 2) DNO 
secretion and TO eversion rates distinctly increase when the larvae reach 
the prepupal stage, yet remain constant throughout the feeding phase 
despite a more than fourfold increase in body mass of P icarus cater- 
pillars ; 3) these temporal and developmental patterns remain unaffected 
if the larvae are reared under food stress, but the amount of secretions 
to be allocated into the mutualism critically depends on the food 
resources available to the larvae (see also Fiedler, 1990 ; Burghardt 
& Fiedler, 1996). 


The temporal patterns of secretory behaviours are concordant with 
the findings of Lemar & Axén (1993) that myrmecophilous caterpillars 
behave “strategically” viz. adjust their DNO secretion and TO ever- 
sion rates to their actual needs. In our experimental conditions, stable 
caterpillar-ant associations established within 1-3 min after the initial 
contact, and after this period there was no need for the caterpillars 
to continue with their potentially costly behaviours at the same high 
rate. Flower-fed mature P icarus larvae spent on average 0.024 ul DNO 
secretions in the initial 3 min compared to 0.009 ul in the subsequent 
12 min. For prepupal caterpillars, the respective estimates are 0.058 ul 
in the initial 3 min and 0.050 ul in the subsequent 12 min. This temporal 
pattern did not depend on the absolute amount of resources available 
to the caterpillars, since we observed it with larvae at all sizes and 
ages, and on the high-quality food (inflorescences) as well as under 
severe nutritional stress (foliage). Furthermore, the same pattern occurs 
in a number of lycaenid butterfly species tested so far under the same 
experimental conditions (Celastrina argiolus: Burghardt, 1994; Zi- 
zeeria knysna : Fiedler & Hagemann, 1995 ; Aricia agestis : Fiedler & 
Hummel, 1995 ; Polyommatus coridon, P. cornelia : Fiedler unpubl.). 
Therefore, this strategy appears to be widespread among butterfly 
species whose larvae live in facultative symbioses with ants. 


DeVries (1988) described the establishment of interactions between ants 
and myrmecophilous caterpillars in the family Riodinidae as an “entice- 


223 


ment and binding” process, where the rewarding with nutritious secre- 
tions is particularly important during the initial phase of an association, 
whereas later on a combined action of all myrmecophilous organs 
ascertains the integrity of the association in time. Our observations 
on a range of lycaenid species, including P. icarus, indicate that a similar 
strategy is prevalent in the butterfly family Lycaenidae as well. To offer 
more and larger droplets after the first contacts between a caterpillar 
and an ant will strengthen the binding between the two partners and 
may even induce the scout ant to recruit additional nestmates (Fiedler 
& Maschwitz, 1988a), whereas later on the secretion rate and droplet 
size are modulated to lower levels. 


Even at the very beginning of an interaction with ants, the caterpillars 
probably do not deliver the complete stored secretion from their DNO 
reservoirs. This conclusion can be drawn when the secretion volumes 
observed in our experiments are compared with morphometric data 
on the size of the DNO in P icarus. According to Malicky (1969 : 
253), each glandular bladder of the DNO ıs 0.72 mm long and 0.2 mm 
wide. The DNO has four such ellipsoidic bladders with a cumulative 
reservoir volume of approximately 0.06 ul. The secretion volumes of 
the initial 3 min in our experiments were distinctly lower than this 
reservoir volume for feeding mature caterpillars (0.024 ul), but almost 
as large (0.058 ul) in prepupae. 


Could the enhanced initial secretion rates be a response to unavoidable 
handling at the beginning of an experiment rather than an adaptive 
trait ? Although handling disturbance might have contributed to the 
pattern observed (cf. Leimar & Axén, 1993), the above calculation 
indicates that at least during the feeding phase larvae retain some 
“currency” for future use. Furthermore, while being constantly attended 
by ants, only 50% of P. icarus larvae responded to tactile disturbance 
with one surplus secretion droplet (Fiedler, unpubl. data), whereas all 
larvae in the experiments detailed above produced on average two 
additional droplets in the initial 3 min interval. Hence, even if response 
to handling did affect our observations, it seems unlikely that this would 
be the sole source of the effect (see detailed discussion in Fiedler & 
Hagemann, 1995). 


Non-feeding prepupal larvae enhance their activity relevant to the 
mutualism for a short period of time (less than 12 h in P icarus). 
Again, this finding is paralleled by results obtained with Zizeeria knysna 
(Fiedler & Hagemann, 1995), Polyommatus cornelia (Fiedler et al., 
1994), P coridon (Fiedler, unpubl.), and Aricia agestis (Fiedler & Hum- 
mel, 1995). Inferring from the significant weight losses of ant-tended 


224 


prepupae, a substantial investment into myrmecophily does probably 
also occur at this developmental stage in the Nearctic Hemiargus isola 
(Wagner, 1993). 


What adaptive significance, if any, may this enhanced prepupal invest- 
ment into myrmecophily have? We suggest that enhanced prepupal 
secretory activity assists to strengthen the caterpillar-ant association 
during the most vulnerable time of the life-cycle. Firstly, prepupae and 
pupae are immobile and thus unable to escape from, or defend against, 
attacks by predators or parasitoids. Secondly, freshly moulted pupae 
have an extraordinarily soft cuticle which renders them highly susceptible 
to fatal damage. Thirdly, all species where enhanced secretory activity 
in the prepupal phase has so far been observed, descend from their 
hostplants for pupation, which then takes place on the soil, in the litter 
or at the hostplant base. In such locations, the likelihood of encounters 
with ground-foraging ants is much higher than on the hostplants. There- 
fore, the prepupae require effective mechanisms to appease ants and 
may gain further selective advantages if ant-associations can be main- 
tained and extended far into the pupal stage (see Fiedler, 1988, and 
Wagner, 1995, for myrmecophily of lycaenid pupae or prepupal larvae). 


In view of earlier experiments on the influence of food quality on the 
ability of P icarus caterpillars to produce DNO secretions (Fiedler, 
1990), it was not surprising that foliage-fed caterpillars were much 
inferior to flower-fed siblings with respect to their myrmecophilous 
properties (see also Burghardt & Fiedler, 1996). Strong effects of host- 
plant quality on the attractiveness of lycaenid caterpillars to ants have 
also been reported for the Australian obligate myrmecophile Jalmenus 
evagoras (Baylis & Pierce, 1991, 1993). The important notion from 
our experiments is that the temporal as well as developmental patterns 
of myrmecophilous behaviours remain unchanged under food stress, 
although the absolute and relative investment into myrmecophily are 
very sensitive to the nutritional environment of the larvae. 


Because we did not quantify the nutrient content in the minute secretion 
droplets, we cannot rule out that changes in droplet size or secretion 
rate might be accompanied by changes in nutrient concentrations. This 
also applies to the experiments of Leimar & Axén (1993). The possible 
variance of food rewards should evidently be studied in the future. 
However, in our experiments we never observed that the ants would 
reject a single secretion droplet. Hence, even if variation in secretion 
quality should have occurred, all droplets were highly attractive to the 
attendant ants. 


22,5 


In our experiments, TO eversion rates almost exactly followed the 
frequency patterns of DNO secretions : an enhanced initial TO activity 
was followed by a rapid decrease, more frequent TO eversions occur- 
red in the prepupal phase, and TO activity was much reduced under 
food stress. Although the exact function of the TOs in lycaenid-ant 
mutualisms 1s still under discussion, there is ample evidence that these 
organs alert and attract tending ants (Fiedler & Maschwitz, 1988b ; 
Ballmer & Pratt, 1992). Fiedler (1991 : 8-29) observed that TO eversion 
rates in P icarus larvae were significantly correlated with the stability 
of their ant-associations. Since the action of the TOs involves the 
production of some communicative signal as well as muscular activity, 
these organs pose an additional energetic cost on the caterpillars, even 
if this cost is likely to be small. It should therefore be adaptive that 
caterpillars restrict TO activity to times when these organs may assist 
in establishing or stabilising caterpillar-ant associations, like at the 
beginning of an interaction or when entering the prepupal phase. Under 
severe food stress, caterpillars should restrict the use of their TOs to 
spare energy, and this is exactly what we observed in our rearings on 
Medicago sativa foliage. In addition, Lemar & Axén (1993) demon- 
strated that TO activity is enhanced when caterpillars are subject to 
a simulated attack. Finally, we observed significant correlations between 
DNO secretion and TO eversion rates across the tested individuals 
(flower-fed prepupae : rs = 0.594, n = 20, p < 0.01 ; foliage-fed pre- 
pupae : rs = 0.584, n = 20, p < 0.05). Collectively, these observations 
suggest that TO eversions may advertise, as an honest signal, the pro- 
fitability of a P icarus larva as nectar resource to its attendant ants. 


Our experimental data reveal that myrmecophilous behaviours of P 
icarus caterpillars show characteristic temporal and developmental 
plasticity, that the patterns emerging from this plasticity are presumably 
adaptive, and that the resources available to the larvae are an important 
constraint on the amount of investment, but do not change the patterns 
of its delivery. These observations are in accordance with previous work 
on strategic behaviour in that mutualism (Leimar & Axén 1993) and 
on developmental costs of myrmecophily (Fiedler & Hölldobler, 1992 ; 
Fiedler & Saam, 1994). Furthermore, the occurrence of similar effects 
in a number of additional facultatively myrmecophilous Lycaenidae 
species also suggest that the patterns observed in P. icarus are of general 
significance for unspecific mutualistic caterpillar-ant interactions, which 
are by far the most widespread type of interactions with ants in the 
butterfly family Lycaenidae (Fiedler, 1991). | 


226 


Acknowledgments 


We thank H.-T. Baumgarten, D. Hagemann, and V. Hummel for assistance 
with caterpillar rearing and help in sampling data. P. Seufert critically com- 
mented on an earlier manuscript version. 


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Syntarucus pirithous (L.) y Polyommatus icarus (Rot.) (Lep., Lycae- 
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MASCHWITZ, U., Wust, M. & ScHurRIAN, K., 1975. Bläulingsraupen als 
Zuckerlieferanten fiir Ameisen. Oecologia 18 : 17-21. 

Pierce, N. E., 1987. The evolution and biogeography of associations between 
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Pierce, N. E., Kircuine, KR. L., BUCKLEY, KR. C., TAYLOR, M. F. J. & BENBow, 
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Pontin, A. J., 1958. A preliminary note on the eating of aphids by ants of 
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Pontin, A. J., 1978. The number and distribution of subterranean aphids 
and their exploitation by the ant Lasius flavus (Fabr.). Ecol. Entomol. 
3 : 203-207. 

SACHS, L., 1992. Angewandte Statistik. 7th edition. Springer Verlag, Berlin/ 
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SCHURIAN, K. G., 1989. Revision der Lysandra-Gruppe des Genus Polyomma- 
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WAGNER, D., 1993. Species-specific effects of tending ants on the development 
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WAGNER, D., 1995. Pupation site choice of a North American lycaenid butter- 
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229 


Nota lepid. 19 (3/4) : 230-242 ; 21.XII.1996 ISSN 0342-7536 


Illustrierter Bestimmungsschlüssel 
für die Präimaginalstadien der Schwärmer Europ 
und Nordafrikas (Lepidoptera : Sphingidae) 


Teil III : Puppen 


Alexander PELZER 


Dorfstr. 20, D-30974 Wennigsen, Bundesrepublik Deutschland 


Summary 


Keys to the preimaginal instars of the hawkmoths of Europe and North Africa 
(Lepidoptera : Sphingidae). Part III : Pupae. 

This is the last of a small series of keys dealing with mature larvae, first 
instar larvae, and pupae. They aim at the identification of living specimens. 
Therefore, all characters that are not visible in the intact animal are omitted. 
It is hoped that the keys will be useful for faunistic purposes and for research 
in parasitoids. 


