Animal Biodiversity and Conservation 36.1 (2013)
13
Dalechampii oak
(Quercus dalechampii Ten.),
an important host plant
for folivorous lepidoptera larvae
M. Kulfan, M. Holecová & P. Beracko
Kulfan, M., Holecová, M. & Beracko, P., 2013. Dalechampii oak (Quercus dalechampii Ten.), an important
host plant for folivorous lepidoptera larvae. Animal Biodiversity and Conservation, 36.1: 13–31.
Abstract
Dalechampii oak (Quercus dalechampii Ten.), an important host plant for folivorous lepidoptera larvae.— We
conducted a structured analysis of lepidoptera larvae taxocenoses living in leaf bearing crowns of Dalechampii
oak (Quercus dalechampii Ten.) in nine study plots in the Malé Karpaty Mountains (Central Europe). The
differences between lepidoptera taxocenoses in individual oak stands were analyzed. A total of 96 species
and 2,140 individuals were found. Species abundance peaked in May, while number of species and species
diversity reached the highest values from April to May and from April to June, respectively. Abundance showed
two notable peaks in lush feeders and in late summer feeders. Lepidoptera taxocenosis in the study plot
Horný háj (isolated forest, high density of ants) differed signiicantly from all other taxocenoses according to
Sörensen’s index of species similarity, species diversity, analysis of similarity on the basis of permutation and
pairwise tests (ANOSIM), seasonal variability of species composition, and NMDS ordination.
Key words: Moths, Caterpillars, Q. dalechampii, Malé Karpaty Mountains, SW Slovakia.
Resumen
El roble de dalechampii (Quercus dalechampii Ten.), una importante planta hospedadora de las larvas de
lepidópteros ilófagos.— Llevamos a cabo un análisis estructurado de las taxocenosis de larvas de lepidópteros que viven en las copas del roble de dalechampii (Quercus dalechampii Ten.) en nueve parcelas del
estudio en los Pequeños Cárpatos (Europa central). Se analizaron las diferencias entre las taxocenosis de
lepidópteros de cada roble. Se hallaron 96 especies y 2.140 individuos. La abundancia de especies alcanzó
su valor más elevado en mayo, mientras que el número y la diversidad de especies fueron máximos desde
abril hasta mayo y desde abril hasta junio, respectivamente. La abundancia mostró dos máximos notables en
las larvas que se alimentan durante la brotación y las que se alimentan al inal del verano. La taxocenosis
de los lepidópteros en la parcela del estudio Horný háj (un bosque aislado con una elevada densidad de
hormigas) diirió signiicativamente de las demás taxocenosis según el índice de Sörensen para la similitud
de las especies, la diversidad de las especies, el análisis de la similitud sobre la base de las pruebas de
permutación y las pruebas de pares (ANOSIM), la variabilidad estacional de la composición de especies y
el escalamiento multidimensional no métrico (NMDS por sus siglas en inglés).
Palabras clave: Polillas, Orugas, Q. dalechampii, Pequeños Cárpatos, Eslovaquia sudoccidental.
Received: 11 VII 12; Conditional acceptance: 27 X 12; Final acceptance: 20 XII 12
Miroslav Kulfan & Pavel Beracko, Dept. of Ecology, Fac. of Natural Sciences, Comenius Univ., Mlynská dolina
B–1, SK–84215 Bratislava, Slovakia.– Milada Holecová, Dept. of Zoology, Fac. of Natural Sciences, Comenius
Univ., Mlynská dolina B–1, SK–84215 Bratislava, Slovakia.
Corresponding author: M. Kulfan. E–mail: kulfan@fns.uniba.sk
ISSN: 1578–665X
© 2013 Museu de Ciències Naturals de Barcelona
Kulfan et al.
14
Introduction
Oaks belong to the woody plants that host the richest
insect assemblages in Central Europe (Patočka et al.,
1999). Lepidoptera larvae have been shown to be the
most important group of oak defoliators (Patočka et
al., 1962, 1999). About 250 lepidoptera species are
known to damage the assimilation tissue of oaks in
Central Europe (Patočka et al., 1999; Reiprich, 2001).
Lepidoptera fauna on some oak species in Central
Europe have been relatively well studied (Patočka et
al., 1962, 1999; Csóka, 1990–1991, 1998a, 1998b;
Kulfan, 1990, 1997; Kulfan, 1992; Kulfan et al., 1997,
2006; Kulfan & Degma, 1999; Turčáni et al., 2009, 2010;
Parák et al., 2012, etc.). Taxocenoses of lepidoptera
caterpillars on three oak species from Slovakia and
the Czech Republic (Quercus robur, Q. petraea and
Q. cerris) have been used to explain why there are so
many species of herbivorous insects in tropical rainforests (Novotny et al., 2006).
However, the lepidoptera fauna related to Q.
dalechampii growths has been poorly explored in
Europe. A total of nine lepidoptera miner species from
families Nepticulidae, Tischeriidae and Gracillariidae
have been recorded on Q. dalechampii in southern
Slovakia (Arboretum Čifáre) (Skuhravý et al., 1998).
Kollár (2007) mentions the species Phyllonorycter
roboris (lepidoptera miner) as a pest of Q. dalechampii
in Slovakia. Stolnicu (2007) studied lepidoptera leaf–
miners on Q. dalechampii in Romania. Kulfan (2012)
partially studied economically most important pest
species on Q. dalechampii in Central Europe.
Dalechampii oak (Quercus dalechampii Ten.) is one of
the most common oaks in Europe and is naturally distributed in Western Italy, Sicily, Greece, Albania, Montenegro,
Macedonia, Bosnia & Herzegovenia, Serbia, Slovenia,
Austria, Hungary, Slovakia, Romania, and Bulgaria.
The main aims of the present study were: (i) to
analyze the structure taxocenoses, alpha diversity and
representation of trophic groups and seasonal guilds of
lepidoptera en bloc on Dalechampii oak; (ii) to complete
data concerning biodiversity of lepidoptera species
feeding on oaks in Central Europe; and (iii) to highlight
the differences among the individual study plots representing various types of oak forests, with emphasis on
fragmentation, forest age and crown canopy.
Material and methods
Material was collected by the beating method into
a tray of 1 m diameter (one quantitative sample =
beating from 25 branches) on nine selected plots
at regular 2–weekly intervals from April to October
2000–2002. Samples were taken from branches at a
height of about 1–2.5 m above ground with varying
exposure to cardinal points. Larvae were identiied
using the keys by Gerasimov (1952), Patočka (1954,
1980) and Patočka et al. (1999). Seasonal guilds of
lepidoptera caterpillars were established according to
Turčáni et al. (2009).
