CONE ENTOMOFAUNA OF NORWAY SPRUCE PICEA ABIES (L.) KARST. IN SELECTED STANDS OF THE TATRA NATIONAL PARK IN POLAND

Marek Kozioł
Department of Forest Entomology, Agricultural University of Cracow, Poland

ABSTRACT

The qualitative and quantitative composition of cone entomofauna of Norway spruce Picea abies (L.) Karst. of the Tatra National Park was determined during 1987–1993. The trophic groups of insects were distinguished and characterized using ecological indexes. Some of the data were analyzed statistically using Kolmogorov-Smirnov, Bartlett’s, and Duncan’s tests (at the significance level α = 0.05). The diversity of the qualitative and quantitative composition of insects inhabiting spruce cones is presented according to their altitudinal distribution. The results obtained fill the gaps in knowledge on seed and cone pests of Norway spruce of the Tatra National Park. Above 67 thousand of insect specimens representing 46 species from 5 orders were collected. From among these species 13 were for the first time reported from Norway spruce cones in the world. Besides, other two species, to date reported from other European countries, were for the first time reported from Norway spruce cones in Poland. A significant effect of altitude on numbers of 9 species was found. Also the difference between the cone entomofauna of Norway spruce in the Western Tatra and that in the High Tatra was determined.

Key words: Norway spruce, Tatra Mountains, cones, entomofauna, altitude.

INTRODUCTION

The actual state of knowledge on insect fauna of the Tatra Mountains needs to be supplemented with new data on ecological insect communities. This especially concerns the phytophagous entomofauna of forest habitats, including cones of Norway spruce Picea abies (L.) Karst., seasonally or accidentally inhabited by various developmental stages of insect species. Studies on entomofauna of spruce cones not only fill the gaps in knowledge on forest insect fauna of the Polish Tatra but also are important from the practical point of view because some insects living in cones are considered to be serious pests of seeds. The knowledge about them makes the basis for elaboration in future of efficient control methods in order to protect seed of the most valuable populations of native races of Norway spruce in the Tatra Mountains.

The entomofauna of Norway spruce cones, composed of insects of various taxonomic groups has been of interest of many investigators. The studies on this group of insects were initiated by Trägårdh [34] and Holste [11]. They were followed by Čermak [7], Bakke [3], Györfi [10], Jakovlev [12], Arend [2], Stadnickij [30,31], Saksons [20], and Milišauskas [17]. Among other papers concerning this group of insects only few contain information including a larger number of species. They usually deal with species important from the economic point of view, i.e. those damaging Norway spruce seeds and cones, as well as with their parasitoids. For example such information has been given by Bakke [4], Cankov [6], Favero and Masutti [9], Annila [1], Roques [19], Nanu et al. [18], and Krištek and Skrzypczyńska [16].

Skrzypczyńska [21,22] published for the first time in Poland the list of the cone entomofauna of Norway spruce from the Beskid Sšdecki mountain range. Moreover, studies dealing with this problem were carried out in the Beskid Œlšski and the Beskid Żywiecki ranges [26,27,28], as well as in the Babia Góra [29], Białowieża [24,25], and Gorce [37] National Parks.

The present paper describes some results of studies which for the first time were carried out in the Tatra National Park. Other results of these studies were published in earlier papers [13,14,15].

The objectives of this study were:

1. to determine the qualitative and quantitative compositions of insects inhabiting cones of Norway spruce Picea abies (L.) Karst. in selected stands of the Tatra National Park,

2. to distinguish and characterize the trophic groups from among the species found, and

3. to determine the diversity of the qualitative and quantitative compositions of Norway spruce cone entomofauna:

• in stands of the Western Tatra and the High Tatra,

• at various altitudes.

METHODS

The field investigations were carried out in the Tatra National Park during 1987 – 1993. Cone samples of Norway spruce Picea abies (L.) Karst. were collected in 42 permanent sample plots situated every 100 m in altitude, from 1000 to 1200 m in the lower mountain zone and from 1300 to 1500 m in the upper mountain zone. Cones were also collected from spruce trees growing above the mountain forest limit, among mountain pine Pinus mugo Turra (Fig. 1).

The observation sites in the High Tatra were situated in the Sucha Woda valley and its branches, the Gšsienicowa valley, the Pańszczyca valley, branches of the Białka valley, the Waksmundzka valley, the Roztoka valley, and also in surroundings of Gęsia Szyja and the Waksmundzka Plain.

The observation sites in the Western Tatra were located in the Koœcieliska valley, the Smytnia Glade, the Tomanowa valley, the Chochołowska valley, the Jarzšbcza valley, and the Wyżna Chochołowska valley.

Also single localities were situated in the Iwanicka Pass and the Kondratowa valley.

In total 5780 cones were collected from 289 trees. Each sample contained 20 cones. In the laboratory, from each cone sample 5 cones were randomly selected for the analysis by cutting. Cones were dissected along the axis. Then the cone scales were separated and seeds were extracted and cut. In total 1445 cones with 317 165 seeds were analyzed. This procedure yielded larvae and pupae of various insect species. They were counted and identified. The remaining 15 cones from each sample were used for mass rearing in photoeclectors or glass jars. All rearing containers were kept in cool rooms with conditions close to natural ones for two years. Each year from the moment of appearance of first insects the rearing was inspected once a week.