Zusammenfassung 


Dieser Schlüssel ist der letzte aus einer kleinen Reihe, die sich mit den erwach- 
senen Raupen, den Eilarven und den Puppen befaBt. Sein Ziel ist das Bestim- 
men lebender Tiere. Daher wird auf alle Merkmale, die am lebenden Tier 
nicht erkennbar sind, bewußt verzichtet. Die Schlüssel könnten vor allem in 
der Faunistik und der Parasitoidenforschung von Nutzen sein. 


Resume 


Cette clé est la dernière d’une petite série qui traite des chenilles — adultes et 
au stade Li — et des chrysalides. Elle a pour objectif de permettre la deter- 
mination des espèces vivantes à ces différents stades. Par conséquent ne sont 
pas pris en considération las caractères invisibles chez l’animal vivant. Ces clés 
pourraient rendre service en faunistique et pour l'étude des parasitoides. 


Einleitung 

Europäische Schwärmerpuppen waren schon um die Jahrhundertwende 
vergleichsweise gut bekannt (Hofmann, 1893 ; Spuler, 1908). In den 
letzten Jahrzehnten sind die Puppen weiterer Arten beschrieben worden 


230 


(z.B. Pittaway, 1979 ; Pelzer, 1982 ; Heinig & Häsler, 1986). Dennoch 
gibt es noch keinen Bestimmungsschlüssel für die europäischen Arten. 
Auf die Merkmale, durch die sich verwandte Arten unterscheiden, 
gehen die vorhandenen Beschreibungen nur in Ausnahmefällen ein (z.B. 
Pelzer, 1988). Erst in jiingster Zeit hat Patocka (1993) eine zusammen- 
fassende Darstellung der mitteleuropäischen Arten versucht. 


Von einfachen Bestimmungsschlüsseln für die Präimaginalstadien, mit 
denen sich bereits ein lebendes Tier bestimmen läßt, könnte neben der 
Systematik und der Faunistik besonders die Ökologie profitieren : 
Schlüpfen statt des Falters einige Fliegen oder Wespen, läßt sich die 
Zuordnung Wirt-Parasitoid nur dann herstellen, wenn die Puppe be- 
stimmbar ist. 


Die Bestimmung von Puppen ist in den meisten Fällen recht einfach. 
Manche Gruppen — etwa die Gattungen Hemaris und Hyles —, die 
als Eilarven überhaupt keine Bestimmungsprobleme aufgeben (Pelzer, 
1995), kombinieren als Puppen jedoch eine insgesamt nur kleine Zahl 
verwertbarer Merkmale mit erheblicher innerspezifischer Variabilität. 


Der Schlüssel ist in erster Linie dafür gedacht, lebende Puppen zu 
bestimmen. Leere, vollständige Puppenhüllen oder tote Tiere werden 
sich in der Regel jedoch ebenfalls bis zur Art, mindestens aber bis 
zur Gattung bestimmen lassen. 


Systematik und geographische Abgrenzung 


Die verwendete Systematik wird im 1. Teil der Reihe erläutert (Pelzer, 
1991) ; sie folgt weitgehend der Auffassung von Rothschild & Jordan 
(1903). Der Name Laothoe tremulae (Fischer von Waldheim, 1830) 
wird durch L. amurensis (Staudinger, 1892) ersetzt (Pittaway, 1993). 


Das im Schlüssel behandelte Gebiet umfaßt Europa bis zum Ural sowie 
Nordafrika nördlich der Sahara. Nach dem gegenwärtigen Kenntnis- 
stand treten 32 Schwärmerarten mehr oder weniger regelmäßig in 
diesem Areal auf (Tab. 1). Hinzu kommen einige Irrgäste aus Nord- 
amerika, Afrika und Asien, die hier nicht behandelt werden (Listen 
in Meerman 1987 und Pittaway 1993). Die Merkmale der wenigen 
Hybriden, die aus dem Freiland bekannt sind, vermitteln stets zwischen 
denen der Elternarten. 


Material und Methoden 


Die Schlüssel basieren überwiegend auf eigenen Daten. Sie beruhen 
auf zahlreichen Zuchten und Freilandbeobachtungen der meisten hier 


231 


behandelten Arten. Bei manchen Artengruppen bestätigen meine Ergeb- 
nisse die von Patocka (1993) herausgearbeiteten Unterscheidungsmerk- 
male (z.B. S. ligustri / H. pinastri, M. tiliae | L. populi), bei anderen 
ergeben sich Differenzen (Gattungen Hemaris, Hyles). Patocka unter- 
schätzte offenbar die nicht unerhebliche Variationsbreite mancher 
Merkmale. 


Von fünf Arten lag mir kein Material vor. Es handelt sich dabei um 
Arten, für die Europa auf dem äußersten Rand ihres Verbreitungsgebiets 


liegt. 
Die Aufnahme der Puppe von R. komarovi in den Schliissel stiitzt sich 


auf die Beschreibung von Pittaway (1979), die der Puppe von A. osiris 
auf die Beschreibung von Fawcett (1901). 


PSt 


VK 


Abb. 1. Schematische Darstellung einer Schwärmerpuppe (A. convolvuli, 2 ; ventral 
und lateral) mit den fiir die Bestimmung wichtigsten Merkmalen 


AS : Antennenscheide, ASt : Abdominal-Stigma, GF : Gleitflache der freien Abdominal- 
segmente, HFS : Hinterfliigel-Scheide, X : Kremaster, MF: Milbenfalle, MS : Mittel- 
bein-Scheide, MTF: Metathorax-Feile, PSt : Prothorax-Stigma, RS : Rüsselscheide, 
VFS: Vorderfliigel-Scheide, VH: Vorderbein-Hiifte, VK: Vorderkôrper (-lange), 
VS : Vorderbein-Scheide. 


232 


Abb. 2-6. Puppen europäischer und nordafrikanischer Schwärmer. Sofern nicht anders 
vermerkt, sind die Tiere in Ventralansicht abgebildet. In Klammern hinter dem 
Artnamen der jeweilige Abbildungsmaßstab. 


Abb. 2 a, b) A. atropos (ventral und dorsal, 1,2 X) ; c) A. convolvuli (lateral, 1,5 X) ; 
d, e) S. ligustri (ventral und lateral, 1,5 X) ; f) A. pinastri (lateral, 2,3 X). 


Nach der zur Verfügung stehenden Beschreibung (Hofmann, 1893) 
dürfte die Puppe von S. gorgoniades bis auf die Färbung der von S. 
kuldjaensis sehr ähneln. Die frühen Stände der letztgenannten Art sind 
vor kurzem detailliert beschrieben worden (Zolotuhin, 1994). Dr. Zolo- 
tuhin stellte mir freundlicherweise eine Puppenhiille von S. kuldjaensis 
zur Verfügung. 


Die Puppe von S. caecus soll der von S. ocellatus ähneln (Graeser, 
1888). — Die Puppe von D. elegans ist noch unbeschrieben. 


Die verwendeten Merkmale werden in Abb. 1 definiert. Zur Bestimmung 
ist eine Handlupe mit etwa 10-facher Vergrößerung ausreichend. 


Die Anordnung der einzelnen Arten auf den Tafeln weicht z.T. von 
der in den Teilen I und II ab, um ähnliche Puppen wieder direkt neben- 
einander zu stellen. 


Tab. 1. Schwärmerarten Europas und Nordafrikas. 


Acherontia atropos (Linné, 1758) Totenkopf 

Agrius convolvuli (Linné, 1758) Windenschwärmer 
Sphinx ligustri Linné, 1758 Ligusterschwärmer 

Hyloicus pinastri (Linné, 1758) Kiefernschwärmer 
Dolbina elegans A. Bang-Haas, 1912 

Marumba quercus (Denis & Schiffermüller, 1776) Eichenschwärmer 
Mimas tiliae (Linné, 1758) Lindenschwärmer 

Smerinthus caecus Ménétriés, 1857 

Smerinthus ocellatus (Linné, 1758) Abendpfauenauge 
Laothoé populi (Linné, 1758) Pappelschwärmer 

Laothoë amurensis (Staudinger, 1892) 

Hemaris fuciformis (Linné, 1758) Hummelschwärmer 
Hemaris tityus (Linné, 1758) Skabiosenschwärmer 
Hemaris croatica (Esper, 1779) 

Daphnis nerii (Linné, 1758) Oleanderschwärmer 
Macroglossum stellatarum (Linné, 1758) Taubenschwanz 
Proserpinus proserpina (Pallas, 1772) Nachtkerzenschwärmer 
Rethera komarovi (Christoph, 1885) 

Sphingonaepiopsis gorgoniades (Hiibner, 1819) 

Hyles lineata (Fabricius, 1775) Linienschwärmer 

Hyles gallii (Rottemburg, 1775) Labkrautschwärmer 
Hyles euphorbiae (Linné, 1758) Wolfsmilchschwärmer 
Hyles nicaea (Prunner, 1798) Nizzaschwärmer 

Hyles centralasiae (Staudinger, 1887) 

Hyles zygophylli (Ochsenheimer, 1808) 

Hyles hippophaes (Esper, 1789) Sanddornschwärmer 
Hyles vespertilio (Esper, 1779) Fledermausschwärmer 
Deilephila elpenor (Linné, 1758) Mittlerer Weinschwärmer 
Deilephila porcellus (Linné, 1758) Kleiner Weinschwärmer 
Hippotion celerio (Linné, 1758) Großer Weinschwärmer 
Hippotion osiris (Dalman, 1823) _ 

Theretra alecto (Linné, 1758) 


234 


Abb. 3 a) M. tiliae (2,3 X); b) L. populi (2,3 X) ; c) L. amurensis (2,1 X) ; d) S. ocel- 
latus (2,0 X) ; e) M. quercus (1,6 X). 


255 


Schlüssel 


Die verwendeten Begriffe werden in Abb. 1 erläutert. Das Kürzei in 
Klammern hinter dem Artnamen ist die jeweils zugehörige Abbildungs- 


nummer. 
1. Rüsselscheide deutlich kürzer als die Flügelscheiden (Abb. 3a-e) : PTE 2 
1’. Rüsselscheide mindestens so lang wie die Flügelscheiden 6 
2. braun, von stumpfer Farbe ; obere Kutikulaschichten abblätternd wie 
Kirschbaumrind® ........ ann ae. IIS 3 
2. schwarz oder sehr dunkel braun ; Kutikula wie poliert glänzend .......... 5 
3. Rüsselscheide so lang wie die Mittelbein-Scheiden (Pfeilspitze) .............. 
SNARE RON RAA «4 de duo RR: à à cas 73, SSSR M. tiliae (3a) 
3’. Rüsselscheide kürzer als die Vorderbein-Scheiden (vergl. Abb. 3b : Pfeil- 
RY DUA |e ee oc ee nn cuove renoue eee EE 4 
4. Kremaster Außenkante glatt Ikonkay.............. L. populi (3b) 
4’. Kremaster-Außenkante mit einem Absatz (Pfeilspitze), verjüngt sich 
daher In 2 Stuten... ee leet eee L. tremulae (3c) 
5. Kutikula der Flügelscheiden glatt, lackglänzend ; Vorderbein-Hüfte nicht 
SICH Dale en een S. ocellatus (3d) (auch S. caecus ?) 
5. Kutikula der Flügelscheiden runzelig ; Vorderbein-Hüfte sichtbar ........... 
ee ee Bee en M. quercus (3e) 
6. Rüsselscheide vorn über die Augen hinaus kiel- oder henkelförmig ver- 
längert (verge Abb. 22 01 Lt RER, RE 7 
6’. Rüsselscheide vorn nicht verlängert, höchstens blasig vorgewölbt (vergl. 
Abb. 68) ee NS <5 14 
7. Rüsselscheide henkelförmig verlängert (vergl. Abb. 2c)....................... 8 
7. Rüsselscheide kielförmig verlängert (vergl. Abb. 6f) .......................... 10 
Rüsselscheide bildet eine Spirale mit einer 3/4-Windung ......................... 
N RER H. convolvuli = 
8. Rüsselscheide viel kürzer RE... PR ee... 
9. dunkelbraun ; Vorderbein-Hüfte sichtbar (Pfeilspitze) ; Rüsselscheide es 
nur am Ende aufleeende........ me... en... S. ligustri (2d, e) 
9. schwarzbraun ; Vorderbein-Hüfte nicht sichtbar ; Rüsselscheide fast in 
ganzer Länge dem Thorax aufliegend 7.2... 2... H. pinastri (2f) 
10,,..einfarbig. schwarz bra © 5,04: 22... ee ee... R. komarovi 
10’. Grundfarbe weißlich bis braun, mit braunen und/oder schwarzen Zeich- 
nungsmüusternn ae 201 ANAS ana De 11 
11. klein; Grundfarbe weißlich ; Rüsselnaht auf ganzer Länge schwarz ; 
Vorderbein-Hüfte nicht sichtbar .............................. M. stellatarum (4d) 
11’ 


236 


. Grundfarbe braun ; Vorderbein-Hüfte z.T. unauffällig, aber sichtbar .. 12 


IN 
N 
FIN 
N 
À 
N 
N 
NN 


_ 


Abb. 4 a) A. tityus (2,8 X) ; b) H. fuciformis (2,8 X); c) H. croatica (Sommer- 
puppe, 2,1 X) ; d) M. stellatarum (2,4 X) ; e) D. nerü (1,4 X) ; c)P. proserpina (2,3 X). 