The complete linkage clustering in combination
with Sörensen’s index and Wishart’s similarity ratio
was used to classify the taxocenoses. Visualization of
dendrograms was done by computer program Syn–tax,
Version 5.0 (Podani, 1993). Diversity of taxocenoses
was characterised using Pielou’s index of equitability,
Shannon–Wiener’s index of total species diversity, and
Simpson’s index of dominance (Poole, 1974; Ludwig
& Reynolds, 1988). Shannon–Wiener diversity indices
were compared using the t–test (Poole, 1974). Ordination was carried out with non–metric multidimensional
scaling (NMDS) using the Bray–Curtis dissimilarity
coeficient. One–way analysis of similarities (ANOSIM)
was used to identify difference in species variability
of the lepidoptera taxocenosis in the study plots during the year. Hierarchical (nested) ANOVA was used
to examine spatial (locality) and temporal (sampling
months) variation in the distribution of the total abundance, number of species, taxa and species diversity
of lepidoptera. The model contained factors (terms)
representing the effects of locality and sampling date
nested in locality. Multiple sample comparisons were
used to identify signiicant differences in the number
of individuals, number of species and species diversity between localities and sampling months. The
hypothesis that occurrences of three types of feeding
specialization are randomly distributed throughout the
vegetation season was tested according to Poole &
Rathcke (1979). Differences of means and dispersion
of species numbers in feedings groups were analyzed
by Tukey’s pairwise comparison and Levene’s test in
ANOVA, respectively. Analyses of variance and Tukey’s
pairwise comparison were used to identify differences
between the number of species and the number of
individuals in seasonal gilds. The nomenclature and
systematic classiication of the lepidoptera species
were used according to Laštůvka & Liška (2011). The
trophic groups of lepidoptera larvae were established
according to Brown & Hyman (1986). The map (ig. 1)
and pedological and phytocoenological characteristics
of the investigated area are given in detail by Zlinská
et al. (2005).
Voucher specimens (in ethanol) are deposited at
the Faculty of Natural Sciences, Comenius University,
Bratislava, Slovakia.
Study area
The lepidoptera larval stages on Quercus dalechampii were studied in the territories of the Protected
Landscape Area of Malé Karpaty and Trnavská
pahorkatina hills situated in the centre of Europe
in the western part of Slovakia.The vast majority of
the plots are located in the southern to northern part
of the Malé Karpaty Mountains (Mts.) at altitudes of
240–350 m a.s.l. and an average annual temperature
of 8–9°C. Study plots in Trnavská pahorkatina hilly
land are situated near the Malé Karpaty Mts. at an
altitude of 240 m. The annual precipitation in both
territories is about 650–800 mm.
Study plots (abbreviation of study plot in parentheses
used in the text):
Vinosady (VI), 48º 19' N, 17° 17' E, 280 m a.s.l.:
a 60–80–year–old forest at the foot of the Kamenica
Animal Biodiversity and Conservation 36.1 (2013)
15
70
Study plot
72
N
Contour line (m a.s.l.)
72
48º 48' N
W
Malé Karpaty Mts. border
500
200
71
E
300
67
73
68
69
500
GRN DFS
Bradlom
Jablonica
300
68
Čachtice
48º 42' N
200
S
Brezová pod
Settlement
Vrbové
200
0
30
300
48º 36' N
Chtelnica
200
74
300
Rohožnik
75
0
500
LL
Smolenice
NK
HH
50
48º 30' N
NA
LH
500
200
20
FU
48º 24' N
0
76
LI
Lozorno
CA
VI
0
5
10
15
20
25 km
30
0
Stupava
Modra
77
300
Sv. Jur
48º 12' N
78
200
48º 06' N
Bratislava
79
17º 00'E
Pezinok
17º 10' E
48º 00' N
17º 20' E
48º 18' N
17º 30' E
17º 40' E
17º 50' E
17º
18º
19º
20º
21º
22º
63 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 94 96 97 98 99 00 01
63
64
65
66
67
68
69
49º70
71
72
73
74
75
76
77
78
79
48º80
81
82
83
GRN DFS: Grid references number of the Databank of Slovak fauna
Map design:
© Marek Šimurka 2005
Fig. 1. Study area with location of the study plots.
Fig. 1. Área del estudio con la ubicación de las parcelas del estudio.
hill, NW and W oriented, with drier subxerophilous
meadows and shrub complexes. Besides Quercus
dalechampii, the tree stratum consists of Q. cerris
and Acer campestre.
Cajla (CA), 48º 20' N, 17° 16' E, 260–280 m a.s.l.:
an 80–100–year–old forest at the foot of the Malá
cajlanská homola hill, S oriented and neighbouring
meadows and vineyards on S and E, from N and W
closed forest complexes. Quercus dalechampii and
Carpinus betulus predominate in the tree layer.
Fúgelka (FU), 48º 22' N, 17° 19' E, 350 m a.s.l.:
an 80–100–year–old forest near the Dubová village,
S oriented. Besides Quercus dalechampii, the tree
stratum consists of Acer pseudoplatanus.
Lindava (LI) (Nature Reserve), 48º 22' N, 17° 22' E,
240 m a.s.l.: an 80–100 (120)–year–old forest near
the village of Píla. Quercus dalechampii and Q. cerris
predominate in the tree layer.
Horný háj (HH), 48º 29' N, 17° 27' E, 240 m a.s.l.:
a larger complex of an island forest 60–80–years old
Kulfan et al.
16
near the village of Horné Orešany, surrounded by
ields and vineyards, W and SW oriented. Quercus
cerris, Q. dalechampii, Carpinus betulus and Fraxinus
excelsior predominate in the tree layer.
Lošonec–lom quarry (LL), 48º 29' N, 17° 23' E,
340 m a. s. l.: an 80–100–year–old forest SW oriented,
neighbouring with mesophilous meadows and pastures.
The tree layer consists of Quercus dalechampii, Q.
cerris and Carpinus betulus. The leaf litter, herbage
undergrowth and trees are strongly covered with calcareous dust from a nearby quarry.
Lošonský háj (LH) (Nature Reserve), 48º 28' N,
17° 24' E, 260 m a.s.l.: an 80–100–year–old oak–hornbeam forest NE oriented, surrounded by closed forest
complexes. Quercus dalechampii, Q. cerris and Carpinus
betulus predominate in the tree stratum.
Naháč–Kukovačník (NA), 48º 32' N, 17° 31' E,
300 m a.s.l.: a small forest island, approximately 40–60–
year–old surrounded by ields and pastures, NE oriented.
Quercus dalechampii, Q. cerris and Carpinus betulus
predominate in the tree layer.
Naháč–Katarínka (NK) (Nature Reserve), 48° 33' N,
17° 33' E, 340 m a.s.l.: a 40–60–year–old forest NW
oriented, surrounded by closed forest ecosystems.
Quercus dalechampii and Carpinus betulus predominate
in the canopy.
Only abbreviations of the study plots are used in
the following text.
The study plots LI and HH are situated in Trnavská pahorkatina hills and the others are in the Malé
Karpaty Mts.
Results
From 2000–2002, a total of 2,140 Lepidoptera larvae
were collected in nine study plots with Quercus dalechampii. They represented 96 species from 17 families
(appendix 1). The families Geometridae, Noctuidae and
Tortricidae encompassed the highest number of species
found (27, 23, and 13, respectively) (appendix 1). The
lowest number of species (18 species) were found in
HH (appendix 1). Six species (Coleophora siccifolia,
Lomographa temerata, Peribatodes rhomboidaria,
Acronicta auricoma, Orthosia opima and Amata phegea)
were found on oaks for the irst time in Slovakia (cf.
