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TABLE 1. Life history traits, outbreak tendency and origin of
the laboratory cultures of pine sawfly species. G. frutetorum Interme- Single eggs
Fig. 1. Percentage of fat allocated to the egg load (mean ±
SE) in females of five pine sawfly species. Significant differ-
ences between means are indicated by different letters (P <0.05, t-tests on pairwise differences between parameter esti-
still containing all the eggs), extracted for 24 h, 48 h or 96 h, so
mates in the General Linearized Model).
we chose 48 h as a sufficient period for fat extraction.
In addition, the average egg size of each species was esti-
tetorum invested about half of their fat in their eggs and (3) the
mated by measuring the lengths and middle widths of 10
gregarious species, G. pallida, less than half of its fat (43%).
eggs/female and 5 females/species. The shape of the eggs of the
Plasticity in the allocation of fat with increase in total fat was
different species is similar (“banana”-like) and the volume (V)
tested by the regression of the total fat content (x = total
of an egg was calculated using V = S(width/2)² length.
fat/[unit BW]) on fat allocation (y = proportion of fat allocated
Evaluation of data
to eggs) in a particular species. The basic allocation pattern
remained unchanged in the gregarious species. However, in the
1. Number of eggs per female and average volume of eggs.
(semi-)solitary living D. similis and G. frutetorum the propor-
2.Total body weight (BW) [mg]: dry weight of soma + dry
tion in the egg load decreased with increasing fat content (D.
weight of egg load before fat extraction. BW was used as an
similis: y = -1.299x + 0.824, F = 10.16, (1, 23), P < 0.005; G.frutetorum: y = -0.701x + 0.679, F = 3.34, (1, 51), P = 0.072).
3. Fat [mg]: absolute amount of fat in soma or egg load. DISCUSSION
4. Fat content (Fat/[unit BW]): relative amount of fat in soma or
egg load in relation to BW. Total fat/[unit BW] is the sum of
The results of this study confirm that pine sawfly species
differ in their pattern of fat allocation to eggs and soma. How-
5. Fat allocation: Proportion or percentage of total fat allocated
ever, there is no clear relationship between the allocation pat-
to the egg load or soma, respectively.
terns and the dichotomy in sawfly oviposition. D. pini and N. sertifer are gregarious and lay their eggs in one large cluster
The statistical analysis was performed using linear regression
(Pschorn-Walcher, 1982; Blümke & Anderbrant, 1997). Sur-
and ANOVA by adopting General Linearized Models. Propor-
vival of larvae increases with colony size (Lyons, 1962),
tions were arcsine-transformed where appropriate. Means were
probably because the feeding of the early instars is enhanced or
compared by t-tests on pairwise differences between parameter
large colonies are better able to defend themselves so reducing
estimates in the General Linearized Model (Crawley, 1993).
the mortality risk per capita (Heitland & Pschorn-Walcher,1993; Codella & Raffa, 1995; Hunter, 2000). The data presented
indicate that these sawfly species invested most of their fat in
The sawfly species studied differed in several somatic and
their egg load, producing more or larger eggs. D. pini had the
reproductive traits (Table 2). The highest egg load was found in
highest number of eggs (especially in comparison to the nearly
D. pini, which was also the largest of the species (as indicated
similar sized D. similis). N. sertifer had the least, but the largest
by total BW). G. pallida and N. sertifer were significantly
eggs of all the species. In contrast to the other European saw-
smaller than the other species and had fewer eggs. However the
flies, this species overwinters in the egg and presumably has to
eggs of N. sertifer were larger than those of the other sawflies
optimize its egg size in terms of energy reserves.
(Table 2: column 5). The sawflies also varied significantly in
Distributing eggs singly or in several small clusters spatially
their total fat content, with females of G. frutetorum accumu-
can be seen as a risk-avoiding strategy, which is often associ-
lating the most fat (Table 2: column 6). The fat content (fat/unit
ated with cryptic colouration and behaviour of solitary feeding
[BW]) of the egg load was significantly different from that of
larvae (Prop, 1960). These species, in contrast to egg-clustering
the soma in the gregarious species D. pini, N. sertifer and G.
species, are likely to spend more time travelling and ovipositing,
pallida, whereas the (semi-)solitary species D. similis and G.
which are energetically costly. The results of this study lend to
frutetorum invested equal amounts of fat in eggs and soma
this contention, as the (semi-)solitary species G. frutetorum and
D. similis allocated comparatively more fat to soma than D. pini
The allocation of fat to the eggs differed between species
and N. sertifer. Moreover, with increasing fat content, both
(Fig. 1, ANOVA: F = 24.53, d.f.= (4, 207), P < 0.01). In princi-
(semi-)solitary species increased their investment of fat in soma.
ple, three species groups can be distinguished: (1) the gregarious
In addition, in the laboratory the longevity of the solitary G. fru-
species N. sertifer and D. pini allocated most of their fat (about
tetorum (10 days) was greater than that of the gregarious D. pini
60%) to their eggs, (2) the solitary species D. similis and G. fru-
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Europe-Russia in the gas market. What kind of partnership does the future hold? Anne-Grete Ellingsen, 2010 Background The oil embargo in 1973 brought the energy policy on the agenda in The European Union for the first time. In 1974 it was agreed to diversify the energy sources and the supply to improve the security of supply. New attempts to create a common energy policy were made in 1
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