237 


12. 


12’. 
13. 
13’. 
14. 


14. 
15. 


15. 


16. 


16’. 
17. 


17% 


18. 


18’. 


19. 


19’. 


20. 


20’. 


238 


Rüsselscheide um höchstens etwa !/2 Augendurchmesser verlängert ....... 
na dos BE ee dea Me LC wos ara RE Ce eee H. celerio (6e) 


Riisselscheide um mindestens 1 Augendurchmesser verlängert ........... 13 
Rüsselscheide um etwa 1 Augendurchmesser verlängert ..... T. alecto (6f) 
Rüsselscheide um etwa 2 Augendurchmesser verlängert ............ H. osiris 
Abdomen seitlich mit kräftigen Stachelsäumen (vergl. Abb. 6c, Pfeil- 
SPÜZEN). neue NN Rene en N ER 15 
Abdomen ohne Stachelsaumer tres. Zee a 16 
klein (Vorderkörper <25 mm) ; Stachelsäume deutlich caudal der ab- 


dominalen Stigmen ; Abdomen unterseits mit dunklen Winkelzeichnun- 
gen (<>) und/ oder hellen Ringen an den Stellen, an denen bei der Raupe 
die Bauchbeine saßen ; Augengrate sehr scharf ............ D. porcellus (6c) 
groß (Vorderkörper > 25 mm) ; Stachelsäume verlaufen durch die abdo- 
minalen Stigmen ; Abdomen unterseits mit verwaschenen Zeichnungen, 
die keine Winkel bilden ; Augengrate unauffällig ........... D. elpenor (6b) 


walzenförmig, erst auf den letzten Abdominalsegmenten plötzlich ver- 
jüngt ; Grundfarbe gelb-bräunlich, Rüsselnaht und Stigmen kontrastie- 
TENdSCHWALZ ET. ce nue ee e EE D. nerii (4e) 
Anders ........ ns. nk done te EN EEEEE 17 


mit Metathorax-Feile (runzelig-rauhe Fläche auf dem Metanotum, Pfeil- 
spitze) ; sehr groß (Vorderkörper > 40 mm) ; Rüsselbasis mit deutlichen 


Querrunzeln. es else A. atropos (2a, b) 
ohne Metathorax-Feile, auf dem Metanotum höchstens glatt-wulstige 
Gebilde. se denses Mensen nennen RE EEE 18 


fast einfarbig braun (auch schwarz?) ; klein bis sehr klein (Vorderkörper 
< 20 mm) ; Vorderflügelscheiden an der Außenseite deutlich eingebuchtet 
(Abb, 4: Peilspitze) : Kremaster lang und spitz 22 u 7 77 19 
ANG CLS) ote ga ..ecnessuonseeeenaeereeenhnanie ten she ce RE IRRE 20 


klein (Vorderkörper knapp 20 mm) ; Milbenfalle ohne Grate ................. 
NER... een RO os RACE Moe ds dem be onde. eee P. proserpina (4f) 
sehr klein (Vorderkörper < 15 mm) ; Milbenfalle mit ca. 3 scharfen Gra- 
UST Sc AUS se Br... RE 2 (S. gorgoniades) 


klein ; Grundfarbe dunkelbraun bis schwarz, evtl. weißlich bereift ; Gleit- 
flächen der freien Abdominalsegmente heller als Grundfarbe des Ab- 
domens ; Kremasteransatz etwa so breit wie das letzte Abdominalsegment 
un ie oc „Luke ehe ee ee ee Hemaris spp. 21 
mittelgroß bis sehr groß ; Grundfarbe entweder hell- bis mittelbraun oder 
verdunkelt (braun bis schwarz), dann aber stets auch mit dunkler Zeich- 
nung ; nie bereift ; Gleitflächen der freien Abdominalsegmente (nur bei 
lebenden Tieren !) dunkler als Grundfarbe des Abdomens ; Kremaster 
viel schmaler als das letzte Abdominalsegment ................. Hyles spp. 23 


Mi 


’ 
2 


7 X) 
3 X) 
239 


I, 
CL, 


2 


1Caea 


XO) 3 ©) lab 


2 


7X); b) H. gallii (helle Form 
H. lineata (2 


2 


d) 


jae (dunkle Form, | 


Abb. 5 a) H. euphorb 
(dunkle Form, 1,8 X) 
ER): 


c) H. zygophylli 
f) A. centralasiae (1 


21. 


21”. 


22, 


227. 


23. 


23’. 


24. 
24’. 
25. 


23%. 


26. 


26’. 


27. 


Ia 


28. 


28’. 
29. 


29’. 


240 


Flügelscheiden bis auf die Aderung glatt; braun, glänzend (Sommer- 
puppen) oder weißlich bereift (Winterpuppen) ............... H. croatica (4c) 
Flügelscheiden runzelig ; schwarzbraun bis schwarz, nie bereift ......... 22 


Prothorax-Stigma mit + elliptischem, glattem Hof, nur etwa zur Hälfte 
verdeckt ; die „Hörnchenplatte* am Vorderende (Pfeilspitze) nur ein 
flaches, abgerundetes Plättehen 22... 22,2 H. fuciformis (4b) 
Prothorax-Stigma mit schmalem, + nierenförmigem Hof, fast völlig ge- 
schlossen ; die „Hörnchenplatte“ am Vorderende ventral in eine Spitze 
AUSGEZOgENN...2..40.40nn ee ea ren RP UE CUS H. tityus (4a) 


relativ schlanke Puppen (Verhältnis Vorderkörperlänge zu Vorderkör- 
perbreite > 2,5) ; hell bis mittelbraun, ohne grobe dunkle Zeichnungs- 
MUSLET „ce neseenneesnanensn case rennen ee see ee tr 24 
relativ plumpe Puppen (Verhältnis Vorderkörperlänge zu Vorderkör- 
perbreite < 2,5) ; sehr hell bis fast schwarz, meist mit auffälligen dunklen 


Zeichnungsmusterm. 7.004002... 0000 CI 27 
blaB gelblich bis hell beigebraun, Fliigelscheiden oft durchscheinend ... 25 
mittelbraun, Flügelscheiden nicht durchscheinend ............................. 26 


mittelgroß (Vorderkörper < 25 mm) ; Kremaster gleichmäßig in eine lange 
Spitze ausgezogen ; Rüssel- und Beinscheiden ohne dunkle Runzelzeich- 
DNS I. nu. a er les eee H. lineata livornica (5d) 
groß bis sehr groß (Vorderkörper > 35 mm); Kremaster abgesetzt, in 
zwei Stufen in eine kurze, stumpfe Spitze auslaufend ; Rüssel- und Bein- 
scheiden mit feiner, dunkler Runzelzeichnung (Lupe !) ..... H. nicaea (Se) 


Rüsselscheide blasig aufgewölbt ; Rüssel- und Beinscheiden ohne dunkle 
Runzelzeiehnung.......... en H. vespertilio (6a) 
Rüsselscheide nicht blasig aufgewölbt ; Rüssel- und Beinscheiden mit 
dunkler Runzelzeichnung ren a H. hippophaes (6d) 


hellbraun ; Flügelscheiden ohne Zeichnung, durchscheinend ................... 
RER RE Re loasnsne: H. centralasiae (Sf) 
Grundfarbe variabel, weißlich bis schwärzlich (nur selten hellbraun) ; 
Flügelscheiden auch bei hellen Formen mit bräunlicher oder schwärzlicher 
Runzelzeichnung, niemals durchseheinend” 22.2222. 20 ee 28 


Milbenfalle am 6. Abdominalsegment (Pfeilspitze) kleiner und schwächer, 
ansonsten aber ähnlich wie am 5. Abdominalsegment gebaut .................. 
BEER ERBEN AONE ee ee H. gallii (5b) 
6. Abdominalsegment ohne Milbenfalle...........2.2.. ee 29 


+ matt ; Runzelzeichnung setzt sich auf den Außensaum der Vorderflügel- 
scheiden fort ; Kremaster meist abgesetzt, in zwei Stufen in eine kurze, 
stumpfe Spitzeauslaufend ca nc. ee H. euphorbiae (5a) 
lackglänzend ; Runzelzeichnung dehnt sich nicht auf den Außensaum 
der Vorderflügelscheiden aus ; Kremaster gleichmäßig in eine lange Spitze 
BUSTEZOBEN Hana ee Se EE H. zygophylli (Sc) 


+ 
N \ 
\N 


NN 


Abb. 6 a) H. vespertilio (1,6 X); b) D. elpenor (1,7 X); c) D. porcellus (2,1 X); 
d) H. hippophaes (1,7 X) ; e) H. celerio (lateral, 1,7 X) ; f) T. alecto (lateral, 1,2 X). 


241 


Dank 


Mein Dank gilt nochmals Herrn Dr. E.A. Loeliger (Oegstgeest [NL]) für 
Lebendmaterial zahlreicher Arten und Herrn E. de Bros (Binningen [CH]) 
für die französische Übersetzung der Zusammenfassung. Herrn Dr. V.V. Zolo- 
tuhin (Uljanovsk [RUS]) danke ich für die Überlassung einer Puppenhiille 
von S. kuldjaensis und Frau I. Paas (Bochum) für die fotografischen Abzüge. 


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der Schwärmer Europas und Nordafrikas (Lepidoptera : Sphingidae). 
Teil I : Erwachsene Raupen. Nota lepid. 14 : 220-233. 

PELZER, A., 1995. Illustrierter Bestimmungsschlüssel für die Präimaginalstadien 
der Schwärmer Europas und Nordafrikas (Lepidoptera : Sphingidae). 
Teil II : Eilarven. Nota lepid. 17 : 141-154. 

Pitraway, A. R., 1979. On Rethera komarovi manifica [sic] (Brandt) (Lepi- 
doptera : Sphingidae). Entomol. Gaz. 30 : 3-6 + Taf. 1. 

Pirraway, A. R., 1993. The hawkmoths of the Western Palaearctic. Martins : 
Harley Books. 