Hrubý, 1964; Patočka et al., 1999). A. phegea is one
of six species presenting irst records of lepidoptera
larvae feeding on oaks. This species probably entered
the oak crown from the surrounding low vegetation
because it has not been found previously on trees
according to the literature (Reiprich, 2001).
The most abundant families were Geometridae and
Noctuidae (appendix 1, table 1). The families Tortricidae and Erebidae achieved relatively high dominance,
mainly due to the species Aleimma loelingiana (Tortricidae) and Lymantria dispar (Erebidae) (appendix 1,
table 1). Species with dominance higher than 10%
were Lymantria dispar in HH, Operophtera brumata
in CA (calamitous oak pests), Cosmia trapezina in
LI, Aleimma loelingiana in FU (an important pest of
oaks) and Cyclophora linearia in HH (cf. Patočka et
al., 1999; appendix 1).
Table 1. Family dominance (%) of lepidoptera
larvae on Quercus dalechampii in the Malé
Karpaty Mountains in 2000–2002 (based on
total number of individuals).
Tabla 1. Dominancia por familia (%) de las larvas
de lepidópteros que se encontraron en Quercus
dalechampii en los Pequeños Cárpatos entre
los años 2000 y 2002 (con respecto al número
total de individuos).
Family / year
2000
2001
2002
Total
Psychidae
0.00
0.17
0.00
0.05
Bucculatricidae
0.00
0.00
0.43
0.19
Gracillariidae
0.17
0.00
0.00
0.05
Ypsolophidae
2.48
2.15
1.29
1.87
Chimabachidae
2.74
1.49
1.12
1.73
Peleopodidae
8.85
1.65
1.25
3.46
Coleophoridae
8.77
3.63
4.09
5.28
Gelechiidae
0.17
0.83
0.22
0.37
Tortricidae
6.79
19.64
Lycaenidae
0.33
0.33
0.43
0.37
Pyralidae
1.82
0.50
1.29
1.21
Drepanidae
0.33
0.33
0.11
0.23
9.68 11.68
Geometridae
32.62
Notodontidae
5.46
0.17
0.22
1.68
Erebidae
7.95
5.94
9.04
7.85
Nolidae
3.48
1.98
1.72
2.29
Noctuidae
No individuals
18.05
604
39.11 42.58 38.79
22.11 26.56 22.90
606
930 2,140
The species Lymantria dispar, Cyclophora linearia,
Pseudoips prasinana and Carcina quercana reached
the highest dominance on the species poorest study
plot HH when compared with other plots (appendix 1).
Characteristic species of the plot LL covered
with calcareous dust are as follows: Tortrix viridana,
Conobathra tumidana, Aleimma loelingiana, Agriopis
leucophaearia and Alsophila aceraria. Three lepidoptera species, Archips podana, Eudemis profundana
and Apocheima hispidaria (appendix 1), were found
only in this plot but abundance was low.
Lepidoptera species Agriopis marginaria, Cosmia
trapezina, Orthosia cruda and Lymantria dispar (apendix 1) were typical of the lighter, sparser and younger
oak stands (study plots NK, LI, CA, VI).
The vast majority of Lepidoptera belonged to the
monovoltine species with main occurrence in spring.
Further oligophagous species (Cyclophora linearia
and Ennomos erosaria) and polyphagous species
(Parectropis similaria and Colocasia coryli) belonged
Animal Biodiversity and Conservation 36.1 (2013)
17
to the bivoltine species. Watsonalla binaria proved to
be trivoltine species (appendix 1).
Most species found belonged to the trophic group of
generalists (64 species). Narrow oligophages (18 species) feeding on oaks are considered to be typical oak
species. Only six species belonged to wider oligophages.
The value of Shannon–Wiener´s diversity index of
the richest lepidoptera taxocenosis (NK, H’ = 3.428)
and the poorest taxocenosis (HH, H’ = 2.505) was
statistically signiicantly different from other taxocenoses (T–test, P < 0.05) (table 2). A detailed algorithm
is given by Poole (1974). The richest taxocenosis
NK includes 462 individuals representing 52 species;
of these, seven species dominate at least 5%. The
poorest taxocenosis HH includes only 44 individuals
belonging to 18 species; 4 of these species dominate
over 5% (appendix 1).
Poor qualitative–quantitative taxocenosis of lepidoptera larvae on island forest HH is also expressed
by Simpson’s index of dominance (c = 0.126) where
dominance is concentrated in a small number of species (appendix 1). In other taxocenoses, dominance is
spread to more co–dominant species (Simpson’s index
of dominance values from 0.044 to 0.086). The value
of equitability was highest at FU, NK and NA (table 2).
A dendrogram based on the qualitative representation (Sörensen’s index, complete linkage) separated the
lepidoptera taxocenosis on the study plot HH (isolated
forest, high density of ants, the lowest diversity of
species) (ig. 2). Based on a qualitative–quantitative
similarity (Wishart’s similarity ratio, complete linkage),
the hierarchical classiication divided the lepidoptera
taxocenoses into two clusters connected on the relatively low level of similarity (ig. 3). The irst cluster consisted of the taxocenoses HH and NA (island forests)
with the lowest igures for abundance and individuals
(44 and 133, respectively). The second cluster had two
subclusters and included other taxocenoses. The irst
subcluster contained the taxocenoses from the denser
and older plots (LL. Study plot affected by calcium dust
deposition and with higher canopy cover of shrub story;
LH. Lot with higher canopy cover of wood species
crowns; and FU. Plot with higher canopy cover of both
shrub story and wood species crowns). The second
subcluster may be formed from the taxocenoses on
lighter and younger plots (NK, LI, CA and VI)
The NMDS showed plot HH was set apart from
all the other study plots (ig. 4). The study plot NA
was also separated (although less marked so) as
conirmed by Wishart’s index.
Table 2. Species diversity test and basic characteristics of caterpillar taxocenoses at study plots in
2000–2002: H'. Shannon’s index of species diversity; e. Pielou’s index of equitability; c. Simpson’s index of
dominance. (T–test values of H' are under the diagonal and degrees of freedom are above it; the testing
process is detailed in Materials and methods; signiicance levels: *** P < 0.001; ** 0.001 < P < 0.01;
* = 0.01 < P < 0.05; ns = 0.05 < P (non–signiicant); for abbreviations of the study plots see Material
and methods).
