distribution of eriophyoid mites (acari: eriophyoidea) on coniferous trees

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Distribution of eriophyoid mites (Acari: Eriophyoidea) on coniferous trees Mariusz Lewandowski Marcin Kozak Received: 25 October 2007 / Accepted: 5 February 2008 / Published online: 21 February 2008 Ó Springer Science+Business Media B.V. 2008 Abstract The aim of the paper was to determine the infestation parameters and species composition of eriophyoid mites for different parts of Norway spruce and Scots pine as well as for different age groups of the trees. The observations on the occurrence of the mites were conducted in 2004 and 2005 in 4 locations distributed in various regions of Poland, accounting for 11 environments (location x year). Three plant age groups were studied: (1) adult trees: 40–60 years old, additionally divided into three levels: top, middle and bottom; (2) young trees: 6–10 years old; and (3) seedlings: 2–3 years old. The same number of species (five) occurred on each coniferous tree, but only one, the rarest, was common on both tree species. Out of 500 samples for each species, mites were found on 279 pine (55.8%) and 252 spruce samples (50.2%). No tendency for the mites to choose any particular level on Scots pine and Norway spruce was observed. In addition, no tendency for the mites to choose trees from any of the age groups was observed for both Scots pine and Norway spruce, in the latter case the result obtained also for mite species subdivided into vagrant and refuge-seeking ones. Final conclusions were that in case of adult trees samples can be taken from the bottom part of a tree; however, sampling from young trees growing among adult trees may be seen as the most efficient sampling method. Keywords Eriophyoidea Pinaceae Spruce Pine Vertical distribution Introduction Phytophagous arthropods can specialize not only with respect to plant taxa they feed upon but also to the particular part of plants. This is especially encountered in groups where size M. Lewandowski (&) Department of Applied Entomology, Faculty of Horticulture and Landscape Architecture, Warsaw University of Life Sciences—SGGW, Nowoursynowska 159, 02-776 Warsaw, Poland e-mail: [email protected] M. Kozak Department of Biometry and Bioinformatics, Faculty of Agriculture and Biology, Warsaw University of Life Sciences—SGGW, Nowoursynowska 159, 02-776 Warsaw, Poland 123 Exp Appl Acarol (2008) 44:89–99 DOI 10.1007/s10493-008-9135-0

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Page 1: Distribution of eriophyoid mites (Acari: Eriophyoidea) on coniferous trees

Distribution of eriophyoid mites (Acari: Eriophyoidea)on coniferous trees

Mariusz Lewandowski Æ Marcin Kozak

Received: 25 October 2007 / Accepted: 5 February 2008 / Published online: 21 February 2008� Springer Science+Business Media B.V. 2008

Abstract The aim of the paper was to determine the infestation parameters and species

composition of eriophyoid mites for different parts of Norway spruce and Scots pine as

well as for different age groups of the trees. The observations on the occurrence of the

mites were conducted in 2004 and 2005 in 4 locations distributed in various regions of

Poland, accounting for 11 environments (location x year). Three plant age groups were

studied: (1) adult trees: 40–60 years old, additionally divided into three levels: top, middle

and bottom; (2) young trees: 6–10 years old; and (3) seedlings: 2–3 years old. The same

number of species (five) occurred on each coniferous tree, but only one, the rarest, was

common on both tree species. Out of 500 samples for each species, mites were found on

279 pine (55.8%) and 252 spruce samples (50.2%). No tendency for the mites to choose

any particular level on Scots pine and Norway spruce was observed. In addition, no

tendency for the mites to choose trees from any of the age groups was observed for both

Scots pine and Norway spruce, in the latter case the result obtained also for mite species

subdivided into vagrant and refuge-seeking ones. Final conclusions were that in case of

adult trees samples can be taken from the bottom part of a tree; however, sampling from

young trees growing among adult trees may be seen as the most efficient sampling method.