ROTHSCHILD, W., JORDAN, K., 1903. A revision of the lepidopterous family 
Sphingidae. Novit. Zool. 9, Suppl. 

SPULER, A., 1908. Die Schmetterlinge Europas. Stuttgart. 

ZOLOTUHIN, V.V., 1994. To the biology and morphology of Sphingonaepiopsis 
kuldjaensis Graeser, 1892. Atalanta 25 : 245-259. 


242 


Nota lepid. 19 (3/4) : 243-260 ; 21.X11.1996 ISSN 0342-7536 


Breeding of an F, of Hyles hybrid vespertilioides 
(Boisduval, 1827) after induction with ecdysone 
of metamorphosis in diapausal pupae 


E. A. LOELIGER* & F. KARRER** 


* Hofdijck 48, NL-2341 ND Oegstgeest, The Netherlands 
** Rebbergstrasse 5, CH-4800 Zofingen, Switzerland 


Summary 


After synchronisation of the emergence of male and female adults of Hyles 
hybrid vespertilioides Bdv. by means of ecdysone injected into four of the 
six female pupae, pairing took place quickly. Oviposition was abundant and 
survival was normal. Fertility of the eggs averaged 50 per cent. Ovipositing 
females as well as about 65 per cent of the larvae preferred Epilobium dodo- 
naei Vill. as host. Phenotypically, F,-larvae and F,-imagines varied marked- 
ly : from the typical A. vespertilio (Esper, 1779) to the almost classical A. 
hippophaes (Esper, 1793). There was, however, a striking correlation between 
host preference and phenotype : caterpillars displaying typical grandparental 
aspect choose the grandparental host and transformed into imagines strongly 
resembling the respective grandparent. Particularly notable among larvae 
raised on Hippophae rhamnoides L. and/or Elaeagnus angustifolia L. were 
those mimicking the uncommon recessive variety of H. hippophaes displaying 
rubiginous colouration and among imagines those closely resembling the rare 
ab. flava Blach. of H. vespertilio. 


Résumé 


Apres synchronisation de l’éclosion des adultes mâle et femelle de Hyles hy- 
bride vespertilioides Bdv. au moyen d’ecdysone injectée en quatre des six chry- 
salides femelles, l’accouplement eut lieu rapidement. La ponte était abondante 
et le degré de survie normal. La fertilité des œufs était d'environ 50 pour 
cent. Les femelles qui pondaient ainsi qu’approximativement 65 pour cent 
des chenilles préféraient Epilobium dodonaei Vill. comme plante-hôte. Le 
phénotype des chenilles F, et des imagos F, variait de façon marquée : de 
H. vespertilio (Esper, 1779) au H. hippophaes (Esper, 1793) presque classique. 
Il y avait néanmoins une corrélation frappante entre la préférence de l’hôte 
et le phénotype : des chenilles arborant l’aspect grandparental typique choi- 
sissaient la plante-hôte grandparentale et se transformaient en imagos res- 
semblant fortement au grand-parent respectif. Particulièrement remarquables 


243 


c 


Fig. 1. Three L; caterpillars, 1 a and 1 b being specimens of F, H. hybrid vespertilioides, 
1 a closely resembling vespertilio, and 1 b that of 1 c which is the propositus of the rare 
rubiginous variety of an hippophaes L; found in the field (Loeliger, 1996). 


244 


Fig. 2. L, caterpillars of F, H. hybrid vespertilioides displaying uncommon appearance, 
2a, 2b, and 2c grown on H. rhamnoides/E. angustifolia and 2 d, 2e, and 2f on 
E. dodonaei/angustifolium. 245 


parmi les chenilles élevées sur Hippophae rhamnoides L. et/ou Elaeagnus 
angustifolia L. étaient celles imitant la variété récessive rare de H. hippophaes 
arborant une coloration rougeatre et parmi les imagos ceux ressemblant de 
près la rare ab. flava Blach. de H. vespertilio. 


Introduction 


Hyles hybrid vespertilioides (Boisduval, 1827) is the product of crossing 
a Hyles hippophaes (Esper, 1793) male with a Hyles vespertilio (Esper, 
1779) female, two closely related species of the sphingid family which 
are practically monophagous and highly ubiquitous in the riverine 
valleys of the northern Mediterranean region. Hybridization must occur 
regularly under field conditions since hybrid caterpillars have been 
found repeatedly along the Rhône River (Boisduval, 1827 ; Feisthamel, 
1827; Denso, 1909; Wenczel, 1983). Hybridization has also been 
obtained in captivity (Benz, 1952 ; Baumgartner, 1971, pers. comm. ; 
personal experience on repeated occasions in the eighties). Reports on 
a second generation (F,), 1.e., the cross vespertilioides x vespertilioides, 
have not yet been published although an F, has been obtained at least 
once (Benz, 1962, unpublished). The adults produced and diary records 
are in the possession of one of us (FK). 


Our interest in an F, generation arose from observations of the here- 
ditary transmission of grandparental peculiarities, such as the complete 
lack of the caudal horn of H. vespertilio caterpillars or the conspicuous 
colouration of adults of the two species involved. Investigation of a 
possible correlation between host preference and phenotype of the cater- 
pillar and/or ovipositing adult also seemed worthwhile. In addition, 
the possible de novo appearance of rare varieties of the grandparental 
species continues to be one of the incentives to undertake hybridization 
experiments. 


The attempt to breed an F, generation of vespertilioides was complicated 
by the fact that metamorphosis of the females is known to be markedly 
retarded (Baumgartner, 1971, pers. comm ; personal observations in 
1971 and 1973), although it may occur simultaneously with that of 
the males (Benz, 1962). 


In this paper the achievement of successful synchronization of the emer- 
gence of male and female adults of vespertilioides by means of the 
injection of ecdysone into female pupae is reported and our observations 
during the breeding of an F, generation of this hybrid are described. 


246 


Materials and methods 


EXPERIMENTAL ANIMALS : the male imago of the hybrid vespertilioides 
cross was an exceptionally strong specimen of H. hippophaes, grown 
in 1993 as the offspring of the mating of a H. hippophaes caucasica 
(Denso, 1913) male, the caterpillar of which was found on Elaeagnus 
angustifolia near Bogazkale (TR) in October 1992, with a H. hippophaes 
hippophaes female originating from the Swiss part of the Rhône Valley 
(courtesy F.Weber, Riehen, CH). This female had produced 680 mainly 
fertile eggs (dissection revealed that only eight eggs had remained in 
situ). The mother of the vespertilioides cross was one of the many 
H. vespertilio-specimens grown in 1993 under field conditions (the F, 
female lived for 4!/2 weeks and laid all of its 250 eggs, exclusively 
on Epilobium dodonaei Vill., the vast majority being fertile). The grand- 
parental vespertilio caterpillars were found near Les Baux in southern 
France (courtesy C. Czipka, Fiirth-Erlangen, D). 


FLIGHT CAGESs : the rectangular wooden floor, 30 x 50 cm, 1s capped 
with a tunnel of gauze, e.g., surgical TubinettelM, stretched over the 
floor and three aluminium arches which have a maximum height of 
30 cm and are attached at regular intervals to the floor along its length. 


A cage generally does not contain more than one or two females plus 
two to three males. The long side is exposed to daylight and, if possible, 
there is some circulation of fresh air. With sprigs of the host along 
the bright side of the cage, the cages also serve as receptacles for ovi- 
position. 


PREPARATION AND INJECTION OF THE ECDYSONE SOLUTION : one milli- 
gram of commercially available alpha-ecdysone crystals is dissolved in 
0.12 ml ethanol or isopropanol. To obtain the full sterilizing capacity 
of the alcohol, 0.08 ml of water are added. Ten minutes later, 0.8 ml of 
water are added to obtain the final alcohol concentration of 12 per cent 
and the final ecdysone concentration of 0.1 per cent. The ingredients 
are mixed thoroughly and the solution is aspirated with a 30 G Micro- 
lance™ needle (Beckton, Dickinson & Comp.) into a one-ml-syringe 
subdivided into 0.02 ml portions (e.g. a Jecton-S™ syringe), each 
portion thus containing 20 mg ecdysone and amounts of diluted alcohol 
too small to damage pupal tissue. 


The optimum amount of hormone to be injected, as assessed from 
the results of treatment of more than 300 specimens, is approximately 
six mg per gramme pupa. Before injection, the vertex of the head of 


247 


Fig. 3. Five imagines resembling H. vespertilio ab. flava Blach., the specimen shown 
in Figure 3 a grown by EAL, and the four displayed specimens shown in Figure 3 c 
belonging to the collection of FK. The specimen presented in Figure 3 a is the same 
as that on display to the bottom left in Figure 4c; the bottom right specimen in 
Figure 3 b is the same as that among the months shown in Figure 4 b. 


248 


Fig. 4. (a) presents two imagines resembling hippophaes (top) and vespertilio (bottom). 
L; of the latter is presented in Figure | a. 
(b) shows the four imagines of the 1994 breeding by FK. The two smaller 
bottom specimens are samples of the Benz collection. 
(c) Overview of the nine imagines of the 1994 breeding by EAL (the 10th, 
a specimen of the flava variety with coiled forewings is not shown). 


the pupa is iodinized. To prevent abdominal movement of the pupa, 
which would increase internal pressure such that blood loss from the 
puncture site would be heavy, the pupa is anaesthetized by exposure 
for 30 to 40 minutes to CO,; for this purpose, the pupa is placed 
in a container on a two to three cm thick layer of linen covering small 
pieces of dry ice. Before the injection, the pupa is laid on the middle 
finger and fixed with the thumb. The needle is inserted along the sagittal 
axis through the iodinized vertex of the head deep into the thorax/ 
abdomen. The predetermined amount of the ecdysone solution is then 
injected slowly. The needle is kept in situ for about 10 seconds. After 
withdrawal of the needle, the puncture site is covered with collodion 
and the pupa is placed on fine peat or sand to avoid unnecessary move- 
ments which may be caused by coarse underlying material. Twenty- 
four hours after injection the pupa can be handled normally. 


REARING THE LARVAE: before use the containers, mainly Petri dishes, 
were thoroughly washed and sterilized with formaldehyde. Every 24- 
48 hours the leaves were changed. Larvae, twigs and leaves were only 
handled after thorough hand-washing ; they were deposited on clean 
sheets of paper. 


Results 


The hybrid vespertilioides : on June 20%, 1994, shortly after midnight, 
copulation took place between a H. hippophaes male (two males had 
been in the cage) with a H. vespertilio female (one adult in the cage 
only). Room temperature was 22°C. Oviposition on E. dodonaei Vill. 
started the day after pairing. By June 30th, a total of 284 eggs had 
been deposited exclusively on the host. Subsequent dissection of the 
female revealed that no eggs were left in the abdomen. The caterpillars 
first grew on E. parviflorum Schreber, until stage L,. Stage L; were 
fed E. angustifolia L. and exposed to a constant temperature of 35°C. 


By July 125, the first larva prepared for pupation, and by July 28*, 
there were 13 male and six female pupae. On July 26‘, the first and 
second male pupae exhibited signs of metamorphosis. By August 2nd 
and 34, five male imagines had emerged ; all other males were still 
in the process of development. The six females remained in diapause. 
In order to induce their metamorphosis, ecdysone was injected into 
four pupae on August 5, the other two pupae were left untreated 
to serve as controls. The injected pupae were maintained at 35°C. 
Developing male pupae were kept at the lowest possible room tem- 
perature ; emerged males were exposed to continuous light to minimize 
flying. By August 8th, eye pigmentation, the first sign of metamorphosis, 


250 


was discerned on translucence, and by August 12th, wing pigmentation 
had become apparent. In the afternoon of August 14%, the four 
imagines emerged almost simultaneously, the forewings being slightly 
deformed in three cases after filling with blood and their scaling being 
more or less defective (unscaled wing patterns display a green glimmer). 
The probosces were insufficiently coiled in two cases. None of the four 
females exhibited overt problems as far as flight, oviposition and feeding 
were concerned. 