Tabla 2. Prueba de la diversidad de especies y características básicas de las taxocenosis de orugas en las
parcelas del estudio entre los años 2000 y 2002. H'. Índice de Shannon para la diversidad de especies;
e. Índice de Pielou para la equidad; c. Índice de Simpson para la dominancia. (Los valores de H' de la
prueba t se encuentran debajo de la diagonal y los grados de libertad, encima; el proceso de la prueba se
detalla en el apartado Material and methods; niveles de signiicación: *** P < 0,001; ** 0,001 < P < 0,01;
* = 0,01 < P < 0,05; ns = 0,05 < P (no signiicativo); para consultar las abreviaturas de las parcelas del
estudio, ver Material and methods).
e
VI
CA
FU
LI
HH
LL
LH
NA
NK
0.851
0.801
0.872
0.809
0.867
0.838
0.849
0.867
0.868
c
0.066
0.086
0.063
0.063
0.126
0.067
0.063
0.066
0.044
H'
3.097
3.065
3.101
3.212
2.505
3.091
3.194
3.197
3.428
VI
3.1
0
541.083
349.481
636.538
55.534
445.853
356.73
228.02
670.81
CA
3.07
0.343ns
0
421.247
582.174
65.467
495.143
431.18
294.16
477.82
FU
3.1
0.048ns
0.346ns
0
396.331
64.992
372.919
350.76
265.84
289.7
LI
3.21
1.356ns
1.495ns
1.127ns
0
59.702
491.438
402.8
259.76
603.69
HH
2.51
3.549***
3.218**
3.429**
4.162***
0
63.662
68.671
78.673
51.015
LL
3.09
0.071ns
0.246ns
0.106ns
1.263ns
3.387**
0
386.06
272.78
381.14
LH
3.19
0.99ns
1.174ns
0.842ns
0.182ns
3.901***
0.957ns
0
284.73
299.12
NA
3.2
0.908ns
1.092ns
0.792ns
0.131ns
3.764***
0.894ns
0.029ns
0
192.36
NK
3.43
4.681***
4.198***
3.784***
2.773**
5.653**
4.013***
2.567**
2.201*
0
Kulfan et al.
18
0.8
Dissimilarity scale
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0.0
HH
FU
NA
LL
CA
LH
NK
LI
VI
Fig. 2. Classiication of lepidoptera taxocenoses on individual study plots according to species presence/
absence (Sörensen’s index).
Fig. 2. Clasiicación de las taxocenosis de lepidópteros en cada una de las parcelas del estudio en
función de la presencia o ausencia de las especies (índice de Sörensen).
Table 3 shows the overall result of the permutation test and pairwise ANOSIMs between all pairs
of groups (provided as post–hoc test). Signiicant
comparisons (at P < 0.05) are shown in bold.
Analysis of similarity based on seasonal variability
of species composition distinguished two signiicant
different lepidoptera taxocenoses. The lepidoptera
taxocenosis of the HH had signiicantly lower abundance and number of species than the taxocenoses
of the other eight study plots (table 4).
Generally, lepidoptera larvae were weakly represented in HH because of the occurrence of numerous
1.0
Dissimilarity scale
0.9
0.8
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0.0
NA
HH
LH
LL
FU
NK
LI
CA
VI
Fig. 3. Classiication of lepidoptera taxocenoses on individual plots according to qualitative–quantitative
similarity (Wishart’s index).
Fig. 3. Clasiicación de las taxocenosis de lepidópteros en cada una de las parcelas del estudio en
función de la similitud cualitativa y cuantitativa (índice de Wishart).
Animal Biodiversity and Conservation 36.1 (2013)
0.16
19
Stress: 0.073
CA
0.12
NMDS_2
0.08
0.04
NK
HH
LI
0
FU
–0.04
LH
–0.08
–0.12
LL
NA
VI
–0.16
–0.2
–0.72 –0.6 –0.48 –0.36 –0.24 –0.12 0
0.12 0.24
NMDS_1
Fig. 4. Nonmetric multidimensional (NMDS) scaling plot based on Bray–Curtis similarities for species
abundance data from nine study plots. (For abbreviations of the study plots see Material and methods.)
Fig. 4. Gráico del escalamiento multidimensional no métrico (NMDS) de las similitudes de Bray–Curtis
a partir de los datos sobre la abundancia de las especies obtenidos en nueve parcelas del estudio (para
consultar las abreviaciones de las parcelas del estudio, véase el apartado Material and methods).
colonies of ants as predators of lepidoptera larvae in
this plot (appendix 1).
The seasonal effect was relected in all three examined parameters of the taxocenosis (table 4). The species
abundance peaked in May, while number of species
and species diversity reached the highest values from
April to May and from April to June, respectively (ig. 5).
Figure 6 shows the number of species and the
number of individuals in seasonal guilds. The number of species and the abundance showed two clear
peaks in lush feeders and in late summer feeders.
The number of lush feeder species was signiicantly
higher than the number of species in other seasonal
guilds (table 5).
Table 3. Results of analysis of similarity (ANOSIM): permutation number: 1,000; mean rank within:
5,047; mean rank between: 5,241; R = 0.037, P < 0.01. (For abbreviations of the study plots see
Material and methods.)
Tabla 3. Resultados del análisis de similitud (ANOSIM): número de permutaciones: 1.000; rango medio
dentro: 5.047; rango medio entre: 5.241; R = 0.037; P < 0,01. (Para las abreviaturas de las parcelas del
estudio, véase el apartado Material and methods).
VI
CA
LI
LL
NA
NK
HH
FU
LH
VI
CA
LI
LL
NA
NK
HH
FU
LH
–
0.16
0.39
0.73
0.15
0.23
0.02
0.06
0.18
–
0.58
0.67
0.77
0.46
0.01
0.68
0.21
–
0.68
0.15
0.32
0.01
0.73
0.17
–
0.38
0.62
0.02
0.83
0.33
–
0.48
0.01
0.18
0.22
–
0
0.3
0.3
–
0.09
0.11
–
0.3
–
Kulfan et al.
20
Table 4. Comparison of abundance, species number and species diversity in spatial and temporal scaling
of studied lepidoptera taxocenoses in the hierarchical (nested) analysis of variance (ANOVA): * P < 0.05,
** P < 0.01; SSq. Sum of squares; MSq. Mean squares; SS. Sampling site; SD. Sampling date.
Tabla 4. Comparación de la abundancia, el número de especies y la diversidad de especies en las
escalas espacial y temporal de las taxocenosis estudiadas de lepidópteros en el análisis jerárquico
(anidado) de la varianza (ANOVA): * P < 0.05, ** P < 0.01; SSq. Suma de los cuadrados; MSq. Media
de los cuadrados; SS. Lugar de muestreo; SD. Fecha de muestreo.
SSq
df
MSq
SS
13336.73
8
1667.09
4.29614
P < 0.01
SD
61095.30
54
1131.39
3.51589
P < 0.01
F–statistic
P–value
Mann–Whitney
pairwise comparison
Abundance
HH < CA, VI, FU, LI, NK, NA, Ll*
June–October < April–May*
Number of species taxa
SS
268.798
8
33.600
3.0216
P < 0.01
HH < CA, VI, FU, LI, NK, NA, Ll*
SD
4419.608
54
81.845
7.3602
P < 0.01
June–October < April–May*
Species diversity
SS
6.50334
8
0.81292
1.2961 P = 0.2571
SD
86.24383
54
1.59711
4.9800
The occurrence in time of lepidoptera species
with two types of feeding specialization (generalists,
narrow oligophagous) was non–randomly distributed
throughout the season (table 6). The number of species in these feeding groups peaked in May. On the
other hand, species in the wider oligophagous feeding
groups exploited time in a random way.