Keywords Eriophyoidea � Pinaceae � Spruce � Pine � Vertical distribution

Introduction

Phytophagous arthropods can specialize not only with respect to plant taxa they feed upon

but also to the particular part of plants. This is especially encountered in groups where size

M. Lewandowski (&)Department of Applied Entomology, Faculty of Horticulture and Landscape Architecture,Warsaw University of Life Sciences—SGGW, Nowoursynowska 159, 02-776 Warsaw, Polande-mail: [email protected]

M. KozakDepartment of Biometry and Bioinformatics, Faculty of Agriculture and Biology, Warsaw Universityof Life Sciences—SGGW, Nowoursynowska 159, 02-776 Warsaw, Poland

123

Exp Appl Acarol (2008) 44:89–99DOI 10.1007/s10493-008-9135-0

Page 2: Distribution of eriophyoid mites (Acari: Eriophyoidea) on coniferous trees

and mobility is limited, such as mites (Krantz and Lindquist 1979). One of the major

groups of phytophagous mites is the superfamily Eriophyoidea, which includes many

species that can cause a direct damage to their host plants and lead to the production of a

variety of galls or other abnormalities and distortions, such as erinea, leaf deformation,

discoloration and russeting (Westphal and Manson 1996). Some eriophyoid mites may

transmit viruses (Oldfield and Proeseler 1996).

Eriophyoid mites are characterized by considerable host specificity. Their active

movement is rather low; within a host they disperse by crawling whereas long distance

movements between plants are possible passively by wind currents or phoresy (Sabelis and

Bruin 1996).

Limited migration capacity of these mites can lead to their uneven distribution within

different plant parts, especially on large perennials, such as coniferous trees. On the other

hand, owing to their perennial nature, coniferous trees can create stable microhabitats for

these small arthropods (Boczek and Shevchenko 1996), facilitating a progressive infes-

tation of the entire tree. Information on eriophyoids’ distribution on coniferous plants and

their possible preference to the age or part of a plant is very limited. Sparse information

available show that eriophyoid mites prefer young tissues of these plants (Boczek and

Shevchenko 1996), and suggests the infestation only of the external shoots of a tree crown.

However, little is known about the vertical distribution of eriophyoids within tree crowns.

In the case of annual plants a tendency for eriophyoid mites to inhabit top plant parts has

been shown. Such preference can be attributed to young tissues, present in top parts of

plants and constituting the best food source (Gibson 1974), as well as to better conditions

for starting the migration (Gibson and Painter 1957; Nault and Styer 1969). In the case of

grass-inhabiting Abacarus hystrix (Nalepa) a greater number of specimens on top leaves

were observed in spring and early summer than on low leaves (Nault and Styer 1969;

Skoracka et al. 2003). A study concerning the distribution of two species of eriophyoid

mites on Cupressus sempervirens L. showed preferences to the top level of plants crown

only in the case of bud species; free living mites were distributed evenly (Castagnoli and

Simoni 2000). The preferences of vagrant eriophyoids to the upper part of plant were

observed on tea shrubs, on which mostly top and middle parts of the plant were infested

(Muraleedharan et al. 1988). Distribution of eriophyoid mite galls on tree crown levels

have been studied in alder (Vuorisalo et al. 1989) and green ash (Wawrzynski and Ascerno

1989), but the results of their studies were opposite, the former showing the mite galls

preference for lower while the latter for upper parts of tree crown. On the other hand, gall-

producing mites Aculus tetanothrix (Nalepa), inhabiting willows, produced smaller number

of galls on the outer than inner leaves of the shoots (Kuczynski and Skoracka 2005).

Simmilar trend was observed on gapevine infested by Calepitrimerus vitis (Nalepa) (de

Lillo et al. 2005).

Presently about 170 eriophyoid species associated with coniferous plants are known,

more than 100 of which have been found on Pinaceae (de Lillo and Amrine 2005,

unpublished data). These eriophyoid mites inhabit various plant parts, from needles and

shoots through buds, inflorescences, fruit, and scales at the base of shoots up to shoot galls.

Based on the literature it is difficult to conclude on the distribution of eriophyoid mites on

coniferous plants, which offer the habitat completely different from those offered by

deciduous trees and shrubs.

A wide diversity of eriophyoids, their different life history as well as a large size of

coniferous trees make the determination of an efficient sampling method difficult. The aim

of the present study was to determine the infestation parameters and species composition of

eriophyoid mites in relation to different (a) parts of plants and (b) age groups of Scotch

90 Exp Appl Acarol (2008) 44:89–99

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pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) H. Karst.). These coniferous

species were chosen for the study since they are commonly used both in forestry and as

ornamental trees in gardens in Poland.