The insects obtained during the study were grouped according to trophic associations as in studies carried out by Skrzypczyńska [21,22], and then described using ecological indexes given by Szujecki [33] (dominance – D, constancy – C, and species diversity – d). The following types of constancy were distinguished: C5 – euconstants (C>75%), C4 – constants (C = 50.01 – 75%), C3 – subconstants (C = 30.01 – 50%), C2 – accessory species (C = 15 – 30%), and C1 – accidents (C<15%). The dominance classes were as follows: eudominants (D>10%), dominants (D = 5.01 – 10%), subdominants (D = 2.01 – 5%), and recedents (D = 1 – 2%).

Some of the results were analyzed statistically. For this purpose results were transformed using the square root, while in the case of percentages an angular transformation was used. The conformity of their distribution with a normal distribution was checked using the Kolmogorov-Smirnov test. The Barlett’s and Duncan’s tests (α = 0.05) were used to study the effect of altitude on variation of variables analyzed.

RESULTS

Qualitative and quantitative analyses of insects found
In total 67 178 insect specimens representing 46 species were obtained by rearing and the analysis of Norway spruce cones. These insects belonged to the following orders: Heteroptera (1 species), Coleoptera (7 species), Hymenoptera (18 species), Lepidoptera (5 species), and Diptera (15 species) (Table 1).

The cones were most numerously inhabited by Diptera (53 880 specimens, i.e. 80.20% of all insects obtained). Quite numerous were Lepidoptera (8 774 specimens – 13.06%) and Hymenoptera (4 434 specimens – 6.60%). Representatives of the remaining orders were not numerous or rare (Coleoptera 0.1%, Heteroptera 0.03%). The following species were for the first time in the world reported from cones of Norway spruce: Trechus striatulus Putz., Oxytelus nitidulus Grav., Corticaria serrata (Payk.), Cryptophagus subfumatus Kraatz., Latridius nodifer Westw., Rhizophagus depressus (Fabr.), Stemacrus femorelis Holmgr, Phaeogenes osculator Thunb., Blacus hastatus Hal., Enizemum nigricornis Ths., Episyrphus cinctellus (Zeet), Conioscinella flavifrons Duda, and Palloptera laetabilis Loew. Moreover, Cydia illutana H.-S (Tortricidae) and Trichomalus campestris (Walker) (Pteromalidae) were for the first time reported from Norway spruce cones in Poland.

Cydia strobilella L. and Kaltenbachiola strobi (Winn.) belonged to constancy class C5 (euconstants), Plemeliella abietina Seitn. and Torymus azureus Boh. to class C4 (constants), Clinodiplosis cilicrus (Kieff.), Tetrastichus strobilanae Ratz., Camptomyia sp., Triplatygaster contorticornis Ratz., and Lestiodiplosis holstei (Kieff.) to class C3 (subconstants), and Asynapta strobi (Kieff.) and Venturia transfuga Grav. to class C2 (accessory species). The remaining species listed in Table 2 belonged to class 1 (accidents).

From among dipterans K. strobi (30 477 specimens) and P. abietina (14 351 specimens) were the most numerous species. Quite numerous were also C. cilicrus (4 054 specimens), Lycoriella solani Winn. (1 488 specimens), L. holstei (1 482 specimens), Camptomyia sp. (967 specimens), and A. strobi (662 specimens).

C. strobilella (8 695 specimens) was the most numerous lepidopteran. The remaining lepidopterans were not numerous or rare (C. illutana, Dioryctria abietella (Den. et Schiff.), Eupithecia analoga Djakonov, Eupithecia abietaria Goeze).

Chalcidoidea: T. azureus (1 279 specimens), T. strobilanae (1 202 specimens), and T. contorticornis (811 specimens), and also the representative of Pteromalidae – Anogmus sp. aff. vala (492 specimens) were the most numerously represented hymenopterans. From among Ichneumonidae V. transfuga turned out to be the most common species (383 specimens).

Corticaria serrata (Payk.) and Ernobius abietis Fabr. were the most numerous coleopterans ( 25 and 21 specimens respectively). Also single specimens of L. nodifer, C. subfumatus, R. depressus, T. striatulus and unidentified representatives of small Staphylinidae were found. O. nitidulus was represented by a single specimen. It should be mentioned that C. subfumatus and L. nodifer have been already reported from the Tatra National Park, but from silver fir cones [23].

Heteroptera were represented by one species – Gastrodes abietum Bergr.

Coenotic characteristics of insects obtained
Insects found in cones and seeds of Norway spruce may be grouped in the following manner:

1. Conophagous insects, larvae or caterpillars of which feed on various parts of cones thus damaging seed scales or seeds (8 species were obtained);

2. Seminiphagous insects, larvae of which feed exclusively inside seeds (2 species);

3. Saprophagous and coprophagous insects, larvae of which feed on dead organic matter and excrements of other insects (10 species);

4. Parasitoids and predators (18 species);

5. Seasonal insects for which cones provide shelter, frequently accidental (9 species) (Tables 2 and 3).

Among insects obtained from spruce cones there were 58.64% of conophages, 21.37% of seminiphages,, 11.09% of saprophages and coprophages, 8.80% of parasitoids and predators, and only 0.1% of seasonal insects. K. strobi and C. strobilella were eudominant species in the group of conophagous insects, while P. abietina was a strong eudominant species in the group of seminiphagous insects.