The two non-treated female pupae remained in diapause. 


Pairing of vespertilioides : The two first-born females (emergence time 
about 16.00 h and 18.30 h, respectively) were kept in separate cages 
together with two males for each. Both copulated on the same day 
shortly after midnight. Pairing lasted one and three hours, respectively. 
The other two females (emergence time about 19.00h and 20.00 h, 
respectively) copulated with different males the next day, also shortly 
after midnight, with similar pairing times. 


OvirosiTion : the first egg was deposited during the night of August 
15th. After eight hours of incubation at 32°C, signs of embryogenesis 
could be discerned microscopically. During the night of August 16‘, 
imago I deposited more than 100 eggs mainly on E. dodonaei. These 
eggs were shipped to collectors and used to obtain an appropriate 
number of caterpillars for observation (n = 50 ; see page seven). From 
August 18th through August 22th, the possibility of a preference for 
one of the parental hosts was investigated. For this purpose, each female 
was kept in a separate cage and the host was changed daily. The first 
night (August 18) H. rhamnoides or E. angustifolia and E. dodonaei 
were presented together (females I and III, Table 1). 


Table 1 


Number of eggs deposited either on the host or on the gauze of the cage. 
The numbers in parentheses refer to the four females separately. 
— : not present during that night. 


Host E. dodonaei H. rhamnoides ‘gauze 
E. angustifolia 
126 (19, 18, 35, 54) 5 ( 8,-, 17, -) SD (Il, 16, 10 &) 
53 (32, 0, 12, 41) SS) (( 9, 30, 1, 13) 


39 ( 3,22, 2,1) = Beh NE) 
39 (18,7,11, 3) | 129 (2, 63, 53,11) 
94 (17, 25, 19, 33) 146 ( 7,100, 14, 25) 


259 (39, > 56,99) | 149 68, 2 40,44) | 457 (40, 2 107, 60) 


251 


Fig. 5. (a) Overview of the 32 imagines raised by Benz in 1962. The numbers run. 
vertically from top to bottom and left to right. The four vertical columns 
contain 8 imagines each. Numbers 2, 3, 4, 6, 13, 18, 21 and 31 (together 
25 per cent) resemble hippophaes, while numbers 11, 12, 16, 17, 20, 22 

52 and 23 (together 22 per cent) are similar to vespertilio. 
(b) shows the four most hippophaes-like specimens (nrs. 2, 3, 13, 18). 


Fig. 6. Adults of the cross vespophaes male x vespertilioides female. Note the novel 
phenotype in the centre of the top row. The two adjacent specimens closely resemble 
hippophaes as did their caterpillars. 


Fig. 7. The seven adults of vespophaes F, in the collection of FK. None is typically 
vespertilio-like. Four resemble specimens of vespophaes F,. The two in the mid-row 
are hippophaes-like, the left one with a highly conspicuous right/left difference. The 
seventh (bottom right) resembles, although only slightly, the novelty shown in Figure 6. 


252 


From the data presented in Table 1 it appears that E. dodonaei was 
preferred for oviposition : the number of eggs deposited on this host 
was about twice that deposited on H. rhamnoides or E. angustifolia. 
Interestingly, the percentage eggs on gauze increased rapidly after the 
first two days. 


The number of eggs deposited separately by the four females was 268, 
630, 309, and 296, respectively, or a total of 1503. The imagines died 
(1) or were killed (IJ, III, IV) 10, 24, 21, and 15 days after emergence. 
On dissection, they contained not more than 2, 0, 0, and 3 eggs, 
respectively. Imago I, which died spontaneously 10 days after emergence, 
had deposited 2/3 of her eggs within three days of pairing. In contrast 
imago II (the female which deposited the most eggs and lived the 
longest) slowly increased the number deposited up to 125 during the 
night of day seven (22/8). The antipathy of this female for H. rham- 
noides and/or E. angustifolia was also striking. 


The imagines were fed a 10 per cent saccharose solution every second 
day. 


PECULIARITIES OF LARVAE AND PUPAE OF THE F, : approximately 75 per 
cent of the eggs deposited by females I and II were fertile versus 25 per 
cent of those from females III and IV. When exposed only to E. dodo- 
naei, about 50 per cent of the larvae were still alive after a fortnight ; 
of those exposed exclusively to E. angustifolia or H. rhamnoides, only 
50 per cent were alive after the first week. Unfortunately, some of 
the L, caterpillars escaped from the Petri dishes, thus excluding exact 
statistics in this respect. 


The length of the caudal horn varied greatly, from being non-existent 
(as in vespertilio) to full length (as in hippophaes). Observation of more 
than 200 larvae revealed that, irrespective of the host, 14 per cent had 
virtually no horn, 36 per cent had a small horn, 36 per cent had a 
horn of medium length, and the remaining 14 per cent had a horn 
about as long as customary for hippophaes larvae. 


Marking and colouration of the caterpillars varied enormously particular- 
ly in phases L; - L;, from a close resemblance to vespertilio (Fig. 1 a) 
to almost indistinguishable from hippophaes, especially the rare rubi- 
ginous variety (Fig. | b and c). However, many of the larvae were also 
similar to the parent hybrid. The majority of the larvae reared on A. 
rhamnoides/E. angustifolia clearly tended to resemble hippophaes, 
whereas the phenotype of larvae growing on E. dodonaei/angustifolium 
looked much more like vespertilio. On several occasions we observed 
accelerated growth of an insufficiently developed vespertilio-like cater- 


254 


pillar when the host was changed from E. angustifolia to E. dodonaei 
or E. angustifolium. Several caterpillars were striking due to their totally 
unfamiliar appearance (Fig. 2). 


CORRELATION BETWEEN HOST PREFERENCE, PHENOTYPE OF THE LARVA 
AND THAT OF THE IMAGO: fifty of the eggs deposited by female I on 
August 164/17" were chosen for raising and observing F, specimens 
under carefully controlled conditions. They were exposed to 33°C and 
a mixture of the grandparental hosts. By August 21%, about one day 
after hatching, 23 larvae had selected E. dodonaei as their host, 14 
H. rhamnoides, and five had not yet made a choice. Eight did not 
hatch. Three days later, 29 surviving larvae were counted: 19 on E. 
dodonaei and 10 on H. rhamnoides. By August 28, 27 larvae 
remained : seven L;, eight L, and three L; on EF. dodonaei, and three 
L;, five L, and one L; on E. angustifolia/ H. rhamnoides (due to a 
shortage of leaves, L; were fed on E. angustifolia and E. angustifolium, 
respectively). On day 10, the first larva ceased feeding and initiated 
pupation. Caterpillars on E. dodonaei pupated one to two days earlier 
than those on E. angustifolia, mainly because growth of the mature 
larvae had decreased temporarily when the host was changed. In total 
23 larvae pupated : 16 of those grown on E. angustifolium (one slightly 
crippled) and 7 of those on E. angustifolia. 


Within a fortnight of pupation, 10 of the 23 pupae (four males and 
six females) underwent metamorphosis, eight having been grown on 
E. dodonaei/angustifolium and two on H. rhamnoides/ E. angustifolia. 
On September 14", the first imago out of the vespertilio-like caterpillars 
of the Epilobium group emerged. After expansion, its forewings 
remained strongly coiled, and its hindwings were yellow instead of red, 
fitting the phenotype of A. vespertilio ab. flava Blach. This flava-type 
variety emerged from four other pupae of this series. The four in the 
possession of FK are shown in Fig. 3 b, in which the specimen to 
the left bottom is the same as that of Fig. 4 b. Eight of the nine cater- 
pillars of the flava-type imagines had been fed on Epilobium. Of the 
five raised by and kept in the collection of EAL only two are shown 
(Fig. 3 a and 4 c). At least five of the caterpillars of the flava type 
had been striking pale yellow tinged green. A conspicuous golden glitter 
of the grayish or brownish scales of the imago was characteristic of 
this de novo variety as was particularly obvious in one case (Fig. 3 a, 
which is the same as the left bottom specimen in Fig. 4 c). Of the 
remaining five caterpillars raised by EAL on E. dodonaei/angustifolium, 
the one that resembled vespertilio the most as caterpillar (Fig. | a) 
gave rise to a slate-grey moth conspicuously similar to its grandmother 


255 


(Fig. 4 a, and Fig. 4 c, top left). The other four had many features 
in commom with their parents, 1.e., vespertilioides F,. Of the two A. 
rhamnoides/ E. angustifolia-fed larvae, the one resembling hippophaes 
the most yielded a moth which closely resembled a pale specimen of 
hippophaes (Fig. 4 a, and Fig. 4 c, bottom right) its larva having been 
olive-green with two pairs of orange ocelli, one at the base of the small 
caudal horn ; it also displayed an orange midline, as so often observed 
on the L; of hippophaes caterpillars. The other imago resembling 
hippophaes (Fig.4 c, midst specimen) emerged from a caterpillar which 
had been green, too, with orange ocelli from head to horn, the latter 
having been of intermediate length. 


Figs. 4 b and 4 c depict F, imagines from the collection of FK and 
EAL, respectively, the FK specimens exclusively grown on E. dodonaei. 
The two vespertilioides-like bottom imagines in Fig. 4 b belong to the 
much smaller specimens of the Benz-collection. The three top specimens 
shown in Fig. 4 c resemble vespertilio as did their caterpillars. 


MENDELIAN TYPE DISTRIBUTION OF THE GRANDPARENTAL IMAGINAL 
PHENOTYPE AMONG SPECIMENS GROWN EXCLUSIVELY ON E. DODONAEI : 
the F, imagines raised by Benz in 1962 were never reported on, probably 
because of insufficient documentation of the breeding. As a supplement 
to our results, however, their presentation may provide additional 
evidence of Mendelian type inheritance of grandparental phenotypes, 
particularly since all of his imagines were from larvae reared exclusively 
on E. dodonaei. (From the records we know that only a few of the 
freshly hatched caterpillars temporarily accepted H. rhamnoides as their 
host). If one studies Figure 5 a , one might be tempted to divide the 
32 imagines very roughly into three categories: one hippophaes-like 
(25 per cent), another vespertilioides-like (53 per cent), and a third one 
vespertilio-like (22 per cent). In Figure 5 b, the four most hippophaes- 
like specimens are presented. 


Discussion 


The present paper is the report of successful breeding of an F, of the 
hybrid vespertilioides in numbers large enough to assess (1) hereditary 
transmission of grandparental morphological and biological peculiarities 
as well as specific markings and colouration ; (2) the possible correlation 
between larval and adult phenotypes ; and (3) the occurrence of de 
novo varieties. Imagines of an earlier successful breeding of this hybrid 
(Benz, 1962, unpublished ; Fig. 5) underscore our findings. 


256 


The success of our breeding is attributable primarily to the excellent 
health of the animals: the grandfather hippophaes was one of the 
numerous “fresh blood” offspring of the mating between a male 
specimen from Anatolia and a female specimen from Switzerland ; the 
grandmother vespertilio was raised in the field, its parents were found 
in southern France. Success would, however, not have been achieved 
without ecdysone-induced synchronization of the emergence of the 
parental males and females. In spite of the fact that the caterpillars 
were exposed to high temperatures, none of the females underwent 
metamorphosis shortly after pupation together with the males. This 
well-known disparity is not necessarily a hybrid-bound phenomenon 
since preferential subitaneous development of exclusively male imagines 
in pupae of H. euphorbiae deserticola caterpillars grown at elevated 
temperatures has also been observed (FK, 1973 ; not published). 