Materials and methods

Observations on the occurrence of eriophyoid mites were conducted from June to Sep-

tember in 2004 and 2005 on Polish native conifers, Norway spruce and Scots pine. The

study was based on the material collected from the following locations distributed in

different regions of the country: shoots of Norway spruce were taken from Koszarawa

Bystra near _Zywiec (Slaskie Province, 49�400 N, 19�240 E) and Lutocin near Sierpc

(Mazowieckie Province, 52�590 N, 19�460 E), whereas shoots of Scots pine from Kuszewo

near Czaplinek (Zachodniopomorskie Province, 53�370 N, 16�170 E), Lutocin near Sierpc

(Mazowieckie Province, 52�590 N, 19�460 E), and Urwitałt near Mikołajki (Warminsko-

Mazurskie Province, 53�480 N, 21�380 E); see Table 1 for sampling dates. Due to workload

connected with the sampling we were not able to collect the samples from the locations at

the same time, for which reason in both years the samples were collected during summer in

order to minimize the seasonal effect on the number of mites. For each plant species 10

stands were examined.

Three plant age groups were studied: (1) adult trees (usually about 40–60 years old); (2)

young trees (usually about 6–10 years old); and (3) seedlings (usually about 2–3 years

old). Additionally, crowns of adult trees were visually divided into three levels: top, middle

and bottom. Ten samples were taken from each level of an adult tree and from a whole

young tree. Stands for sampling were selected in the middle of a forest consisting of

randomly selected trees of all these age groups. Each stand was represented by 1 adult tree,

1 young tree and 10 seedlings. A single sample consisted of a randomly collected 15-cm

shoot. For seedlings, an entire plant comprised a sample.

Mites that occurred on needles, twigs, buds and under scales at the base of a current

year’s shoots were counted and collected. For pine additionally 10 current-year sheaths of

needles were collected. Each sample was examined under the stereo-microscope. Mites

were mounted in modified Berlese medium (Amrine and Manson 1996) and identified to

species.

To show differences in the lifestyle of mites in relation to the host plant, the following

definitions were adapted after Sabelis and Bruin (1996): vagrant - species living freely on a

plant surface, and refuge-seeking - species occurring in shelters on a plant, e.g. buds and

under sheaths.

Statistical analysis

The following parameters were used to describe a degree of eriophyoid mites’ infestation

(Bush et al. 1997):

(1) Prevalence, expressed in percentage values, is interpreted as the probability of finding

a mite on a randomly selected sample. Prevalence for trees (Pt)—the number of trees

infested relative to the number of trees examined. Prevalence for samples (Ps)—the

number of samples infested relative to the number of samples examined.

Exp Appl Acarol (2008) 44:89–99 91

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(2) Intensity (I)—the number of mites relative to the number of samples infested.

Expressed as number of individuals (mites) per unit space (sample), it is interpreted

as the population density in occupied habitat patches.

(3) Density (D)—the number of mites relative to the number of samples checked. This

measure has the same dimension as intensity, but measures the population density

across all (theoretically) available habitat patches. Density can be considered as the

product (or interaction) of prevalence and intensity.

Confidence intervals for prevalence were calculated using the formula for an estimator of

probability of success in a binomial distribution, the distribution that this measure follows.

Confidence intervals for intensity and density were computed using the bootstrap-T method (Efron

and Tibshirani 1993) using bootstrap package of R language (R Development Core Team 2006).