In the group of saprophagous and coprophagous insects C. cilicrus, L. solani and Camptomyia sp. belonged to eudominant species, and A. strobi was a dominant species.

In the group of parasitoids and predators the following were eudominant species: L. holstei, T. azureus, T. strobilanae, and T. contorticornis, while A. sp. aff. vala and V. transfuga were dominant species.

The eudominant species in the group of seasonal insects were as follows: C. serrata and G. abietum, while C. subfumatus was a dominant species. The remaining species listed in Table 2 may be classified as subdominant and recedent species.

The index of species diversity reached the highest value for the group of parasitoids and predators (d = 1.96), and the lowest value for the group of seminiphagous insects (d = 0.10) (Table 2).

Diversity in qualitative and quantitative compositions of insects inhabiting Norway spruce cones

In the Western Tatra and the High Tatra
In total 37 318 insect specimens representing 43 species were obtained from cones collected in the Western Tatra (278 specimens per cone sample on the average). From cones collected in the High Tatra 29 860 specimens representing 39 species were obtained (194 specimens per cone sample).

In the Western Tatra as well as in the High Tatra Diptera were the most numerous group (79.37% and 81.25% of all collected insects respectively) (Table 1, Fig. 2). In the Western Tatra the following species were more frequent and more numerous: L. holstei, A. strobi, and Camptomyia sp., while in the High Tatra: L. solani, Strobilomyia anthracina Czerny, and P. laetabilis. From among less numerous species , C. flavifrons and E. cinctelus were found only in cones of the Western Tatra, and representatives of the genus Drosophila in cones of the High Tatra.

The cones from the Western Tatra were colonized by Lepidoptera to a greater extent than the cones from the High Tatra (14.17% and 12.17% respectively).

The percentage of Hymenoptera obtained from cones of the Western Tatra (8.35%) was higher than that obtained from cones of the High Tatra (4.41%). From among hymenopterans obtained from the Western Tatra cones, T. azureus, T. strobilanae, T contorticornis, V. transfuga, and A. sp. aff. vala were quite frequent and quite numerous. A. piceae, T. campestris, E. nigricornis, and M. strobilobius were found only in the Western Tatra cones, while P. osculator and B. hastatus only in the High Tatra cones.

From among Coleoptera found in cones from the High Tatra, C. serrata turned out to be a quite numerous species, while G. abietum (Heteroptera) was quite numerous in cones from the Western Tatra.

In both Tatra ranges K. strobi and C. strobilella belonged to the group of euconstants (Table 2). From among constants, besides the mutual species P. abietina, also T. azureus belonged to this group in cones in the Western Tatra, while in the High Tatra it was less frequent. There were more subconstants in cones of the Western Tatra. Besides mutual species, i.e. Camptomyia sp., C. cilicrus, and T. strobilanae, also the following species belonged to the group of subconstants in cones of the Western Tatra: A. strobi, T. contorticornis, V. transfuga, and L. holstei. While in cones of the High Tatra there were considerably more accessory species (L. holstei, T. contorticornis, V. transfuga, A. strobi, and S. anthracina). In cones of the Western Tatra A. sp. aff. vala was an accessory species. From among accidental species occurring in cones in the Western Tatra the following species should be mentioned: Anogmus hohenheimensis Ratz., Torymus caudatus (Boh.), S. anthracina, L. solani, C. illutina, and G. abietum, and in cones of the High Tatra: L. solani, A. hohenheimensis, A. sp. aff. vala, L. strobilorum, and C. serrata.

There were more conophagous species in cones collected in the Western Tatra (Table 2, Fig. 3). In spite of the presence of mutual eudominants in this trophic group (C. strobilella and K. strobi), there were more individuals of C. illutana and E. abietaria in cones from the Western Tatra, while in cones of the High Tatra S. anthracina was more numerous. The percentage of seminiphagous species represented by P. abietina was higher in cones from the High Tatra, where also single specimens of M. strobilobius were found. The saprophagous species were more numerous in cones in the High Tatra than in the Western Tatra (14.39% and 8.45% respectively). The following were the mutual eudominants of this trophic group: C. cilicrus and L. solani. In addition, in the Western Tatra Camptomyia sp. and A. strobi were the eudominant species, which in the High Tatra were less numerous . In the High Tatra Drosophila sp. was a subdominant species among saprophages, while P. laetabilis belonged to the recedents. Parasitoids and predators were more numerous in cones of the Western Tatra (11.4%) than in cones of the High Tatra (5.54%), but the species diversity of this trophic group was higher for insects obtained from cones collected in the High Tatra. L. holstei, T. azureus, T. strobilanae, and T. contorticornis were the mutual eudominants of this trophic group. In cones from the High Tatra A. sp. aff. vala and V. transfuga were also eudominant species, while in cones from the High Tatra they belonged to dominant species. A. hohenheimensis was a mutual subdominant species in the group of parasitoids and predators.

The calculated indexes of species diversity turned out to be higher in the group of seasonal insects found in cones from the Western Tatra. In this group of insects C. serrata was a mutual eudominant, while in cones of Western Tatra also G. abietum was an eudominant species. C. subfumatus and L. nodiffer were found in both the High Tatra and the Western Tatra.