As soon as it became evident that the females would remain in diapause, 
appropriate amounts of ecdysone were injected into four of the six 
female pupae. From large experience we knew that it would take about 
10 days to complete metamorphosis if the injected pupae were stored 
at 35°C. This time it took 11 days. The accuracy of the hormone 
dosage had been such that the four females emerged almost simul- 
taneously. Only one displayed no flaws. But the slight distortion and 
incomplete scaling of the forewings of the other three did not per- 
ceptibly interfere with their biological activities although flying might 
have been hampered slightly, which would explain the rapid drop in 
the percentage eggs deposited on the host relative to the gauze. The 
excellent vitality of the ecdysone-treated females was demonstrated by 
the rapid and uncomplicated pairing as well as the abundance and 
completeness of oviposition. 


To evaluate the mode of inheritance of grandparental morphological 
characteristics, caudal horn length seemed to be the most appropriate 
factor. Interlarval differences were gradual, from non-existent to full 
length. There was no clear host dependence, although some of the 
caterpillars with complete absence of the horn looked like typical 
vespertilio (Fig. 1 a) and those with the longest horns often resembled 
hippophaes (Fig. 1 b and c). 


Hereditary transmission of grandparental biological behaviour, such as 
feeding at night or hiding during daytime, which is so typical of the 
L, of vespertilio, or open sunning in the L; of hippophaes, would have 
been difficult to investigate. A much easier factor to assess was imaginal 
and larval host preference for oviposition and feeding, respectively. 
According to the figures presented in Table 1, the preference for E. 


DST 


dodonaei for oviposition was obvious : twice as many eggs were de- 
posited on this host compared to H. rhamnoides/E. angustifolia. On 
days when only the latter plants were available, about 20 per cent 
more eggs were deposited on gauze than on days when E. dodonaei 
was available. 


About two-thirds of the caterpillars selected E. dodonaei out of a 
mixture of the grandparental hosts. Among those confronted with only 
one type, the percentage acceptance was initially high ; but the 50 per 
cent survival time for those on E. dodonaei was about twice that for 
larvae on H. rhamnoides. Unfortunately, exact figures cannot be 
presented since an unknown number of freshly hatched caterpillars 
escaped from the Petri dishes. 


Hereditary transmission of marking and colouration was fascinating. 
Among caterpillars (Figs. 1 and 2), diversity was almost infinite, such 
that a classical Mendelian type of grouping into grandfather, parental, 
and grandmother larval phenotypes would have been a mere fabrication, 
although many specimens displayed the rather vague markings and 
beige-greyish colouration exhibited by the parent caterpillars and the 
resemblance of quite a number of larvae with the grandparental pheno- 
type was conspicuous (Fig. 1 a and b). There was a preponderance 
of olive-green and ochreous brown in larvae grown on A. rhamnoides/ 
E. angustifolia. Several exceptions to this rule exhibited retarded 
growth, which accelerated after switching hosts. 


Among adults, phenotypic identification was easier. About half resemble 
the parent H. hybrid vespertilioides. Of the remnant, some are very 
similar to hippophaes, others to vespertilio. Most interesting is the fact 
that the more the imago resembles the grandparental phenotype, the 
closer the caterpillar had done the same. This is beautifully illustrated 
by the imagines presented in Figure 4 a: the caterpillar of the upper 
moth had displayed all features typical for hippophaes, while the 
caterpillar of the lower specimen looked very much like an L; of 
vespertilio (Fig. 1 a). Among the excellently preserved imagines of 
vespertilioides F, grown by Benz in 1962 (see Fig. 5 a) about 25 percent 
each resembled the two grandparental phenotypes. The four most 
resembling hippophaes are presented separately in Figure 5 b. The rest 
was vespertilioides-like. It is noteworthy that Mendelian segregation 
appears to be independent on the host caterpillars had been fed with, 
since Benz had all raised on E. dodonaei exclusively. Unfortunately, 
Benz had made no attempt to correlate larval with original phenotype. 
Our observation of such a correlation, therefore, must be considered 
novel. It contradicts observations and interpretations made earlier with 
F, hybrids (Fischer, 1931). 


258 


Most interesting, finally, was the occurrence of specimens which closely 
resembled rare varieties of the grandparental species. Among the 
caterpillars, there were the diffusely rubiginous ones, 1.e., those whose 
colour was complementary to the classical olive-green of hippophaes. 
Caterpillars with such colouration are very rarely found in the field. 
Experimentally, it proved to be an autosomal recessive trait (Loeliger, 
1996). Among the adults, those with yellow hindwings are striking. 
Whether these specimens are identical to the ab. flava Blach. of vesper- 
tilio referred to in handbooks remains to be detected. Specimens found 
in the field might, at least in part, be the offspring of primary or 
secondary hybrids occurring under natural circumstances. 


A thorough study of the reports of the first hybrid caterpillars of 
vespertilioides observed near Grenoble (Boisduval, 1927; De Feist- 
hamel, 1827) reveals that those found by Boisduval might, at least 
partly, have been secondary bastards or perhaps even F, vespertilioides 
(Kysela, 1908). This is not a far-fetched supposition since at the end 
of the 19th century, primary and secondary bastards of Hyles hybrid 
epilobii have been found in the field (Mory, 1901 and 1903 ; Lippe, 
1902). 


ADDENDUM 


In 1995, we were successful with the breeding of H. hybrid vespophaes 
(Denso, 1909 ; Benz, 1948) and its F, , and with that of the cross of 
a vespophaes male with one of the two non-injected females of vesper- 
tilioides. Of both these broods, the results were closely similar to those 
described above, confirming our conclusions. There were two differences, 
however. 


First, not a single specimen of the flava variety was observed among 
the 10 and 16 adults obtained from these F, hybrids, respectively. 
Second, an apparently novel phenotype emerged, particularly obvious 
among the seven adults obtained by EAL of the vespophaes x vesper- 
tilioides hybrid, the large specimen in the centre of the top row 
presented in Fig. 6. On closer inspection, the forewings of this novelty 
appear to resemble those of the flava variety of the F, of vespertilioides. 
Particularly notable is the white instead of darkbrown spot. The 
specimen to the bottom right displays resemblance to this novel-type 
variety, as does the bottom right specimen of the seven F, adults of 
vespophaes grown by FK (Fig. 7). Among the latter, note especially 
the highly uncommon right/left colour difference presented by the mid- 
left specimen. 


259 


For the two additional references and the two additional figures see 
under the respective headings. 


Acknowledgements 


We are much indebted to Mrs G. P. Bieger-Smith for the correction of English. 


References 


BAUMGARTNER, A., 1971. Rundschreiben über eine erfolgreiche Zucht von 
H. hybr. vespertilioides. Not officially published. 

Benz, F, 1948. Zur Kenntnis zweier Schwärmerhybriden. Celerio hybr. 
vespophaes Denso und Celerio hybr. galivornica Kunz u. Grosse. Nach- 
richten des Entomlogen-Vereins Basel und Umgebung 5 : 5-8 ; 10. 

BENz, F., 1952. Celerio hybr.vespertilioides. Mitt. ent. Ges. Basel 2 : 71. 

Bots-DuvaL, M.J.-A., 1827. Sphinx vespertilioides. Ann. Soc. Linn. Paris 
6: 114-116. 

DE FEISTHAMEL, M., 1827. Description d’une nouvelle espece de Sphinx, 
nommée Sphinx Amelia. Bull. Sci. Nat. Geol. 11 : 162. 

Denso, P., 1909. Katalog der Schwärmerhybriden. Bull. Soc. lép. Genève 
123203522 

Denso, P., 1913. Celerio hippophaes. Deutsche ent. Zschr. Iris 27 : 22-45. 

FISCHER, E. , 1931. Artbastarde von Schmetterlingen und ihre F, - und Rück- 
kreuzungsgenerationen. Viertelj.schr. Naturf. Ges. Zürich 76 : 214-302. 

KYsELA, E. , 1908. Beitrag zur Kenntnis der Deilephila-Hybriden. Mitt. ent. 
Ver. Polyxena 2 : 63-82. 

Lippe, G., 1903. Deil. hybr. epilobii B. und die von ihm abgeleiteten secundären 
Bastarde hybr.eugeni My. und hybr.lippei My.. Ent. Zschr. 15 :38. 
LOELIGER, E. A. , 1996. Rubiginous larvae of Hyles hippophaes (Esper, 1793), 
an autosomal recessive variety (Lepidoptera : Sphingidae). Nota lepid. 

18 (3/4) : 304-305. 

Mory, E. , 1901. Ueber einige neue schweizerische Bastarde des Sphingiden- 
Genus Deilephila und die Entdeckung abgeleiteter Hybriden in der 
Natur, sowie Beschreibung einer neuen Varietät von Deilephila vespertilio 
Esp.. Mitt. schweiz. ent. Ges. 10 : 333-360. 

Mory, C. E. , 1903. Revision der bis jetzt bekannten von hybr. epilobii B. 
abgeleiteten Bastarde. Mitt. schweiz. ent. Ges. 10 : 460-469. 


260 


Nota lepid. 19 (3/4) : 261-263 ; 21.X11.1996 ISSN 0342-7536 


Short communication — Kurze Mitteilung — En bref 


Breeding status and range expansion of Danaus chrysippus (Linnaeus, 
1758) in the Algerian Sahara (Lepidoptera : Nymphalidae Danainae) 


B. SAMRAOUI 


University of Annaba, 4, rue Hassi-Beïda, Annaba, Algeria 


Summary 


The breeding status of Danaus chrysippus (Linnaeus, 1758) in the Algerian 
Sahara is confirmed and its larval foodplant, Calotropis procera, is reported 
on. Moreover, the species has apparently increased in numbers and considerably 
expanded its range over the last decades. 


Résumé 


La reproduction de Danaus chrysippus (Linnaeus, 1758) dans le Sahara algérien 
est confirmée, la plante-hôte étant Calotropis procera et, sur base de données 
antérieures, l’extension de sa répartition géographique est estimée. 


Key words : Danaus chrysippus, Calotropis procera, Sahara, reproduction, 
range expansion 


The Plain Tiger or African Monarch, Danaus chrysippus (Linnaeus, 
1758), is a most widespread butterfly occurring in Asia, Australia and 
the whole of Africa. As is often the case with migratory butterflies, 
no subspecies have been accepted, but three forms or morphs are 
recognized : chrysippus, alcippus and dorippus. Studies of the poly- 
morphism of D. chrysippus have led to suggestions that the species 
might have originated in the Far east with subsequent colonisation 
of Africa. The geographical variation of the polymorphism of the 
species stems from ecogeographical adaptations of pre-existing morphs 
(Pierre, 1980). 


Previous papers (Samraoui et al., 1992 ; Samraoui, 1993 ; Tennent, 1995) 
have summed up our knowledge of the presence and of the migrations 
of the Plain Tiger in North Africa and in particular in the Algerian 
Sahara. Both forms (chrysippus and alcippus) are known to occur on 


261 


the Canary Islands (Owen & Wiemers, 1992) and mainland North 
Africa. The Plain Tiger’s populations from Fuerteventura were qualified 
as relicts that are now adapted to the endemic Caralluma burchardii 
(Owen & Wiemers, 1992). The last authors also pointed out the dif- 
ferences between the f. alcippus arising from monomorphic populations 
of West Africa and that existing in the polymorphic population of 
Uganda. The f. alcippus is rare in Algeria but seems more abundant 
in Morocco (Tennent, pers. comm.). It is probable that the Moroccan 
f. alcippus populations derive from West African populations and this 
possibility certainly exists (Samraoui, 1993). Sporadic migrations of 
the f. alcippus in North Africa (Tunisia and Algeria) have been reported 
(Chnéour, 1954) and can be found in the data compiled by Pierre (1973). 