To study the vertical distribution of eriophyoid mites on crown levels of adult trees of a

particular tree species, the following transformation was applied:

Zij ¼ Xij

X3

i¼1

Xij

!�1

;

Table 1 Occurrence of eriophyoid species on pinaceous trees in the localities of study

Eriophyoid species Microhabitats Locality, date of sampling

Picea abies

Calepitrimerus lutocinusLewandowski, 2006

Vagrant on needles and twigs Lutocin, 25.09.2004; 30.07.2005;27.08.2005; 23.09.2005

Nalepella shevtchenkoiBoczek, 1969

Vagrant on needles and twigs Koszarawa, 25.08.2004

Lutocin, 25.09.2004; 23.09.2005

Phyllocoptes farkasiBoczek, 1969

Vagrant on needles Lutocin, 23.09.2005

Phyllocoptes piceaeSoika, 1999

Vagrant on needles and twigs Koszarawa, 25.08.2004; 29.06.2005;01.07.2005; 02.07.2005

Lutocin, 25.09.2004; 30.07.2005;27.08.2005; 23.09.2005

Trisetacus relocatesBagnyuk andShevtchenko, 1982

Under scales at the baseof the current year’s shoots

Koszarawa, 25.08.2004; 29.06.2005;01.07.2005; 02.07.2005

Lutocin, 25.09.2004; 30.07.2005

Pinus sylvestris

Phyllocoptes farkasiBoczek, 1969

Vagrant on needles Kuszewo, 08.08.2005

Platyphytoptus sabinianaeKeifer, 1938

Sheaths of needles Urwitałt, 04.08.2004; 22.08.2005

Lutocin, 25.09.2004

Kuszewo, 08.08.2005

Setoptus multigranulatusCastagnoli, 1973

Sheaths of needles Lutocin, 25.09.2004

Urwitałt, 22.08.2005

Setoptus piniBoczek, 1964

Sheaths of needles Urwitałt, 04.08.2004; 22.08.2005

Lutocin, 25.09.2004

Kuszewo, 08.08.2005

Trisetacus silvestrisCastagnoli, 1973

Sheaths of needles Urwitałt, 04.08.2004; 22.08.2005

Lutocin, 25.09.2004

Kuszewo, 08.08.2005

92 Exp Appl Acarol (2008) 44:89–99

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where Xij is the number of mites observed on the ith level (i = 1 [botton], 2 [middle], 3

[top]) on 10 samples from the jth tree, and Zij is the fraction of Xij in the number of all

mites observed on 30 samples from the jth tree. The Zij’s for a particular tree show the

vertical distribution of the mites on this particular plant species. The Zij’s being similar for

the three levels and this distribution being similar for all trees studied, no tendency of the

mites to choose any of the levels would be detected. Note that observations of number of

mites on the three levels from a particular tree are correlated. Therefore, as the Zs followed

the analysis of variance (ANOVA) assumptions (results not presented), we applied

ANOVA for paired data. To perform the analysis, growth package (Lindsey 2007) of R

language (R Development Core Team 2006) was applied.

To compare adult and young trees subject to the mites’ occurrence, mean number of

mites in a sample from an adult and a young tree was calculated; for an adult tree, the mean

was calculated based on 30 samples whereas for a young tree, on ten samples. For Scots

pine, both variables (mean occurrence of mites in a sample from a tree for adult [first

variable] and young trees [second variable]) followed the normal distribution, as revealed

by Shapiro-Wilk test (Shapiro and Wilk 1965) as well as tests for skewness and kurtosis.

However, because their variances were not equal (as the means were calculated based on a

different number of samples: 30 for adult and 10 for young trees), we applied a t-test with

the Welch modification to the degrees of freedom, available in R language. For both

vagrant and refuge-seeking species observed on Norway spruce, the variables were not

normally distributed according to Shapiro-Wilk test. As the box-cox transformation (Quinn

and Keough 2002, sect. 4.3.1) failed to normalize the distributions of the variables, we

applied the Wilcoxon rank sum test with continuity correction to compare the medians. As

each test based on ranks, it assumes equal variances of the variables compared. None-

theless, with the main difference in the variances due to the trees with the largest values,

the Wilcoxon test should work properly for our data.

For all the analyses, R language (R Development Core Team 2006) was used, and the

significance level chosen was a = 0.05.

Results

Five species of eriophyoid mites were found on both Scots pine and Norway spruce

(Table 1). A total of 2263 (range 0–124, mean 4.5 per 10 sheaths of needles, SD 10.5)

specimens of mites were found on pine and 6926 (range 0–472, mean 13.9 per sample, SD

38.8) on spruce. Out of 500 samples for each tree species, mites were found on 279 pine

(55.8%) and 252 spruce samples (50.2%).