Diversity depending on altitude
The largest number of insects was obtained from cones collected at altitudes from 1400 to 1500 m (309 specimens per cone sample on the average), and the smallest number at altitudes from 1000 to 1100 (194 specimens per cone sample on the average).

Diptera were the most numerous in cones collected from trees at altitudes 1000, 1500, and above 1500 m (Fig. 2). The largest number of Lepidoptera was present in cones collected at altitudes 1100, 1200, and 1300 m. Hymenoptera were most numerous in cones collected at altitudes 1100, 1400, and above 1500 m. Coleoptera were most common in cone samples collected from trees growing at 1000, 1300, and 1400 m in altitude.

The conophagous insects were most numerous in upper parts of the upper mountain zone, in cones collected from trees growing at altitudes 1400, 1500, and above 1500 m (Fig. 3). The seminiphagous insects prevailed in cones collected in lower parts of the lower mountain zone at altitudes 1000 and 1100 m. The saprophagous and coprophagous insects were most numerous in cones at altitudes 1000 and 1200 m. The parasitoids and predators were quite numerous at altitudes 1100 and 1200 m of the lower mountain zone, and in the case of the upper mountain zone at altitudes 1400 and above 1500 m.

A more detailed analysis of qualitative and quantitative altitudinal distribution included all insect species found in greater numbers in all observation areas. The percentages of these species in spruce cones collected at various altitudes are presented in Figs. 4 and 5 in order to show existing diversity.

From among the species from the family Cecidomyiidae, C. cilicrus, Camptomyia sp., and A. strobi were most numerous in cones collected from trees growing at altitudes 1000 – 1200 m.

P. abietina did not show any distinct altitudinal quantitative diversity. Generally, this species was slightly more numerous in cones collected in the lower mountain zone. This seminiphagous insect was most numerous in cones collected at altitude of 1000 m (High Tatra and Western Tatra) and at altitude of 1400 m (High Tatra).

S. anthracina was most frequently found in cones collected in the upper mountain zone, especially at altitudes 1300 and 1500 m. However, taking into account a small number of specimens obtained (129), this problem requires further investigations.

The results of statistical analyses showed that altitude has a significant effect on variation of numbers of the following insect species inhabiting Norway spruce cones: C. strobilella, K. strobi, T. azureus, T. strobilanae, T. contorticornis, A hohenheimensis, A. sp. aff. vala, V. transfuga, and L. holstei.

C. strobilella in Norway spruce stands of the Tatra Mountains prefers the upper part of the lower mountain zone (1200 m) and the lower and middle parts of the upper mountain zone (1300 and 1400 m). In the High Tatra this tortricid was most numerous at altitude of 1200 m, and in the Western Tatra at altitude of 1300 m.

K. strobi appeared to be a species occurring in greater numbers in the upper mountain zone, especially at altitude of 1400 m. In the High Tatra it was most numerous at the mountain forest limit (>1500 m), and in the Western Tatra at altitude of 1400 m.

There seem to be a significant relationship between the insect numbers and the height above the sea level in the case of individual species of Chalcidoidea. T. contorticornis turned out to be more numerous than other Chalcidoidea at the mountain forest limit (1500 m). This was confirmed by results of the Duncan’s test. The highest mean numbers of this species obtained from cone samples collected at altitude of 1500 m (9.6) and above 1500 m (12.7) were significantly different from the means for cone samples collected at altitudes from 1000 to 1300 m (mean from 0.28 to 2.4).

The numbers of T. strobilanae, as well as its percentage among Chalcidoidea, were significantly higher in the middle part of the upper mountain zone (1400 m in altitude).

T. azureus occurred especially abundantly in stands of the lower mountain zone. The mean numbers of this species obtained from cones collected at altitudes 1000 and 1200 m were significantly different from the means obtained for cones collected in the upper mountain zone.

A. sp. aff. vala was quite numerous in cones in stands of the upper mountain zone (1400, 1500, and >1500 m). Mean numbers of this species obtained from samples collected above 1500 m were significantly different from the means for cones collected from trees growing at altitudes 1100 – 1300 m and 1500 m.

A less numerous A. hohenheimensis also seemed to be more strongly associated with stands of the lower mountain zone. Mean numbers of this species obtained from cones collected at altitudes 1000 and 1200 m were significantly different from the mean numbers found in cones collected at altitudes 1300 and 1500 m. Perhaps this was connected with a slightly greater numbers at these altitudes of P. abietina, the main host of this parasitoid.

From among Ichneumonidae, V. transfuga was most numerous at similar altitudes as its host C. strobilella, i.e. 1200 – 1400 m. The mean numbers of specimens obtained from cone samples collected at altitude of 1400 m were significantly different from mean values for this species obtained for cones collected at altitudes 1000, 1100, 1500, and above 1500 m, but there were no significant differences in the case of altitudes 1200 and 1300 m.

The predatory dipteran L. holstei was distinctly more numerous in cones collected from trees growing at elevations 1000 – 1200 m. The mean numbers of specimens of this species present in cones collected at altitude of 1200 m were significantly different from means obtained from cones collected at the remaining altitudes.