Discussion
Taxocenoses of lepidoptera larvae observed on
Quercus dalechampii in Malé Karpaty Mts. can be
compared with taxocenoses on Q. cerris that were
studied under similar conditions. A comparison shows
similarities and differences (cf. Kulfan et al., 2006).
Species richness was higher on Q. dalechampii
(96 species on Q. dalechampii compared to 58 species on Q. cerris). The lowest number of species in
both types of taxocenoses was found in the study
plot HH. Lymantria dispar and Operophtera brumata
belonged to the most abundant species both on Q.
dalechampii and on Q. cerris. In the study plot HH,
L. dispar reached a higher dominance on Q. cerris
than on Q. dalechampii (cf. Kulfan et al., 2006).
Regarding cumulative dominance, the families Ypsolophidae, Pyralidae and Drepanidae predominated on
Q. cerris. On the contrary, the families Peleopodidae
and Chimabachidae were noticeably more common
on Q. dalechampii (cf. table 1, Kulfan et al., 2006).
In general, when compared with other areas of
Slovakia, the abundance of lepidoptera larvae found
on Q. dalechampii corresponds to the latent phase of
P < 0.01
––
August–October < April–June**
the gradation cycle on oaks. No marked outbreaks of
folivorous lepidoptera larvae have been observed on
oaks in Slovakia since 1990 (cf. Kulfan, 1990, 1998,
2002; Kulfan, 1992; Kulfan et al., 1997, 2006).
The values of species diversity of lepidoptera
taxocenoses on Q. dalechampii are characterized
by a greater variance than the values of diversity of
taxocenoses on Q. petraea in the Malé Karpaty Mts.
Diversity of lepidoptera taxocenoses on Q. petraea
reached annual values ranging from 3.042 to 3.296
(Kulfan, 1990). The smallest diversity of lepidoptera
taxocenoses on oaks in the Malé Karpaty Mts.was
found on Q. cerris and it achieved a value of 2.230
(Kulfan et al., 2006) for the three–year period.
As a rule, there is notable spring peak in abundance of lepidoptera caterpillars on oaks in Central
Europe (Kulfan, 1992; Kulfan, 1983, 1990; Parák
et al., 2012, etc.). This was also conirmed by the
research on Q. dalechampii. Other peaks in caterpillar abundance on Q. dalechampii were not found
throughout the growing season. Two peaks in the
number of lepidoptera caterpillars during the season
with prevalence in spring time were found on some
oak species in Central Europe (Kulfan, 1992; Kulfan,
1983, 1990). The abundance of lepidoptera taxocenoses on Q. petraea throughout the growing season
in the Malé Karpaty Mts in Slovakia has been found
to have a very marked peak in spring (May–June)
and a less noticeable peak in autumn (September).
However, it is interesting that the autumn peak of total
abundance of lepidoptera caterpillars on Q. petraea
in the old oak stand in 1978 was more noticeable
when compared to the spring peak (cf. Kulfan, 1983).
Animal Biodiversity and Conservation 36.1 (2013)
21
1
Diversity
0.8
0.6
0.4
0.2
0
IV
V
VI
VII
VIII
Ix
x
Number of individuals
200
160
120
80
40
0
Number of species
IV
V
VI
VII
VIII
Ix
x
40
39
20
10
0
IV
V
VI
VII
VIII
Ix
x
Fig. 5. Box plots showing the monthly variation of diversity, abundance and number of species of lepidoptera
taxocenosis: IV–X. Months of presence of lepidopteran caterpillars during the season.
Fig. 5. Diagramas de caja en los que se muestra la variación mensual de la diversidad, la abundancia y
el número de especies de la taxocenosis de los lepidópteros: IV–X. Meses de presencia de las orugas
lepidópteras durante la estación.
This was probably caused by unfavorable weather
conditions in spring.
Yoshida (1985) in northern Japan presented the
highest abundance of lepidoptera caterpillars on oaks
in summer. This difference compared to our results
may be caused by different climate, because the
frosts in May in northern Japan have a large impact
on leaf phenology, which is associated with the development of the spring taxocenoses of caterpillars.
In oak forest on Mont Holomontas (Mediterranean
area, Greece) even three peaks in insect abundance
(consisting mainly of lepidoptera larvae) on six oak
species were found (Kalapanida & Petrakis, 2012).
Not only the species abundance but also the species richness and diversity of lepidoptera species on
Q. dalechampii culminated in the vernal aspect. The
marked increase of species diversity of lepidoptera
taxocenoses on Q. dalechampii was in spring. A similar
Kulfan et al.
22
Number of species
50
40
30
20
10
Number of individulas
0
FIF
SF
LSF
AF
FIF
SF
LSF
AF
500
400
300
200
100
0
Fig. 6. Box plots showing the effects of seasonality on species number and abundance of lepidoptera
taxocenosis: FIF. Flush feeders; SF. Summer feeders; LSF. Late summer feeders; AF. Autumn feeders.
Fig. 6. Diagramas de caja en los que se muestra los efectos de la estacionalidad en el número de especies y en la abundancia de la taxocenosis de los lepidóptero: FIF. Se alimentan durante la brotación;
S. Se alimentan en verano; LSF. Se alimentan al inal del verano; AF. Se alimentan en otoño.
Table 5. Results of one–way ANOVA on differences between seasonal guilds in the number of species taxa
and number of individuals. The post hoc multiple sample comparison test (Tukey's pairwise comparison) for
differences in mean number of species taxa and number of individuals between seasonal guilds: * P < 0.05,
** P < 0.01; FIF. Flush feeders; LSF. Late spring feeders; SF. Summer feeders; AF. Autumn feeders.
Tabla 5. Resultados de la ANOVA simple de las diferencias existentes entre los gremios estacionales en
cuanto el número de taxones y el número de individuos. La prueba múltiple de comparación a posteriori de
Tukey (comparación por pares de Tukey) de las diferencias en el número medio de taxones y el número
de individuos entre gremios estacionales: * P < 0,05; ** P < 0,01; FIF. Se alimentan durante la brotación;
LSF. Se alimentan al inal de la primavera; SF. Se alimentan en verano; AF. Se alimentan en otoño.
F–statistic
df
P–value
Tukey's pairwise
comparison
Number of species in seasonal guilds
85.49
35
P < 0.01
SF, LSF, AF < FIF**
Number of individuals in seasonal guilds
25.44
35
P < 0.01
SF, LSF, AF < FIF**
SF, AF < LSF**
Animal Biodiversity and Conservation 36.1 (2013)
23
Table 6. Results obtained from the Poole–Rathcke method used to segregate the moths in time. The
null hypothesis (H1) states that the dispersion is not signiicantly different from random and the second
null hypothesis (H2) that the two means and dispersions are not signiicantly different: N. Number of
species; OV. Observed variance; EV. Expected variance; DR. Dispersion ratio; RD. Random dispersion
(signiicance of H1); HM. Homogeneity of means; HD. Homogeneity of dispersion (signiicance of H2).