Since single specimens of Phyllocoptes farkasi Boczek were found on both plant spe-

cies, they were not taken into the analysis. In the case of remaining mite species the evident

predomination of mites of the genus Trisetacus Keifer was observed. On spruce Trisetacusrelocatus Bagnyuk and Shevtchenko (70.5%) outnumbered the second-abundant Phyllo-coptes piceae Soika (26.5%) while on pine Trisetacus silvestris Castagnoli (77.7%)

dominated over the second-abundant Platyphytoptus sabinianae Keifer (18.4%).

For Scots pine ANOVA for paired data did not reveal any significant differences

(P = 0.61) in the vertical distribution of mites on the trees. All mites were refuge-seeking

species. A similar result was obtained for the vagrant (P = 0.34) and refuge-seeking mite

species (P = 0.13) for Norway spruce.

The t-test showed that there was no significant difference (P = 0.50) between the mean

occurrence of mites on samples from the adult and young Scots pine trees. The same result

Exp Appl Acarol (2008) 44:89–99 93

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was obtained using the Wilcoxon test for both vagrant (P = 0.85) and refuge-seeking mite

species (P = 0.60) for Norway spruce.

Levels of tree crowns did not differ in infestation parameters for both spruce and pine

(Tables 2 and 4). No differences were detected also for age groups (Tables 3 and 4). Only

for spruce seedlings confidence intervals showed the lower values of prevalence and

density for the eriophyoid mites inhabiting scales at the base of shoots than for young and

adult trees (Table 3) and in the case of pine age groups slightly lower values of prevalence

were observed for seedlings (Table 4).

No significant differences among the crown levels of spruce adult trees in infestation

parameters were found for T. relocatus as well as P. piceae. For Calepitrimerus lutocinusLewandowski single specimens were found only in bottom parts of trees. Nalepellashevtchenkoi Boczek specimens were found in top and bottom parts of trees, also in a small

number (Table 5).

For spruce the infestation parameters of two dominant species, T. relocatus and P.piceae, did not significantly differ also in particular tree age groups. Single specimens of C.lutocinus and N. shevtchenkoi were found on spruce, except for the former on young trees,

where it occurred quite frequently. In addition, N. shevtchenkoi was not observed on

seedlings (Table 5).

Table 2 Infestation parameters and their confidence intervals (CI) for eriophyoid mites found in differentlevels of spruce trees

Pt 95% CI Ps 95% CI I 95% CI D 95% CI

Vagrant mites

Top 90.0 56.0–100 42.0 24.8–66.9 9.7 5.5–26.9 6.6 3.2–32.4

Middle 60.0 26.0–88.0 39.0 15.2–61.7 4.8 2.6–7.5 2.3 1.1–5.1

Bottom 70.0 35.0–93.0 42.0 20.3–68.9 9.9 4.4–19.9 5.8 2.1–11.3

Mites living under scales at the base of the current year’s shoots

Top 30.0 7.0–65.0 13.0 2.1–114.2 15.3 3.0–129.2 3.9 0.0–323.0

Middle 40.0 12.0–74.0 21.0 9.5–52.2 46.5 15.7–168.7 14.9 3.5–106.8

Bottom 40.0 12.0–74.0 25.0 6.2–74.6 70.9 34.4–238.5 22.5 5.0–322.3

Pt, Prevalence for trees [%]; Ps, Prevalence for samples [%]; I, Intensity; D, Density

Table 3 Infestation parameters and their confidence intervals (CI) for eriophyoid mites found in particularage groups of spruce trees

Pt 95% CI Ps 95% CI I 95% CI D 95% CI

Vagrant mites

Adult tree 90.0 56.0–100 41.0 25.7–66.0 7.5 4.1–15.5 4.9 2.4–14.3

Young tree 90.0 56.0–100 45.0 31.7–64.6 7.7 4.5–18.3 5.2 2.8–14.9

Seedlings 20.0 13.0–29.0 20.0 13.0–29.0 6.2 4.6–8.7 1.2 0.6–2.4

Mites living under scales at the base of the current year’s shoots

Adult tree 40.0 12.0–74.0 19.7 6.8–56.7 54.0 19.8–186.0 13.7 3.0–125.2

Young tree 60.0 26.0–88.0 22.0 7.3–44.3 33.7 15.1–160.2 6.1 2.1–13.5

Seedlings 11.0 6.0–19.0 11.0 6.0–19.0 8.0 5.0–12.5 0.9 0.4–0.9

Pt, Prevalence for trees [%]; Ps, Prevalence for samples [%]; I, Intensity; D, Density

94 Exp Appl Acarol (2008) 44:89–99

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No significant differences in infestation parameters of eriophyoid species studied in

relation to different height levels and among different age groups of pine were observed.