DISCUSSION

The results presented in this paper showed that cone entomofauna of Picea abies (L.) Karst., in the Tatra National Park is rich in respect of quality as well as quantity. In total, 67 178 insect specimens were obtained from cone and seed analyses and rearing. They represented 46 species from 5 orders. Hymenoptera and Diptera were the most diverse groups. Out of the species found during this study 13 were for the first time in the world reported from Norway spruce cones. Moreover, a conophagous species Cydia illutana H.-S. was reported from Norway spruce cones for the first time in Poland. To date, this species was reported from other European countries [7,8]. Also parasitoid Trichomalus campestris (Walker) was for the first time in Poland reported from cones of Norway spruce.

In the case of the dipteran Palloptera laetabilis Loew., for the first time found in cones of Norway spruce, additional studies should be undertaken in order to explain its bioecological association with cones.

Anogmus sp. aff. vala, a most numerous representative of the family Pteromalidae (Chalcidoidea) in Norway spruce cones, is probably, according to Dr. Boučka (information from the letter), a new species.

The colonization of Norway spruce cones by K. strobi was in the Tatra National Park exceptionally high, to date not encountered on such a level elsewhere in Poland [14], while C. strobilella and P. abietina turned out to be very serious pests of spruce seeds and cones in stands under investigations [13,14,15]. A seminiphagous species M. strobilobius was not numerous, similarly as in other sites in Poland [21,22,25,27,28,29,37].

Similarly as in the Beskid Sšdecki [21,22], the Beskid Slaski, and the Beskid Żywiecki [27] mountain ranges C. cilicrus and L. solani were in the Tatra National Park the eudominant species in the group of saprophagous and coprophagous insects.

A predatory dipteran L. holstei, quite abundantly encountered during this study, probably greatly limited populations of species from families Cecidomyiidae and Sciaridae mentioned above. It is interesting that in cones in the Beskid Sšdecki, the Beskid Żywiecki, the Gorce National Park, and the Babia Góra National Park ichneumon Liotryphon strobilorum (Thomson) and braconid Baecis abietis Ratz. (unfound during the present study) were much more numerous than V. transfuga [21,22,25,27,28,29,7].

The ichneumon P. osculator, a known parasitoid of Zeiraphera griseana Hb. [5], and for the first time obtained from rearing of Norway spruce cones, turned out to be a new parasitoid of C. strobilella. The author has not found any earlier information concerning this problem. The ichneumon S. femorelis and braconid B. hastatus appeared to be the species for the first time reported from cones of Norway spruce. The determination of hosts of these parasitoids would require undertaking of further studies.

The cones collected in Norway spruce stands of the upper mountain zone were much more abundantly colonized by insects than cones collected in lower parts of the lower mountain zone. The cone entomofauna in the upper mountain zone was also much more diverse than that of the lower mountain zone.

Dipterans and hymenopterans prevailed in the upper mountain zone, while lepidopterans and coleopterans at medium altitudes (1200 – 1300 m).

Cydia strobilella in the Tatra National Park preferred altitudes of the upper part of the lower mountain zone (1200 m) and the lower part of the upper mountain zone (1300 m). This confirmed conclusions of other authors that this species is rather associated with mountain stands of medium altitudes [36,35,32].

Kaltenbachiola strobi turned out to be the species occurring more numerously in the upper mountain zone. This is in agreement with results of Cankov [6], who found that this species in Bulgaria was also more numerous at higher altitudes.

T. strobilanae occurred at higher altitudes also in Norway [4].

According to Bakke [4] in Norway A. vala and T. azureus are more common in lowland and in southern part of the country. T. azureus, a numerous species in this study, seemed to be associated with Norway spruce stands of the lower mountain zone in the Tatra National Park, while A. sp. aff. vala occurred more numerously at mountain forest limit (1500 m).

SUMMARY OF RESULTS AND CONCLUSIONS

1. Founding of 46 insect species (67 178 specimens) from 5 orders showed that the cone entomofauna of Norway spruce Picea abies (L.) Karst. in the Tatra National Park is relatively rich.

2. From among insects found the following species were for the first time reported from Norway spruce cones: Trechus striatulus Putz., Oxytelus nitidulus Grav., Corticaria serrata (Payk.), Latridius nodifer Westw., Rhizophagus depressus (Fabr.), Cryptophagus subfumatus Kraatz., Stenomacrus femorelis Holmgr., Phaeogenes osculator Thunb., Blacus hastatus Hal., Enizemum nigricornis Ths., Episyrphus cinctellus (Zeet), Conioscinella flavifrons Duda, and Palloptera laetabilis Loew. Moreover, Cydia illutana H.-S. and Trichomalus campestris (Walker) were for the first time found in Norway spruce cones in Poland.

3. Diptera, Lepidoptera, and Hymenoptera appeared to be the most numerous insects in cones investigated during this study. Hymenoptera and Diptera were the most diversified orders in respect of species composition.

4. The following were the most abundantly occurring species: Kaltenbachiola strobi (Winn.), Plemeliella abietina Seitn., Cydia strobilella L., Clinodiplosis cilicrus Kieff., Lycoriella solani Winn., Lestdiplosis holstei Kieff., Torymus azureus Boh., and Tetrastichus strobilanae Ratz.

5. It may be supposed, on the basis of the index of constancy (C), that the following species were the most characteristic for the examined cones: Cydia strobilella L., Kaltenbachiola strobi Winn., Plemeliella abietina Seitn., and Torymus azureus Boh.

6. The conophagous and seminiphagous species were the most numerous insects among trophic groups distinguished, while the group of parasitoids and predators was most diverse in respect of species composition.