(* P < 0.05, ** P < 0.01, ns. Non–signiicant)
Tabla 6. Resultados obtenidos con el método de Poole–Rathcke empleado para segregar las polillas en
el tiempo. La hipótesis nula (H1) airma que la dispersión no es signiicativamente distinta de la aleatoria
y la segunda hipótesis nula (2), que las dos medias y las dos dispersiones no son signiicativamente
diferentes: N. Número de especies; OV. Varianza observada; EV. Varianza esperada; DR. Razón de
la dispersión; RD. Dispersión aleatoria (signiicación de H1); HM. Homogeneidad de las medias; HD.
Homogeneidad de la dispersión (signiicación de H2). (* P < 0,05; ** P < 0,01; ns. No signiicativa).
N
OV
EV
DR
RD
Generalists
64
9.87
0.19
51.94736842
**
Narrow oligophagous
18
18.7
0.27
69.25925926
**
Wider oligophagous
6
15.33
0.82
18.69512195
ns
Feeding specialization
HM
HD
Feeding specialization compared
Generalist / narrow oligophagous
Q = 8.54 P < 0.01
W = 2.14 P = 0.15
Generalist / wider oligophagous
Q = 8.74 P < 0.01
W = 13.55 P < 0.01
Narrow oligophagous / wider oligophagous
Q = 0.19 P = 0.98
W = 14.92 P < 0.01
course of diversity was observed on four oak species
in the Borská nížina Lowland in Slovakia. (Kulfan &
Degma, 1999).
Southwood et al. (2005) found distinct seasonal
patterns in species richness of the arthropod fauna
on four oak species in the U.K. In terms of species
richness, the values showed a general trend peaking
in summer and early autumn, but biomass peaked
in May on the native oak species, mainly due to
lepidoptera larvae.
A relatively steady decrease in the individuals from
early spring to autumn is well known from the 'Quercus type' of host tree (Niemelä & Haukioja, 1982).
These authors suggested that this effect was due to
a decline in available resources. Feeny (1970) and
Kamata & Igarashi (1996) stated that tougher leaves
with a higher tannin concentration contributed to the
lower richness of Lepidoptera later during the growing
season. A negative correlation between some specialist oak feeders and condensed tannins in the canopy
of Quercus alba and understorey of Q. velutina was
found (Forkner et al., 2004). Their results generally
indicated a negative response from both specialists
and generalists to condensed tannins.
A higher number of lush feeders in spring compared to the number of species in other seasonal
guilds on Q. dalechampii was also found on three oak
species in Borská nížina lowland (western Slovakia,
Central Europe), where the greatest proportion of
lush feeders was found on Q. robur (cf. Turčáni et
al., 2009).
Acknowledgements
This work was supported by the Slovak Grant Agency for Science (Grant Nos 1/0124/09, 1/0137/11,
2/0035/13 and 1/0066/13). We are grateful to Jaroslav
Fajčík for technical assistance.
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Appendix 1. The list of the lepidoptera species recorded in the nine study plots in the Malé Karpaty
Mountains on Quercus dalechampii with dominance (%), months of occurrence of larvae (MO), trophic
group (TG: S2. Narrow oligophagous; S3. Wider oligophagous species; G. Generalists; U. Unknown),
and larval trophic specialization and seasonal guilds (SG: FIF. Flush feeders; LSF. Late spring feeders;
SF. Summer feeders; AF. Autumn feeders). (For abbreviations of study plots see Material and methods.)
Families and species
VI
CA
FU
LI
0.0
0.0
0.0
0.3
0.0
0.0
0.0
0.0
0.0
0.0
0.6
0.0
Psychidae
Sterrhopterix fusca (Haworth, 1809)
Bucculatricidae
Bucculatrix ulmella Zeller, 1848
Gracillariidae
Phyllonorycter sp.
Ypsolophidae
Ypsolopha alpella (Denis et Schiffermüller, 1775)
1.8
0.0
0.0
0.9
Ypsolopha parenthesella (Linnaeus, 1761)
0.0
0.0
0.0
0.0
Ypsolopha ustella (Clerck, 1759)
0.3
1.1
0.0
0.3
Chimabachidae
Diurnea fagella (Denis et Schiffermüller, 1775)
0.0
0.7
0.6
0.0
Diurnea lipsiella (Denis et Schiffermüller, 1775)
0.6
2.8
1.2
0.6
0.6
2.8
3.0
4.6
Coleophora ibipennella Zeller, 1849
0.0
0.0
0.0
0.3
Coleophora kuehnella (Goeze, 1783)
0.0
0.0
0.0
0.0
Coleophora lutipennella (Zeller, 1838)
0.6
4.2
6.6
4.6
Coleophora siccifolia Stainton, 1856
0.0
0.0
0.0
0.3
Anacampsis timidella (Wocke, 1887)
0.0
0.0
0.6
0.0
Carpatolechia decorella ( Haworth, 1812)
0.0
0.4
0.0
0.0
Peleopodidae
Carcina quercana (Fabricius, 1775)
Coleophoridae
Gelechiidae
Psoricoptera gibbosella (Zeller, 1839)
0.0
0.0
0.0
0.3
Stenolechia gemmella (Linnaeus, 1758)
0.0
0.4
0.0
0.0
Aleimma loelingiana (Linnaeus, 1758)
7.7
0.0
12.0
3.4
Archips crataegana (Hübner, 1799)
0.6
0.0
0.0
0.3
Archips podana (Scopoli, 1763)
0.0
0.0
0.0
0.0
Tortricidae
Eudemis profundana (Denis et Schiffermüller, 1775)
0.0
0.0
0.0
0.0
Pammene albuginana (Guenée, 1845)
0.0
0.0
0.0
0.0
Pandemis cerasana (Hübner, 1786)
0.0
0.7
2.4
0.0
Pandemis corylana (Fabricius, 1794)
0.0
0.4
0.0
0.3
Pandemis heparana (Denis et Schiffermüller, 1775)
0.0
0.4
0.0
0.3
Ptycholoma lecheana (Linnaeus, 1758)
0.0
0.0
0.0
0.3
Spilonota ocellana (Denis et Schiffermüller, 1775)
0.0
2.1
0.6
0.0
Tortricodes alternella (Denis et Schiffermüller, 1775)
1.8
1.4
3.6
0.9
Tortrix viridana (Linnaeus, 1758)
4.9
0.7
1.2
0.3
Zeiraphera isertana (Fabricius, 1794)
1.8
0.4
1.8
0.3
Animal Biodiversity and Conservation 36.1 (2013)
27
Apéndice 1. Lista de las especies de lepidópteros registradas en Q. dalechampii en las nueve parcelas del
estudio ubicadas en los Pequeños Cárpatos con la dominancia (%), los meses de presencia de las larvas
(MO), el grupo tróico (TG) (S2: oligófagas estrictas; S3: especies oligófagas más amplias; G: generalistas; U:
desconocido) y la especialización tróica de las larvas y los gremios estacionales (SG: FIF. Se alimentan durante
la brotación; LSF. Se alimentan al inal de la primavera; SF. Se alimentan en verano; AF. Se alimentan en
otoño). (Para consultar las abreviaturas de las parcelas del estudio, véase el apartado Material and methods).