Taking into account the estimated confidence intervals, significant differences for preva-

lence and density were observed only for the most numerous species, T. silvestris, and for

prevalence of Setoptus species between adult trees and seedlings (Table 6).

Table 4 Infestation parameters and their confidence intervals (CI) for eriophyoid mites found on particularlevels of adult trees and age groups of pine trees

Pt 95% CI Ps 95% CI I 95% CI D 95% CI

Vertical distribution

Top 100 – 58.0 46.0–69.0 7.0 4.4–11.6 5.3 2.9–9.5

Middle 100 – 54.0 39.0–71.0 5.4 4.5–7.1 4.8 2.4–5.8

Bottom 100 – 71.0 60.8–79.8 5.6 4.6–9.5 4.3 3.4–6.1

Age groups

Adult tree 100 – 64.0 50.4–75.4 6.2 4.8–8.0 4.5 3.2–7.1

Young tree 80.0 44.0–97.0 53.0 33.0–67.8 9.9 5.6–13.6 6.2 3.3–11.4

Seedlings 34.0 25.0–44.0 34.0 25.0–44.0 8.7 5.8–14.6 3.0 1.9–5.0

Pt, Prevalence for trees [%]; Ps, Prevalence for samples [%]; I, Intensity; D, Density

Table 5 Infestation parameters and their 95% confidence intervals for eriophyoid species infested sprucetrees

Species of eriophyoid mites

T. relocatus P. piceae C. lutocinus N. shevtchenkoi

Vertical distribution of mites on adult trees

Pt Top 40.0 12.0–74.0 90.0 56.0–100 0.0 – 10.0 0.0–45.0

Middle 40.0 12.0–74.0 50.0 19.0–81.0 0.0 – 0.0 –

Bottom 40.0 12.0–74.0 60.0 26.0–88.0 20.0 3.0–56.0 30.0 7.0–65.0

I Top 21.7 1.1–100.1 11.5 5.1–26.9 0.0 – 1.0 –

Middle 42.5 7.1–183.2 5.3 1.9–10.1 0.0 – 0.0 –

Bottom 70.1 25.4–267.0 9.8 2.4–41.0 1.0 – 2.3 0.2–44.1

D Top 7.9 0.0–171.0 6.5 1.3–54.6 0.0 – 0.02 –

Middle 13.6 1.8–108.1 2.3 0.7–5.7 0.0 – 0.0 –

Bottom 22.5 7.7–269.5 5.4 1.1–29.3 0.02 – 0.3 0–2.9

Distribution of mites on different age groups of trees

Pt Adult tree 40.0 12.0–74.0 100 – 20.0 3.0–56.0 30.0 7.0–65.0

Young tree 60.0 26.0–88.0 80.0 44.0–97.0 50.0 19.0–81.0 50.0 19.0–81.0

Seedlings 11.0 6.0–19.0 20.0 13.0–29.0 4.0 1.0–10.0 0.0 –

I Adult tree 54.4 5.3–297.5 11.3 5.4–20.2 1.0 – 2.3 0.6–67.5

Young tree 27.4 8.4–121.9 5.4 1.9–24.5 6.1 3.4–15.3 1.6 1.0–7.4

Seedlings 8.0 5.4–11.8 5.8 4.0–8.6 1.8 1.0–3.6 0.0 –

D Adult tree 14.7 2.3–174.9 4.7 1.5–22.6 0.01 – 0.1 0.0–1.1

Young tree 4.9 1.4–13.1 3.5 0–31.7 1.4 0.1–8.4 0.3 0.1–0.6

Seedlings 0.9 0.5–1.7 1.2 0.8–1.8 0.1 0.0–0.2 0.0 –

Pt, Prevalence for trees [%]; I, Intensity; D, Density

Exp Appl Acarol (2008) 44:89–99 95

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Discussion

In the study on eriophyoid mites’ distribution on spruce and pine, two most common

coniferous trees in Poland, ten eriophyoid species were found, five for each plant species.