7. The recognition of trophic associations of Drosophila sp., Palloptera laetabilis Loew., Stenomacrus femorelis Holmgr., and Blacus hastatus Hal. in Noeway spruce cones requires further investigations, because these species are probably associated bioecologically with cones.

8. The insect numbers as well the number of species in Norway spruce cones in the Western Tatra were distinctly greater in comparison with entomofauna obtained from cones collected in the High Tatra.

9. Among Diptera the following species were more frequent and more numerous in the Western Tatra: Lestodiplosis holstei Kieff., Asynapta strobi Kieff., and Camptomyia sp., while in the High Tatra: Lycoriella solani Winn., Strobilomyia anthracina Czerny, and Palloptera laetabilis Loew. The following species were found in cones from the Western Tatra only: Conioscinella flavifrons Duda and Episyrphus cinctellus (Zeet), and in cones from the High Tatra only: Drosophila sp.

10. There were more Lepidoptera in cones collected in the Western Tatra, including less numerous species Cydia illutana H.-S. and Eupithecia abietaria Goeze.

11. Hymenoptera were more abundant in cones from the Western Tatra. Out of them the following species were more frequent and more numerous than in the High Tatra: Torymus azureus Boh., Tetrastichus strobilanae Ratz., Triplatygaster contorticornis Ratz., Venturia transfuga Grav., and Anogmus sp. aff. vala (Walker). The species found in cones of the Western Tatra only were as follows: Anogmus piceae (Ruschka), Trichomalus campestris (Walker), Enizemum nigricornis Ths., and Megastigmus strobilobius Ratz., and in the High Tatra only: Phaeogenes osculator Thunb., and Blacus hastatus Hal.

12. Corticaria serrata (Payk.) (Coleoptera) in cones from the High Tatra and Gastrodes abietum Bergr. (Heteroptera) in cones from the Western Tatra turned out to be the quite numerously occurring species.

13. The height above the sea level had a significant effect on numbers of the following insect species: Cydia strobilella L., Kaltenbachiola strobi Winn., Lestodiplosis holstei (Kieff.), Torymus azureus Boh., Tetrastichus strobilanae Ratz., Triplatygaster contorticornis Ratz., Anogmus hohenheimensis Ratz., Anogmus sp. aff. vala (Walker), and Venturia transfuga Grav.

14. Cydia strobilella L. preferred altitudes of the upper part of the lower mountain zone (1200 m) and the lower part of the upper mountain zone (1300 m), while its parasitoid Venturia transfuga Grav. altitudes 1200 – 1400 m.

15. Kaltenbachiola strobi (Winn.) occurred in the highest intensity in the upper mountain zone, while its parasitoids Torymus azureus Boh. and Anogmus hohenheimensis Ratz. in the lower mountain zone, Triplatygaster contorticornis Ratz. at the mountain forest limit, Tetrastichus strobilanae Ratz. and Anogmus sp. aff. vala (Walker) in the upper mountain zone.

16. Lestidiplosis holstei (Kieff.) occurred more numerously in stands of the lower mountain zone.

ACKNOWLEDGEMENTS

I would like to express my gratitue for the first identification and a verification of my identification of selected insects to: Bolesław Burakowski, PhD (Zoology Institute of Polish Academy of Science, Warsaw) (Coeloptera), Doc.Miroslav Čapek, PhD (Vyzkumny ústav lesného hospodarstva, Banská Štavnica, Slovak Republic) (Braconidae), Josef Šedivy, Phd (Ustav Ochrany Rostlin, Praha - Czech Republic) (Ichneumonidae), Zdenek Bouček, PhD (British Museum - Natural History, London) (Chalcidoidea), Jerzy Lis, Phd (Museum of Natural History – Bytom) (Heteroptera), Andrzej Palaczyk, MSc (Institute of Systematic Zoology and Animal Ecology of Polish Academy of Science, Cracow) (Diptera), Janusz Sawoniewicz, PhD (Zoology Institute of Polish Academy of Science, Warsaw) (Ichneumonidae).

I feel particularly obliged to Prof. Małgorzata Skrzypczyńska, PhD for all the help provided as well as her kindness

REFERENCES

1. Annila E. 1981. Kuusen käpy- ja siementuholaisten kannanvaihtelu [Fluctuations in cone and seed insect populations in Norway spruce]. Comm. Inst. For. Fenn. 101, 1-32 [in Finnish].

2. Arend H. 1967. Über den tierischen Abbau von Fichtenzapfen [The breakdown of Spruce cones by animals]. Z.ang. Ent. 59, 74-109 [in German].

3. Bakke A. 1955. Insects reared from spruce cones in northern Norway 1951. Norsk Ent. Tidsskr. 9, 152-212.

4. Bakke A. 1963. Studies on the spruce-cone insects Laspeyresia strobilella (L.) (Lepidoptera, Tortricidae), Kaltenbachiola strobi (Winn.)(Diptera, Itonididae) and their parasites (Hymenoptera) in Norway. Meddels. Frau. Det Norsk. Skogfors. Nr. 67, 1-151.

5. Baltensweiler W. 1958. Zur Kenntnis der Parasiten des Grauen Lärchenwicklers (Zeiraphera griseana Hübner) im Oberengadin [Monograph on the Knowledge of Parasites Zeiraphera griseana Hübner in Oberengadin]. Mitt. Eidgen. Anst. Forstl. Vorsw.34, 339-477 [in German].