HH
LL
LH
NA
NK
MO
TG
SG
0.0
0.0
0.0
0.0
0.0
5
G
FIF
0.0
0.0
0.0
1.5
0.4
6
G
LSF
0.0
0.0
0.0
0.0
0.0
7
U
SF
2.3
0.0
1.1
3.0
2.0
5–6
S2
FIF
0.0
0.0
1.1
0.0
0.0
5
G
FIF
2.3
0.9
1.1
0.8
0.4
5–6
G
FIF
4.5
0.5
0.0
3.8
0.4
6–9
G
SF
0.0
0.5
1.1
0.8
3.9
5–8
G
FIF
6.8
3.2
1.6
2.3
3.5
5–8
G
LSF
0.0
0.0
0.0
0.0
0.0
5
G
FIF
0.0
0.0
0.0
0.0
0.2
5
S2
FIF
0.0
6.0
2.7
3.8
5.0
4–6
S2
FIF
0.0
0.0
0.0
0.0
5.2
4–5
G
FIF
0.0
0.0
0.0
0.0
0.0
5
S2
FIF
0.0
0.0
0.0
0.0
0.0
5
G
FIF
2.3
0.0
0.5
0.0
0.0
5
G
FIF
0.0
0.0
1.1
0.0
0.0
5–6
S2
FIF
0.0
9.7
5.4
10.5
0.0
4–5
S2
FIF
0.0
0.0
1.1
0.0
0.4
5–6
G
FIF
0.0
0.5
0.0
0.0
0.0
5
G
FIF
0.0
0.5
0.0
0.0
0.0
5
S2
FIF
0.0
0.0
0.0
0.0
0.2
5
S2
FIF
0.0
0.0
3.8
1.5
0.7
5–7
G
FIF
0.0
0.5
0.0
0.0
0.0
5
G
FIF
0.0
1.4
0.5
0.0
0.7
5–6
G
FIF
0.0
0.0
0.0
0.8
0.2
5
G
FIF
0.0
0.0
0.0
0.0
0.0
5
G
FIF
4.5
1.4
1.1
1.5
0.9
5
G
FIF
0.0
9.3
0.0
2.3
1.3
4–5
S2
FIF
0.0
0.0
1.1
0.0
0.9
4–5
S2
FIF
Kulfan et al.
28
Appendix 1. (Cont.)
Families and species
VI
CA
FU
LI
0.3
0.0
1.2
0.3
Lycaenidae
Favonius quercus (Linnaeus, 1758)
Pyralidae
Acrobasis repandana (Fabricius, 1798)
0.0
0.0
0.0
0.0
Acrobasis tumidana (Denis et Schiffermüller, 1775)
0.0
0.0
0.0
0.0
Phycita roborella (Denis et Schiffermüller, 1775)
0.9
0.4
0.0
0.9
0.0
0.4
0.6
0.0
Agriopis aurantiaria (Hübner, 1799)
2.5
0.7
0.6
0.6
Agriopis leucophaearia (Denis et Schiffermüller, 1775)
3.1
1.4
1.8
6.8
Agriopis marginaria (Fabricius, 1776)
3.4
4.2
3.6
8.0
Drepanidae
Watsonalla binaria (Hufnagel, 1767)
Geometridae
Alcis repandata (Linnaeus, 1758)
0.0
0.0
0.0
0.0
Alsophila aceraria (Denis et Schiffermüller, 1775)
0.3
0.4
0.0
1.5
Alsophila aescularia (Denis et Schiffermüller, 1775)
4.9
2.1
0.6
1.8
Apocheima hispidaria (Denis et Schiffermüller, 1775)
0.0
0.0
0.0
0.0
Biston betularia (Linnaeus, 1758)
0.0
0.4
1.2
0.0
Biston strataria (Hufnagel, 1767)
0.0
0.0
0.0
0.0
Campaea margaritaria (Linnaeus, 1761)
1.8
0.0
0.0
0.3
Colotois pennaria (Linnaeus, 1761)
1.5
0.4
0.0
1.8
Cyclophora linearia (Hübner, 1799)
0.3
3.2
8.4
2.5
Cyclophora punctaria (Linnaeus, 1758)
0.0
0.4
0.0
0.0
Ennomos autumnaria (Werneburg, 1859)
0.0
0.4
0.0
0.0
Ennomos erosaria (Denis et Schiffermüller, 1775)
0.9
0.0
0.6
0.3
Ennomos quercinaria (Hufnagel, 1767)
0.0
0.4
0.0
0.0
Epirrita dilutata (Denis et Schiffermüller, 1775)
1.2
7.7
1.2
0.9
Erannis defoliaria (Clerck, 1759)
1.8
2.1
0.0
0.6
Hypomecis punctinalis (Scopoli, 1763)
0.6
0.0
0.0
0.3
Lomographa temerata (Denis et Schiffermüller, 1775)
0.0
1.1
0.0
0.6
Lycia hirtaria (Clerck, 1759)
0.0
0.0
0.0
0.3
Operophtera brumata (Linnaeus, 1758)
11.4
22.9
10.2
8.0
Parectropis similaria (Hufnagel, 1767)
0.0
0.0
0.0
0.0
Peribatodes rhomboidaria (Denis et Schiffermüller, 1775)
0.0
0.0
0.0
0.0
Phigalia pilosaria (Denis et Schiffermüller, 1775)
0.0
0.0
0.0
0.0
Selenia lunularia (Hübner, 1788)
0.0
0.0
0.0
0.3
Selenia tetralunaria (Hufnagel, 1767)
0.0
0.0
0.0
0.0
Drymonia ruicornis (Hufnagel, 1766)
0.3
0.0
2.4
0.3
Phalera bucephala (Linnaeus, 1758)
0.0
0.0
0.0
0.0
Notodontidae
Spatalia argentina (Denis et Schiffermüller, 1775)
0.0
0.0
0.0
0.0
Thaumetopoea processionea (Linnaeus, 1758)
0.0
0.0
0.0
0.6
Animal Biodiversity and Conservation 36.1 (2013)
HH
LL
LH
0.0
0.0
0.5
0.0
0.0
0.0
2.3
0.0
NA
29
NK
MO
TG
SG
0.0
0.6
5
S2
FIF
0.0
0.8
0.0
5
S2
FIF
1.6
0.0
0.7
5
S2
FIF
0.5
1.1
0.0
0.9
4–5,9
S2
FIF
0.0
0.9
0.0
0.8
0.0
6,8,10
S3
AF
0.0
3.7
2.2
0.8
1.3
4–6
G
FIF
2.3
7.9
7.1
1.5
8.5
4–5
S3
FIF
2.3
0.9
7.6
1.5
5.7
4–5
G
FIF
0.0
0.0
0.5
0.0
0.0
9
G
AF
0.0
2.8
0.5
0.8
0.7
4–5
G
FIF
0.0
5.6
5.4
3.0
3.1
4–6
G
FIF
0.0
0.5
0.0
0.0
0.0
5
G
FIF
2.3
0.9
0.0
0.0
0.0
8–10
G
AF
0.0
0.0
0.5
0.0
0.0
5
G
FIF
0.0
1.4
2.2
3.8
1.3
4–9,11
G
LSF
0.0
0.0
1.6
1.5
3.3
4–5
G
FIF
11.4
3.2
3.3
8.3
2.8
6–10
S3
LSF
0.0
0.0
0.5
0.0
0.0
6–7
S3
LSF
0.0
0.9
0.0
1.5
0.0
5–7
G
LSF
0.0
0.0
0.0
0.0
0.0
5,9
S3
FIF
0.0
0.0
0.5
0.0
0.0
5–6
G
FIF
2.3
0.5
0.5
2.3
1.5
4–5
G
FIF
0.0
0.5
0.0
0.0
0.7
4–5
G
FIF
0.0
0.0
0.0
0.0
1.3
6–9
G
AF
0.0
0.0
0.0
0.0
0.4
7–8
G
SF
0.0
0.0
0.0
0.0
0.4
5
G
FIF
4.5
15.7
15.8
5.3
9.2
4–5
G
FIF
0.0
0.9
0.0
0.0
0.0
7,10
G
AF
0.0
0.0
0.0
0.8
0.0
11
G
AF
0.0
0.0
0.0
0.0
0.2
5
G
FIF
0.0
0.0
0.0
0.0
0.0
6
G
LSF
0.0
0.0
0.5
0.0
0.0
6
G
LSF
0.0
0.0
0.0
0.0
0.9
5
S2
FIF
4.5
0.0
0.0
0.0
0.0
7
G
SF
0.0
0.0
0.5
0.0
0.2
6–7
G
SF
0.0
0.0
0.0
0.0
4.4
5–6
S2
FIF
Kulfan et al.