Three of them, Setoptus multigranulatus Castagnoli, T. relocatus and T. silvestris were new

for Poland. Phyllocoptes farkasi has so far been found on three pine species: P. sylvestis,

Pinus nigra J. F. Arnold and Pinus jeffreyi Grev. and Balf. (Boczek 1969; Soika and

Łabanowski 1999). The presence of this eriophyoid on Norway spruce should not be

thought of as a proof for this plant species to be a host for this mite, because this infestation

might have been accidental (with just single specimens having been found). For Norway

spruce T. relocatus was a predominant species. Its significant share in the species com-

position of the eriophyoid mites inhabiting spruce resulted from the fact that, as the species

living in refugia (under the buds’ scales, at the base of the current-year shoots), it

developed more numerous colonies than the vagrant species did (Bagnyuk and Shevchenko

1982). This was also confirmed by the values of infestation parameters. Prevalence seemed

lower for T. relocatus than for P. piceae, the latter being the second abundant species, but

intensity seemed higher for the former. These differences, however, were not shown by the

statistical analysis owing to a wide range of the mite counts observed, which resulted from

numerically heterogeneous colonies.

Table 6 Infestation parameters and their 95% confidence intervals for eriophyoid species that infested pinetrees

Species of eriophyoid mites

T. silvestris P. sabinianae Setoptus spp.

Vertical distribution of mites on adult trees

Pt Top 90.0 56.0–100 40.0 12.0–74.0 50.0 19.0–81.0

Middle 100 – 40.0 12.0–74.0 20.0 3.0–56.0

Bottom 100 – 20.0 3.0–56.0 30.0 7.0–65.0

I Top 6.7 3.8–11.2 3.0 1.2–11.5 1.9 1.3–6.0

Middle 5.2 4.3–6.9 2.6 0.7–5.4 1.5 0.9–9.3

Bottom 5.8 4.5–7.1 1.8 1.2–4.8 1.8 0.9–10.3

D Top 4.5 2.1–8.8 0.5 0.1–3.1 0.3 0.1–1.8

Middle 3.6 1.7–7.2 0.1 0.06–0.4 0.04 –

Bottom 4.2 2.9–5.8 0.1 0.01–0.8 0.1 0–0.3

Distribution of mites on different age groups of trees

Pt Adult tree 100 – 50.0 19.0–81.0 60.0 26.0–88.0

Young tree 80.0 44.0–97.0 60.0 26.0–88.0 30.0 7.0–65.0

Seedlings 25.0 17.0–35.0 20.0 13.0–29.0 4.0 1.0–10.0

I Adult tree 5.9 4.5–7.8 2.7 1.7–6.7 2.1 1.2–4.1

Young tree 7.2 3.3–34 7.2 2.2–61.0 1.7 0.0–2.6

Seedlings 4.6 3.4–6.4 9.6 5.4–20.1 9.0 0.5–24.3

D Adult tree 4.1 2.6–6.2 0.2 0.1–0.9 0.1 0.0–0.6

Young tree 4.4 2.2–19.6 1.6 0.1–17.3 0.1 0.0–0.2

Seedlings 1.1 0.8–1.7 1.9 0.9–4.8 0.4 0.0–9.9

Pt, Prevalence for trees [%]; I, Intensity; D, Density

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Apart from P. farkasi, found only on few pine trees, all eriophyoid species were found

on plants in each age group studied. For both pine and spruce, only N. shevtchenkoi,inhabiting spruce, was not found on seedlings; moreover, the values of density parameters

of particular mite species did not significantly differ for adult and young trees of both

species. Based on these results one may presume that the development of eriophyoids

begins on seedlings or young trees and their population increases with the tree’s devel-

opment. This probably explains almost the same species composition of eriophyoids for all

the age groups and plant species studied as well as similar mite density on young and adult

trees. For this reason it seems likely that the inhabiting of new trees by these mite species

consists of falling from adult trees on seedling and young trees growing close around. This

movement strategy for these eriophyoids may be as important for short distance move-

ments as passive dispersal by wind currents or phoresy for long distance movements.