6. Cankov G. V. 1974. Semejadi i technite entomofagi pro smrca Picea excelsa (Link.) v njakoj tipove gora v gorsko stopanstvo “Vasil Kolarov” [Seed Insects And Their Parasites in Norway Spruce (Picea excelsa (Link.) in Selected Forest Types in the “Vasil Kolarov” Forest Farm.]. Gorsko stop. nauka 11, 61-67 [in Bulgarian].

7. Čermak K. 1952. Hmyzi škudci semen našich lesnich dřevin [Insects attacking seeds of Czechoslovakian forest species]. Prace vyzk. ust. lesn. I. Lesn. Knih. 5, 5-87 [in Czech].

8. Champon J. P., Roques A., Genestier G. 1992. Observations sur quelques Lépidoptéres Tortricidae peu observis ou nouveaux pour la faune de France [Observations on some Lepidoptera Tortricidae rarely seen or new to the fauna of France]. Nouv. Revue Ent.(N.S.), T.9, Fasc 2, 107-117 [in Franch].

9. Favero R. D., Masutti L. 1974. Animali e strobili di abete rosso in alcuni boschi delle Alpi Orientali [Animals and cones of Norway Spruce in some forests of the eastern Alps]. Monti e Boschi, 25, 3-16 [in Italish].

10. Györfi J. 1956. Nadelholzzapfen- und Nadelholzsamenschädlinge und ihre Parasiten [Cone and seed pests of conifers and their parasites]. Acta Agron. Acad. Scient. Hung. 5, 321-373 [in German].

11. Holste G. 1922. Fichtenzapfen- und Fichtensamenbewohner Oberbayerns [Inhabitants of Cones and Seeds of Spruce in Oberbayerns] Z. ang Ent. 8, 125-160 [in German].

12. Jakovlev B. P. 1961. Vrediteli šišek i semjan eli [Parasites of Spruce Cones And Seeds]. Petrozavodsk, Gosud. Izd. Karel’skoj ASSR [in Russian].

13. Kozioł M. 1997. Influence of altitude on adult emergence, abundance and dynamics of Cydia strobilella populations in the Tatra National Park, Poland. Proceedings of the 5th Cone and Seed Insects Working Party Conference (IUFRO S7.03-01), September 1996, Monte Bondone, Italy. Padova, Institute of Agricultural Entomology. University of Padova, 75-85.

14. Kozioł M. 1998. Influence of altitude on adult emergence and dynamics of. Kaltenbachiola strobi. (Winnertz). (Diptera, Cecidomyiidae). populations and its parasites in the Tatra National Park, Poland. Anz. Schädlingskde., Pflanzenschutz, Umweltschutz. 71, 121-127.

15. Kozioł M. 2000. Cono- and seminiphagous insects of Norway spruce Picea abies (L.) Karst. And their parasitoids in lower and upper montane zone of the Tatra National Park in Poland. J. Appl. Ent. 124, 259-266.

16. Křistek J., Skrzypczyńska M. 1992. Živočišní škudci semen, šišek a plodů lesních dřevin [Organisms Damaging Seeds, Cones And Fruits of Forest Trees]. In, J. Křistek. Škudci semen, šišek a plodů lesních dřevin. Brazda, Praha, 11-101 [in Czech].

17. Milišauskas Z. 1976. Lietuvos TSR paprastosios egles (Picea abies Karst.) kankoŕežių entomofauna [The insect fauna of cones of Norway spruce (Picea abies Karst.) in the Lithuanian SSR]. Acta ent. Lituan. Vilnius 3, 37-47 [in Lithuania].

18. Nanu N., Lăcătusu M., Tudor C. 1986. The balance factor in the conifer merocenoses in Romania. Proc. 2nd Conf. IUFRO Cone and Seed Insects Working Party. S2.07-01, Briançon, (France), Sept. 3-5, 1986, 91–99 [in Romanien].

19. Roques A. 1983. Les insectes ravageurs des cones et graines de conifers en France [Insect pests of cones and seeds of conifers in France]. INRA Paris [in Franch].

20. Saksons JA. L. 1973. Entomofauna generativnych organov sosny (Pinus silvestris L.) i eli (Picea abies Karst.) v Latvinskoj SSR [The entomofauna of the generative organs of Scots pine (Pinus silvestris L.) and Norway spruce (Picea abies Karst.) in the Latvian SSR]. Zašč. Lesa, Riga, 29–52 [in Russian].

21. Skrzypczyńska M. 1982. The entomofauna of the cones of spruce, Picea abies ( L.) Karst. in Poland. Zeit. ang. Ent. 94, 21-32.

22. Skrzypczyńska M. 1986. Insects of cones and seeds of Norway spruce, Picea abies (L.)Karst. in Poland. Proceedings of the 2nd Conf.IUFRO W.P S2.07-01, Briançon, France, 27-38.

23. Skrzypczyńska M. 1988. Entomofauna of cones of fir (Abies alba Mill.) in the Tatra National Park in Poland. J. Appl. Ent. 105, 217-222.