In the Central Sahara (Tassili N’Ajjer and Hoggar), the form chrysip- 
pus is common while the alcippus form is very rare but present. In 
the oasis of Djanet, we have collected larvae and pupae of the Plain 
Tiger on Calotropis procera, thus confirming its breeding status in 
the Algerian Sahara. It leaves open the question of the foodplant in 
the northern oases (Touggourt, Laghouat), where C. procera is rather 
rare if not inexistent (Samraoui et al., 1992). Previously overlooked 
references point to a range. expansion of D. chrysippus as revealed 
by the following statement : “Danaus chrysippus, Vespéce éthiopienne 
commune, s'étend au nord jusque dans la région méridionale du bassin 
de l’oued Mya, mais n’atteint ni Touggourt ni Ghardaïa” (Rothschild, 
1922). It is noteworthy that Chnéour (1948) did not record the Plain 
Tiger in Tunisia nor did the French expedition of 1949 to the Tassili 
N’Ajjer (Rungs, 1958), so it might safely be assumed that the species 
has substantially increased in numbers. Following the numerous 
colonies that managed to gain a foothold in Europe (Bretherton, 1984 ; 
Martin & Gurrea, 1988 ; King, 1991), it is probable that the expansion 
is Slowed down by the availability of suitable larval foodplants. We 
do not know the reason for this range expansion and it 1s interesting 
to point out that several species of insects (Samraoui, in prep.) have 
also pushed up further their northern limit. We might speculate that, 
in the event of global warming, D. chrysippus might well gain a firm 
foothold in Europe (Owen, 1991). 


References 


CHNEOUR, A., 1948. Contribution à l’étude des Macrolépidoptères de Tunisie 
(suite). Bull. Soc. Hist. Nat. Afr. N. 39 : 70-96. 

CHNEOUR, A., 1954. Macrolépidoptères de Tunisie. I, IL. Rhopalocera, Grypo- 
cera. Bull. Soc. Sci. nat. Tunis. 7 : 207-239. 


262 


Kina, G., 1954. The Spanish whereabouts of the Monarch and Plain Tiger. 
Bull. amat. Ent. Soc. 50 (374) : 48. 

MARTIN, J. & GURREA, P., 1988. Establishment of a population of Danaus 
plexippus (Linnaeus, 1758) (Lep.: Danaidae) in Southwest Europe. 
Entomologist’s Rec. J. Var. 100 : 163-168. 

Owen, D. F., 1991. Can Danaus chrysippus (L.) (Lepidoptera : Danaidae) 
establish itself in Europe ? Entomologist’s Gaz. 42 : 37-39. 

Owen, D. F. & Wiemers, M. 1992. The butterflies of Fuerteventura. 
Entomologist’s Gaz. 43 : 87-92. 

PIERRE, J., 1973. Etude du polymorphisme chez Danaus chrysippus L. (Lépi- 
doptère Danaïde) : aires de répartition et catégories infraspécifiques en 
Afrique. C.r. Séanc. Acad. Sc. Paris 276 : 2685-2687. 

PIERRE, J., 1980. Variation géographique du polymorphisme et du mimétisme 
de Danaus chrysippus et d’Hypolimnas misippus (Lépidoptères Rho- 
palocères) en Afrique et en Asie. C.r. somm. Séanc. Soc. Biogéogr. 486 
(1978) : 179-187. 

ROTHSCHILD, W., 1922. L’Algérie et sa faune ; an address read before the 
entomological society of London, Annual Meeting 18th. January 1922. 
Bull. Soc. Hist. Nat. Afr. N. 13 : 146-154. 

Runes, C. 1958. Lépidoptères du Tassili N’Ajjer. Inst. Rech. Sahar. Univ. 
Alger. 3 : 167-176. 

SAMRAOUI, B., 1993. Migration of the African Monarch Danaus chrysippus 
(L.) and the African Migrant Catopsilia florella (Fab.) in Mauretania 
(Lepidoptera : Danaidae, Pieridae). Nota lepid. 16 (1) : 68-70. 

SAMRAOUI, B., BENYACOUB, S. & MENAI, R., 1992. Danaus chrysippus (L.) : 
Possible breeding status and new sightings (Lepidoptera, Danaidae). 
Nota lepid. 14 (4) : 348-350. 

TENNENT, J., 1995. Danaus chrysippus Linnaeus, 1758 ; a review of records 
and present status in the Maghreb countries of Morocco, Algeria and 
Tunisia (Lepidoptera, Danainae). Nota lepid. 17 (3/4) (1994) : 201-216. 


263 


Nota lepid. 19 (3/4) : 264-267 ; 21.X11.1996 ISSN 0342-7536 


Book reviews — Buchbesprechungen — Analyses 


A Catalogue of the type-specimens of Parnassius in the Zoological 
Museum of the Kiev University (Lepidoptera, Papilionidae). Vadim 
V. TSHIKOLOVETS. 75 pages. Editeur Yu. P. NEKRUTENKO. Publié par 
l’auteur V.V. Tshikolovets, P.O. Box 650/5 UA-252032 Kiev Ukraine, 
Nov. 1993. En anglais, broché 16,5 X 26 cm. 


Nous avons reçu de l’auteur un exemplaire de son Catalogue en anglais (édité 
a Kiev par le Dr. Yuri P. Nekrutenko, membre de la Soc. Eur. Lepid.). Nous 
tenons a signaler 4 tous nos membres, en particulier aux amateurs de Parnas- 
siinae, une telle nouveauté. Il s’agit en effet d’un heureux début puisque l’on 
cherche ainsi à donner aux chercheurs du monde entier un accès direct au 
riche matériel-type des collections de l’ancienne Union Soviétique, reflet de 
la grande tradition entomologique slave, matériel resté trop longtemps à l'écart 
des sources d’information universelles. L’intention est même d'étendre les 
publications de ce genre à d’autres familles, d’autres genres et d’autres musées 
(sous le titre : A Catalogue of the type-specimens of the Lepidoptera). 


Dans les pages du Catalogue proprement dit, on trouve les renseignements 
de base sur les types de 129 espèces («group taxa») de Parnassiinae, soit au 
total 460 specimens, tels qu’ils ont été désignés dans les collections et/ou 
présentés comme types dans les descriptions originales. Les noms créés à 
l’origine comme infrasubspécifiques sont également compris. L’auteur espère 
pouvoir publier une partie illustrée : planches avec les specimens types de 
tous les taxa mentionnés, ce qui est malheureusement impossible pour le 
moment vu les difficultés techniques et financières. 


Les taxa se suivent dans l’ordre alphabétique, avec la référence a la description 
originale, le status a l’origine et, si nécessaire, les changements ultérieurs de 
rang et de status. Pour chaque spécimen, les données sont tirées directement 
des étiquettes. Les publications les plus importantes en rapport avec le matériel- 
type sont citées pour chaque taxon sous «Additional published data». 


Les 49 pages du Catalogue proprement dit (p. 9-57) sont suivies de trois appen- 
dices : 


1. Checklist des espèces de Parnassiinae (group taxa) par ordre systématique 
(Bryk, 1935) (6 pages). 

2. Liste des specimens types de Parnassiinae acquis par le Zool. Mus. Kiev 
Univ. (ZMKU) (2 pages). 

3. Liste des specimens types espérés au ZMKU mais qui ne s’y trouvaient pas 
(1 page). 

Le travail se termine par 8 pages de références bibliographiques. 

Emmanuel DE Bros 


264 


Die Tagfalter der Tiirkei unter Berücksichtigung der angrenzenden 
Länder. Gerhard HESSELBARTH, Harry van OorscHot & Sigbert 
WAGENER. )22,5,.cm. X 35..cm,, vols. 1 and.2,: 1354.p.,,.75, text: figs, 
21 tables, 2 colour maps, 36 colour plates (incl. 306 figs), General Part 
(Geography, Geology, Climate, Vegetation, Ecology, Biogeography, 
Glossary), Special Part (Introduction, Abbreviations used, Checklist, 
Systematic Part (Hesperiidae, Papilionidae, Pieridae), Lycaenidae, in- 
cluding a contribution by K. Fiedler : Associations of lycaenid butterflies 
with ants in Turkey), Nymphalidae, Unconfirmed Records and Potential 
Migrants)), Locality List, Collectors List, Bibliographic References, 
Indices of Zoological and Botanical Names) ; Vol. 3 : 847 p., 128 colour 
plates (imagines), 13 black and white plates (eggs, larvae and pupae), 
IV + 342 distribution maps, Selbstverlag Sigbert Wagener, Hemdener 
Weg 19, D-46399 Bocholt, 1995, DM 780,00. 


This monumental work is the outcome of almost 20 years of intensive field 
work (over 60 trips), study of huge quantities of material from private and 
museum collections and bibliographic research, and the final result entirely 
fulfills the high expectations one was having in waiting such a long time for 
the publication of this long-awaited monograph. 


The General Part, entirely written by Sigbert Wagener, sets the scenery in 
which the subsequent parts of this work should be placed, reviewing the 
geography, the geological history and the climate of Turkey from the Tertiary 
up to the present, whereby data from various sources are reviewed and 
compiled into a comprehensive and exhaustive treatise on the subject. This 
is followed by equally interesting reviews on the flora and phytogeography 
of Turkey and on the ecology of Turkish butterflies. The last chapter of this 
part treats the biogeography of the Turkish butterflies, from resp. the choro- 
logical (faunal elements), ecological and historical (presumed Pleistocene 
refugia) view-points. In separate headings, special attention is paid to the 
butterflies of the Aegean Islands and Cyprus (their distribution is summarized 
on Table 27 on pp. 1110-1113 of Vol. 2; unfortunately, several errors and 
omissions have been noted on the latter table). A brief glossary closes the 
General Part. 


The introduction to the Special Part deals a.o. with the genese of the work 
and with taxonomic and nomenclatorial conceptions. A checklist precedes 
the actual Systematic Part : 345 species and, including subspecies, 416 taxa, 
were recognized from Turkey by March 1995. The subsequent 950 pages 
(pp. 159-1109) treat each of these individual taxa in considerable detail, not 
only reviewing existing knowledge, but especially adding a wealth of new data 
not only on taxonomic insight, though also on their bionomics (including 
e.g. the first-ever publication of several chromosome numbers) and distribution. 
For over 70 taxa, original rearing data and descriptions of early stages are 
presented for the first time. The text part is illuminated with 36 colour plates, 


265 


showing photographs of an exceptional quality of biotopes and adult living 
butterflies, their eggs, larvae and pupae. 


This work is no less than an epoch-making taxonomic revision of a good deal 
of the West Palaearctic butterflies, whereby the situation in adjacent countries 
(Balkans and Greece, former U.S.S.R., Iran, Iraq and the Levant) is brought 
into the discussion. No new species-group taxa are described in this book, 
but a great many subspecies (and sometimes even species) are newly sunk 
in synonymy, beside new type-designations and new or revised taxonomic 
combinations. The authors have always used the material itself as the very 
basis for their taxonomic evaluations and the resulting text found its way 
from often memorable, passionate and mutually enriching discussions with 
fellow colleagues. As such it is really the result of a balanced analysis of 
problems, the fruit of team-work in the very best sense of the word. On more 
than one occasion, the authors have given up presenting a solution to a 
problematic situation rather than offering a false answer. In such cases (perhaps 
the most fascinating passages of the whole work !) the problem is exposed 
in its full extent, in the present state of knowledge, in order to enable any 
future reviser to build on it. 


The third volume is for the biggest part taken up by the 128 colour plates, 
showing for each taxon material of both sexes (in the very most cases on both 
upper- and underside), often in good series in order to illustrate significantly 
their geographical and individual variation. These plates are, without exag- 
geration, of a fantastic quality ! They are followed by a series of black-and- 
white plates of equally high standard, showing SEM photographs of (part of) 
eggs, larvae and pupae. The distribution in Turkey of the individual taxa 
is shown on 342 maps, each symbol reflecting the occurrence of each individual 
taxon within a 10 X 10 km grid. 