The lack of differences in eriophyoids’ vertical distribution on particular parts of adult

trees could result from the number of the samples taken and the low density of eriophyoids

that was observed. Perhaps increasing the number of samples would reveal some prefer-

ences of these mites to specific parts or plant age; on the other hand, such high P-values

obtained for different hypotheses indicate a strong credibility of the here-formed conclu-

sions. Moreover, because of the labor- and time-consuming methods of collecting the

samples and analyzing the material, increasing the number of samples was impossible.

Vertical distribution of eriophyoid mites on coniferous plants has been studied only on

C. sempervirens. Observations on two eriophyoid species inhabiting this plant showed that

Trisetacus juniperinus (Nalepa) preferred buds in the top zones of tree crown, while

Epitrimerus cupressi (Keifer), a vagrant species, did not show this tendency (Castagnoli

and Simoni 2000). This may suggest that the preference of an eriophyoid mite species to a

plant crown level may be strictly related to the growth strategy of this species. This was

confirmed by other studies on the distribution of galls produced by eriophyoids on

deciduous trees. For Eriophyes fraxiniflora Felt the number of galls was the greatest on the

top level of tree crown (Wawrzynski and Ascerno 1989); the opposite tendency, however,

was observed in the study on distribution of galls produced by Eriophyes laevis (Nalepa)

on leaves of alder trees (Vuorisalo et al. 1989).

Uneven distribution of eriophyoids within a plant can be also related to food quality and

dispersion phenomenon. Eriophyoid mites prefer young plant tissues (Boczek and Shev-

chenko 1996); on herbaceous plants they are more numerous on top than low parts (Gibson

1974), which additionally facilitates their dispersion (Gibson and Painter 1957; Nault and

Styer 1969). In addition, the results of studies concerning the vertical distribution of galls-

producing eriophyoids (Wawrzynski and Ascerno 1989; Vuorisalo et al. 1989; Kuczynski

and Skoracka 2005) as well as the vagrant species inhabiting crop trees and shrubs

(Muraleedharan and Ambalatharsu 1983; Muraleedharan et al. 1988; 1994; de Lillo et al.

2005) might suggest that also on spruce and pine some preferences to infest a particular

part of a plant could be expected. For Camellia sinensis (L.) O. Kuntz a higher number of

eriophyoid mites were observed on the top and middle than on low parts of bushes;

however, each of the eriophyoid species preferred a different plant part (Muraleedharan

et al. 1988). The lack of preference for inhabiting specific tree parts may in the case of

coniferous plants result from the fact that these trees offer the eriophyoids stable and long-

lasting conditions for living and reproducing, which makes the mites not need to migrate to

seek winter shelters (Boczek and Shevtchenko 1996). Moreover, young tissues appearing

each year on terminal parts of shoots make it unnecessary for the eriophyoid mites to move

towards the top plant parts.

Exp Appl Acarol (2008) 44:89–99 97

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From our results it follows that in studies to eriophyoid mites on pine and spruce,

samples can be taken from the bottom part of a tree. Because of the lack of differences in

the eriophyoids’ densities among the parts of plants investigated, revealed in the present

study, the adapted sampling method may be considered reliable in determining both

species composition and density of eriophyoids. Nonetheless, with no significant differ-

ences in density and species composition found between young and adult trees and with

sampling from young trees less labor-consuming than from adult trees, sampling from

young trees that grow among adult trees may be seen the most efficient sampling method.

Acknowledgments We wish to thank Prof. Jan Boczek (WULS-SGGW, Warsaw, Poland), Prof. Kro-pczynska-Linkiewicz (WULS-SGGW, Warsaw, Poland) and dr Anna Skoracka (Adam MickiewiczUniversity, Poznan, Poland) for their valuable suggestions on the manuscript. The study was financiallysupported by the Warsaw University of Life Sciences - SGGW, grant no. 50404120011. We would also liketo thank the two anonymous reviewers for their detailed and helpful comments on the first version of thispaper.

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