24. Skrzypczyńska M. 1989. Studies on the occurrence and parasitization of the spruce seed pest, Megastigmus strobilobius Ratz. (Hym.,Torymidae) and Plemeliella abietina Seit. (Dipt., Cecidomyiidae) in Poland during a poor seed year. J. Appl. Ent.108, 138-143.

25. Skrzypczyńska M. 1994. Über die in Zapfen und Samen der Fichte Picea abies (L.) Karst. fressenden Insekten und ihre Parasitoiden in Nordost-Polen [Insects feeding in the cones and seeds of Norway spruce Picea abies (L.) Karst. and their parasitoids in north-eastern Poland]. Anz. f. Schädl. 67, 45-46 [in German].

26. Skrzypczyńska M., Bysko G. 1997. Owady zasiedlajšce szyszki swierka pospolitego Picea abies(L.) Karst. w wybranych drzewostanach Beskidu Slaskiego [Insects inhabiting Cones of Norway Spruce Picea abies (L.) Karst. in Selected Stands of the Beskid Slaski Mts.]. Sylwan 141, 6, 27-34 [in Polish].

27. Skrzypczyńska M., Kozioł M., Kosibowicz M. 1998. Szkodliwe owady i ich parazytoidy zasiedlajšce szyszki œwierka pospolitego Picea abies (L.) Karst. w Beskidzie Slaskim i Beskidzie Żywieckim [Pest Insects and Their Parasitoids Inhabiting Cones of Common Spruce Picea abies (L.) Karst. in Beskid Slaski Mts. And Beskid Żywiecki Mts.]. Wiad. Entomol. 17 supl. 43 Zjazd PTE – Streszczenia zgłoszonych posterów, 186 [in Polish].

28. Skrzypczyńska M., Kozioł M., Kosibowicz M. 1999. Owady uszkadzajšce szyszki i nasiona œwierka pospolitego Picea abies (L.) Karst. w wybranych drzewostanach Beskidu Œlšskiego i Żywieckiego w różnym stopniu skażonych emisjami przemysłowymi [Insects damaging cones and seeds of Norway spruce Picea abies (L.) Karst. in the selected stands of the Beskid Slaski Mts. And the Beskid Żywiecki Mts., polluted at various degree by industrial emissions]. Acta Agr. et Silv. Ser. Silv. 37, 65-78 [in Polish].

29. Skrzypczyńska M., Kożuch J. i Gibas S. 1994. Conophagous and seminifagous insects of Norway spruce Picea abies (L.) Karst. in Babia Góra National Park and its surroundings. J. Appl. Ent.118, 321-326.

30. Stadnickij G. V. 1969. Parazitičeskie chal’cidy i proktotrupidy (Hymenoptera; Chalcidoidea, Proctotrupoidea) v šiškach eli obyknovennoj [Parasitic Chalcidoidea And Proctotrupoidea in Common Spruce Cones]. Sborn. nauč.-issl. rab. po lesn. chozj. vyp. 12, 229-243 [in Russian].

31. Stadnickij G. V. 1971. Charakteristika šišek eli evropejskoj (Picea abies [L.] Karst.) kak staci obitanija i naseljajuščego ich entomokompleksa [Characterization of Common Spruce Cones (Picea abies [L.] Karst.) As a Place of Habitation and Feeding Ground]. Ent. Obozr. 50, 85-94 [in Russian].

32. Stadnitzky G. V., Kapten J. L. 1989, The spatial structure of population and measure of agregation and sampling of cone and seed insects. Proc. of the 3rd Int. Conf. IUFRO “Cone and Sees Insects”, Victoria, B.C. Canada, 167-173.

33. Szujecki A. 1980. Ekologia owadów lesnych [Ecology of forest insects]. Warszawa, PWN [in Polish].

34. Trägårdh I. 1917. Undersökingar över gran-och tallkottarnas skadeinsekter [Insect Pests of Cones And Seeds of Norway Spruce]. Medd. Skogsförsöksanst. 13-14, 1141-1204 [in Norwegian].

35. Trosset L., Roques A. 1986. Analyse de la distribution altitudinale des insectes ravageures des cones d`epicéa (Picea abies Karst.) dans les Alpes françaises du Nord [Analysis of the altitudinal distribution of the insect pests of cones of spruce (Picea abies Kast.) in the northern French Alps]. Proc. 2th Conf. IUFRO Cone and Seed Insects Working Party. S2.07-01. Brainçon (France), 83-90 [in Franch].

36. Zašev B. 1959. Vŷvoj obal’ovača šiškovego Laspeyresia (Grapholitha) strobilella L. v Bulharsku a jeho škodlivost na urode semena roku 1955 [The increasing importance of Laspeyresia (Grapholitha) strobilella L. in Bulgaria, and its damage to the 1955 seed harvest]. Lesn. čas. 5, 397–408 [in Bulgarian].

37. Witteczek K. 1998. Owady uszkadzajšce szyszki i nasiona swierka pospolitego Picea abies (L.) Karst. w wybranych drzewostanach Gorczańskiego Parku Narodowego [Insects damaging the cones and seeds of the Norway spruce Picea abies (L.) Karst. in selected wood stands in the Gorce National Park]. Parki Nar. Rez. Przyr. 17, 111-123 [in Polish].

Responses to this article, comments are invited and should be submitted within three months of the publication of the article. If accepted for publication, they will be published in the chapter headed 'Discussions' and hyperlinked to the article.