This is, without the slightest doubt, one of the finest works on lepidopterology 
ever written so far : a milestone that forces humility, respect and admiration. 
No doubt anyone will not agree with each and every taxonomic conclusion 
arrived at for one or another single taxon. This certainly in no way diminishes 
the value of this work, but rather adds to its interest. The authors deserve 
our warmest congratulations and feelings of gratitude for offering the 
lepidoptertists’ community this marvellous achievement. The only regret that 
I can formulate is the choice of German instead of English as the language. 
The latter option would certainly have been more appropriate for giving it 
the widest possible distribution. The price of 780,- DEM is democratic 
considering what this book offers and anyone who is interested in lepidop- 
terology simply has to read it. If you are somewhat financially limited (no 
abnormality these days) and you can only afford to buy one single butterfly 
book within the next few years, this is the one ! 

Alain OLIVIER 


266 


Le Sesiidae della fauna italiana (Lepidoptera). Memorie Soc. Ent. ital., 
Genova, 70 (1991) : 279-312. Carlo PRoLA & Sergio BEER. 


A la fin de 1991 est sorti, dans les «Memorie della Societa entomologica italiana 
di Genova», un long article écrit par deux lépidoptérologistes italiens très 
connus et très actifs, Carlo Prola et Sergio Beer, concernant une famille de 
lépidoptères généralement peu étudiée, les Sesiidae. Il s’agit là d’un important 
travail faunistique et en particulier d’une vraie révision du groupe entier pour 
ce qui concerne l’Itahe. L'article vise à fournir seulement, comme du reste 
l’indiquent clairement les auteurs eux-mêmes, un résumé des progrès fait par 
la Sesiologie italienne durant les quelque soixante-dix années qui nous séparent 
de l’œuvre de Ragusa, c’est-à-dire de la publication du seul catalogue italien 
qui concerne les Sesiidae. L’exposé des taxa est précédé d’une introduction 
historico-bibliographique approfondie et intéressante qui fait le point sur le 
progrès des connaissances du peuplement italien de cette famille si peu étudiée 
et de plus fournit des informations sur ses vicissitudes systématiques et nomen- 
clatoriales si compliquées, à partir de De Prunner (1798) jusqu'aux travaux 
de Laëtüvka (1982-1990), dont la classification a été utilisée par les auteurs. Les 
Sesiidae sont aujourd’hui compris dans la superfamille Sesioidea dans laquelle 
on considère aussi, parmi les autres, les familles des Choreutidae et des Bracho- 
didae. 


Dans ce travail, les auteurs traitent 52 espèces de Sesiidae, appartenant aux 
genres Sesia, Paranthrene, Synanthedon, Bembecia, Pyropteron, Synansphecia 
et Chamaesphecia. Pour chaque espèce on a eu le soin d’indiquer les syno- 
nymies désormais stabiles, la distribution, les plantes nourricières, la période 
de vol et, enfin, la liste des localités de capture. 


En 1995, toujours dans les «Memorie della Società entomologica italiana», 
au vol. 73, les mêmes auteurs ont publié un deuxième travail sous le titre 
«I feromoni in Lepidotterologia e per la conoscenza delle Sesiidae italiane». 
Dans cette nouvelle contribution, Prola et Beer nous présentent un essai qui 
est le fruit de leur connaissance approfondie en matière de communication 
entre les organismes animaux, dont ils font une synthèse claire et complete. 
Grace a la mise a leur disponibilité d’un certain nombre d’ attractifs chimiques 
spécifiques pour les Sesiidae, fournis par le Max Planck Institut für Verhaltens- 
physiologie de Seewiesen (Allemagne), les auteurs ont eu la possibilité d’inté- 
grer les données sur le peuplement italien de cette famille. 


L'usage d’attractifs chimiques pour la capture de lépidoptères est déjà beau- 
coup effectué à l'étranger dans les domaines de la faunistique et de la systé- 
matique, tandis qu’en Italie cet usage se cantonne surtout au domaine de 
la lutte biologique en agriculture appliquée. 


Suite à cette nouvelle contribution, la faune italienne des Sesiidae augmente de 
six espèces non encore signalées de la péninsule italienne, ce qui amène le total 
des espèces y recensées actuellement a 58. De plus, il a été possible d’ac- 
croitre considérablement le nombre de données sur le peuplement de bon 
nombre d’autres espèces, jusqu’à présent considérées très rares ou du moins 


assez localisées. : : 
Pietro PASSERIN D’ENTREVES 


267 


Nota lepid. 19 (3/4) : 268-270 ; 21.XII.1996 ISSN 0342-7536 


Vol 19 — 1996 


Dates of publication — Publikationsdaten — Dates de publication 


19 (1/2): 21.X1.1996 pp. 1-140 
19 (3/4): 21.XIL.1996 pp. 141-270 


Contents — Inhalt — Sommaire 


BURGHARDT, F. & FIEDLER, K. : Myrmecophilous behaviours in 
caterpillars of the butterfly, Polyommatus icarus (Rottem- 
burg, 1775) : temporal patterns and age dependency (Lepi- 
doptera : Lycaenidae) 

CEBOLLA, M. — cf. STEFANESCU, C. 

DE Prins, R. — cf. OLIVIER, A. 

FIEDLER, K. — cf. BURGHARDT, F. 

GARCIA-BARROS, E. — cf. GARCIA PEREIRA, P. 

GARCIA PEREIRA, P. & GARCIA-BARRoS, E. : Spot distribution 
in Maniola jurtina (Linnaeus, 1758) (Lepidoptera : Nympha- 
lidae, Satyrinae) in Central Spain 

HATTENSCHWILER, P. : Sciopetris karsholti, eine neue Psychide aus 
Tunesien (Lepidoptera, Psychidae) 

HAUSMANN, A.: The morphology of the geometrid moths of the 
Levant and neighbouring countries. Part I: Orthostixinae 
and Geometrinae 

Hausmann, A.: Systematic list of the geometrid moths of the 
Levant and neighbouring countries. Part I: Orthostixinae 
and Geometrinae 

KARRER, F. — cf. LOELIGER, E. A. 

Lockwoop, M. — cf. STEFANESCU, C. 

LOELIGER, E. A. & KARRER, F.: On the induction of metamor- 
phosis of Lepidoptera by means of ecdysone and 20-hydroxy- 
ecdysone 

LoELIGER, E. A. & KARRER, F. : Breeding of an F, of Hyles hybrid 
vespertilioides (Boisduval, 1827) after induction with ecdysone 
of metamorphosis in diapausal pupae 

OLIVIER, A. & De Prins, R. : The butterflies of the Greek island 
of Kés: a synthesis (Lepidoptera : Hesperioidea & Papilio- 
noidea) 


268 


3/4 


3/4 


243 


185 


PELZER, A. : Illustrierter Bestimmungsschlüssel für die Präimagi- 
nalstadien der Schwärmer Europas und Nordafrikas (Lepi- 
doptera : Sphingidae) 

SAMRAOUI, B. : Breeding status and range expansion of Danaus 
chrysippus (Linnaeus, 1758) in the Algerian Sahara (Lepi- 
doptera : Nymphalidae Danainae) 

STEFANESCU, C., Lockwoop, M. & CEBOLLA, M. : The butterfly 
assemblages of northeastern Spanish wetlands 

VANHOLDER, B.: The migration of Danaus plexippus (Linnaeus, 
1758) during October 1995 in the UK (Lepidoptera : Nym- 
phalidae, Danainae) 

Wust, P. : New and rare species of Heterocera (Lepidoptera) and 
some notes on their distribution in different habitats in the 
Nestos Delta, Macedonia, Greece 


Book reviews — Buchbesprechungen — Analyses 


A Catalogue of the type-specimens of Parnassius in the Zoological 
Museum of the Kiev University (Lepidoptera, Papilionidae) 

Die Tagfalter der Türkei unter Berücksichtigung der angrenzenden 
Lander 

Le Sesiidae della fauna italiana (Lepidoptera) 


3/4 


3/4 


3/4 


1/2 


3/4 


3/4 


3/4 
3/4 


230 


261 


165 


129 


142 


264 


265 
267 


269 


New taxa described in Vol. 19 
Neue Taxa in Band 19 beschrieben 
Nouveaux taxa décrits dans le Vol. 19 


GEOMETRIDAE 


Pseudoterpna coronillaria halperini Hausmann, 1996 
Microbaena Hausmann, 1996 

Microbaena pulchra minor Hausmann, 1996 
Proteuchloris Hausmann, 1996 

Thetidia persica Hausmann, 1996 

Victorinella Hausmann, 1996 

Victoria wiltshirei Hausmann, 1996 

Eucrostes indigenata lanjeronica Hausmann, 1996 
Microloxini Hausmann, 1996 

Acidromodes Hausmann, 1996 

Hemidromodes unicolorata Hausmann, 1996 


PSYCHIDAE 


Sciopetris karsholti Hattenschwiler, 1996 


270 


NOTA LEPIDOPTEROLOGICA 


A quarterly journal devoted to Palaearctic lepidopterology 
Published by Societas Europaea Lepidopterologica 


Manuscripts should be sent to the editor : Alain Olivier, Luitenant Lippenslaan 43B14, 
B-2140 Antwerpen, Belgium 


Instructions for authors 


Kopien dieser Hinweise in deutscher Sprache sind beim Redaktor erhältlich. 
Copies de ces instructions en français sont disponibles auprès de l’éditeur. 


Full articles and short communications (max. 2 typed pages) on any aspect of 
Palaearctic lepidopterology will be considered for publication. Full articles will be 
reviewed by two referees. Publication languages are English, French and German. 
Every effort should be made to carry out linguistic corrections before submitting the 
manuscript, otherwise considerable delays can be expected. The editors reserve the 
right to make textual corrections that do not alter the author’s meaning. 


All manuscripts should be clearly typed with double spacing and wide margins, and 
submitted in triplicate. In addition to the original figures, three copies should be 
supplied in a format not exceeding A4. A computer diskette (preferably IBM, but 
Macintosh also accepted) should also be sent where possible. Full articles must include 
a summary of not more than 200 words. A translation of the summary in at least 
one other European language will be added by the editors, if not already supplied 
by the author. Current issues of the journal should be checked for style and format. 
Do not hyphenate words at the right-hand margin or type surnames in capitals. 
References should be styled as follows : 


Heath, J., 1985. New species of Micropterix Hübner (Lepidoptera, Zeugloptera : 
Micropterigidae) from Greece and Cyprus. Nota lepid. 8 : 336-340. 

Heath, J., Pollard, E. & Thomas, J.A., 1984. Atlas of butterflies in Britain and Ireland. 
158 pp. Viking Press, Harmondsworth. 


Legends to figures should be typed on a separate sheet. All figures should be suitably 
mounted and numbered where necessary. When adding numbers and letters, any final 
reduction in size should be allowed for. Drawings should be in black waterproof ink. 
Photographs for black & white reproduction should be glossy positive prints. The 
cost of coloured plates must be borne by the author (currently from about FB11,000 
for a half- or full-page). 


The first mention of any insect should include the full scientific name with the author 
and year of description. Alternatively, the nomenclature used should follow a recent 
list or other suitable work and this must be cited. New descriptions must conform 
with the current edition of the International Code of Zoological Nomenclature. We 
strongly urge deposition of types in major museums and all type depositions must 
be cited. 


Manuscripts not conforming with these instructions may be returned. 


Twenty-five reprints of each article will be supplied free of charge to the first author. 
Additional copies may be ordered on a form enclosed with the proofs. 


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