Dalechampii oak (Quercus dalechampii Ten.), an important host plant for folivorous lepidoptera larvae. M. Kulfan, M. Holecová & P. - PDF

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.,

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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., Dalechampii oak (Quercus dalechampii Ten.), an important host plant for folivorous lepidoptera larvae. Animal Biodiversity and Conservation, 36.1: 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 flush feeders and in late summer feeders. Lepidoptera taxocenosis in the study plot Horný háj (isolated forest, high density of ants) differed significantly 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 filófagos. Llevamos a cabo un análisis estructurado de las taxocenosis de larvas de lepidóp teros 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 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 final del verano. La taxocenosis de los lepidópteros en la parcela del estudio Horný háj (un bosque aislado con una elevada densidad de hormigas) difirió significativamente 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 Bratislava, Slovakia. Milada Holecová, Dept. of Zoology, Fac. of Natural Sciences, Comenius Univ., Mlynská dolina B 1, SK Bratislava, Slovakia. Corresponding author: M. Kulfan. E mail: ISSN paper: X ISSN digital: X 2013 Museu de Ciències Naturals de Barcelona 14 Kulfan et al. 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, , 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 rainfor ests (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 distrib uted 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 repre senting 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 Samples were taken from branches at a height of about m above ground with varying exposure to cardinal points. Larvae were identified 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). Ordina tion was carried out with non metric multidimensional scaling (NMDS) using the Bray Curtis dissimilarity coefficient. One way analysis of similarities (ANOSIM) was used to identify difference in species variability of the lepidoptera taxocenosis in the study plots dur ing the year. Hierarchical (nested) ANOVA was used to examine spatial (locality) and temporal (sampling months) variation in the distribution of the total abun dance, 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 significant differences in the number of individuals, number of species and species diver sity 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 classification 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 (fig. 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 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 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 year old forest at the foot of the Kamenica Animal Biodiversity and Conservation 36.1 (2013) Study plot Contour line (m a.s.l.) Malé Karpaty Mts. border Settlement GRN DFS N 48º 48' N 72 W E S Brezová pod 73 Bradlom Jablonica Vrbové Čachtice 48º 42' N Chtelnica 48º 36' N Rohožnik LL 500 Smolenice HH NA LH NK 48º 30' N 76 Stupava Lozorno º 00'E 17º 10' E CA 300 Bratislava Sv. Jur 48º 12' N 48º 06' N 48º 00' N FU Modra VI Pezinok GRN DFS: Grid references number of the Databank of Slovak fauna LI km 48º 24' N 48º 18' N 17º 20' E 17º 30' E 17º 40' E 17º 50' E 17º 18º 19º 20º 21º 22º º º 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, m a.s.l.: an 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 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 (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 years old 16 Kulfan et al. near the village of Horné Orešany, surrounded by fields 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, 17º 23' E, 340 m a. s. l.: an 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 cal careous dust from a nearby quarry. Lošonský háj (LH) (Nature Reserve), 48º 28' N, 17º 24' E, 260 m a.s.l.: an year old oak horn beam 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, 17º 31' E, 300 m a.s.l.: a small forest island, approximately year old surrounded by fields and pastures, NE oriented. Quercus dalechampii, Q. cerris and Carpinus betulus predominate in the tree layer. Naháč Katarínka (NK) (Nature Reserve), 48 33' N, 17º 33' E, 340 m a.s.l.: a 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 Trnavs ká pahorkatina hills and the others are in the Malé Karpaty Mts. Results From , a total of 2,140 Lepidoptera larvae were collected in nine study plots with Quercus dalechampii. They represented 96 species from 17 families (appen dix 1). The families Geometridae, Noctuidae and Tor tricidae 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 first time in Slovakia (cf. Hrubý, 1964; Patočka et al., 1999). A. phegea is one of six species presenting first 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 Tortrici dae and Erebidae achieved relatively high dominance, mainly due to the species Aleimma loeflingiana (Tor tricidae) 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 loeflingiana 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 (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 Total Psychidae Bucculatricidae Gracillariidae Ypsolophidae Chimabachidae Peleopodidae Coleophoridae Gelechiidae Tortricidae Lycaenidae Pyralidae Drepanidae Geometridae Notodontidae Erebidae Nolidae Noctuidae No individuals ,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 loeflingiana, Agriopis leucophaearia and Alsophila aceraria. Three lepidop tera 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 (apen dix 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 spe cies) 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 significantly different from other taxoceno ses (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 lepi doptera 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 spe cies (appendix 1). In other taxocenoses, dominance is spread to more co dominant species (Simpson s index of dominance values from to 0.086). The value of equitability was highest at FU, NK and NA (table 2). A dendrogram based on the qualitative representa tion (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) (fig. 2). Based on a qualitative quantitative similarity (Wishart s similarity ratio, complete linkage), the hierarchical classification divided the lepidoptera taxocenoses into two clusters connected on the relati vely low level of similarity (fig. 3). The first cluster con sisted of the taxocenoses HH and NA (island forests) with the lowest figures for abundance and individuals (44 and 133, respectively). The second cluster had two subclusters and included other taxocenoses. The first 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 (fig. 4). The study plot NA was also separated (although less marked so) as confirmed by Wishart s index. Table 2. Species diversity test and basic characteristics of caterpillar taxocenoses at study plots in : 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; significance levels: *** P 0.001; ** P 0.01; * = 0.01 P 0.05; ns = 0.05 P (non significant); 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 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 significación: *** P 0,001; ** 0,001 P 0,01; * = 0,01 P 0,05; ns = 0,05 P (no significativo); para consultar las abreviaturas de las parcelas del estudio, ver Material and methods). VI CA FU LI HH LL LH NA NK e c H' VI CA ns FU ns 0.346ns LI ns 1.495ns 1.127ns HH *** 3.218** 3.429** 4.162*** LL ns 0.246ns 0.106ns 1.263ns 3.387** LH ns 1.174ns 0.842ns 0.182ns 3.901*** 0.957ns NA ns 1.092ns 0.792ns 0.131ns 3.764*** 0.894ns 0.029ns NK *** 4.198*** 3.784*** 2.773** 5.653** 4.013*** 2.567** 2.201* 0 18 Kulfan et al Dissimilarity scale HH FU NA LL CA LH NK LI VI Fig. 2. Classification of lepidoptera taxocenoses on individual study plots according to species presence/ absence (Sörensen s index). Fig. 2. Clasificación de las taxocenosis de lepidópteros en cada una de las parcelas del estudio en función de la presenc
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