30
Appendix 1. (Cont.)
Families and species
VI
CA
FU
LI
0.0
0.4
0.0
0.0
Erebidae
Amata phegea (Linnaeus, 1758)
Calliteara pudibunda (Linnaeus, 1758)
1.2
0.0
0.0
0.9
Lymantria dispar (Linnaeus, 1758)
16.3
1.8
3.0
11.4
Orgyia antiqua (Linnaeus, 1758)
0.0
0.0
0.0
0.0
Nolidae
Bena bicolorana (Fuessly, 1775)
0.3
0.4
0.0
0.3
Nycteola revayana (Scopoli, 1772)
0.0
0.0
0.6
0.0
Pseudoips prasinana (Linnaeus, 1758)
1.2
4.2
3.0
2.8
Acronicta auricoma (Denis et Schiffermüller, 1775)
0.0
0.7
0.0
0.3
Agrochola helvola (Linnaeus, 1758)
0.0
0.0
0.0
0.0
Amphipyra pyramidea (Linnaeus, 1758)
0.0
0.0
0.6
1.5
Noctuidae
Colocasia coryli (Linnaeus, 1758)
0.0
0.4
0.0
0.0
Cosmia pyralina (Denis et Schiffermüller, 1775)
5.2
2.5
0.0
0.0
Cosmia trapezina (Linnaeus, 1758)
5.5
12.0
12.0
14.5
Dichonia convergens (Denis et Schiffermüller, 1775)
2.8
0.7
1.8
0.6
Dryobotodes eremita (Fabricius, 1775)
0.0
0.0
4.2
0.0
Eupsilia transversa (Hufnagel, 1766)
0.6
2.8
1.2
0.3
Lithophane ornitopus (Hufnagel 1766)
0.6
2.1
3.0
0.9
Moma alpium (Osbeck, 1778)
0.0
0.0
0.0
0.0
Noctuidae species 1
0.0
0.0
0.0
0.0
Noctuidae species 2
0.0
0.0
0.0
0.0
Noctuidae species 3
0.0
0.0
0.0
0.0
Noctuidae species 4
0.0
0.0
0.0
0.9
Noctuidae species 5
0.0
0.0
0.0
0.3
Noctuidae species 6
0.0
0.0
0.0
0.0
Noctuidae species 7
2.2
0.0
0.0
0.0
Orthosia cerasi (Fabricius, 1775)
5.5
2.1
0.0
3.4
Orthosia cruda (Denis et Schiffermüller, 1775)
2.2
1.1
1.8
2.5
Orthosia gothica (Linnaeus, 1758)
0.0
2.1
0.0
0.6
Orthosia incerta (Hufnagel, 1776)
0.0
0.4
0.0
0.0
Orthosia opima (Hübner, 1809)
0.0
0.0
2.4
3.4
No individuals
325
284
167
325
No species / taxons
38
46
35
53
Animal Biodiversity and Conservation 36.1 (2013)
HH
LL
LH
2.3
0.0
0.0
NA
31
NK
MO
TG
SG
0.0
0.0
4–5
G
FIF
0.0
0.0
0.0
0.8
0.0
6–8
G
SF
29.5
0.0
2.2
18.0
3.5
4–7
G
FIF
0.0
0.0
0.0
0.0
0.2
6
G
LSF
0.0
0.0
0.0
0.8
0.0
4,8
S2
FIF
0.0
0.0
0.0
0.0
0.0
5
S3
FIF
6.8
0.9
1.1
1.5
1.1
6–10
G
LSF
0.0
0.0
0.0
0.0
0.0
5–6
G
LSF
0.0
0.0
0.5
0.0
0.0
5
G
FIF
0.0
0.0
0.5
0.0
0.2
5
G
FIF
0.0
0.5
0.0
0.8
0.0
6,8
G
LSF
4.5
0.5
0.0
0.0
1.5
4–5
G
FIF
0.0
5.6
9.8
0.0
5.9
4–5
G
FIF
0.0
0.5
0.0
3.0
1.3
4–5
G
FIF
0.0
0.5
0.0
0.8
0.0
5
S2
FIF
4.5
2.3
0.5
3.0
0.7
4–6
G
FIF
0.0
0.9
0.5
2.3
0.2
4–6
G
FIF
0.0
0.5
0.0
0.8
0.4
7–8
G
SF
0.0
0.0
0.0
0.8
0.0
5
U
FIF
0.0
0.0
0.0
0.8
0.0
5
U
FIF
0.0
0.0
0.0
1.5
0.0
5
U
FIF
0.0
0.0
0.0
0.0
0.0
4
U
FIF
0.0
0.0
0.0
0.0
0.0
4
U
FIF
0.0
0.0
1.6
0.0
0.0
9
U
AF
0.0
0.0
0.0
0.0
0.0
4
U
FIF
0.0
3.2
6.5
0.0
1.1
4–7
G
FIF
0.0
1.4
0.0
0.0
7.0
4–6
G
FIF
0.0
0.0
0.0
0.8
0.4
5–6
G
FIF
0.0
0.0
0.0
0.0
0.0
5
G
FIF
4–6
G
FIF
0.0
0.0
0.5
0.0
2.8
44
216
184
133
462
18
40
43
40
52