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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- 7$%/(  $YHUDJH VRPDWLF DQG UHSURGXFWLYH WUDLWV RI QHZO\ HPHUJHG IHPDOHV RI ILYH SLQH VDZIO\ VSHFLHV 9DOXHV DUH PHDQV “ 6' Q QXPEHU RI IHPDOHV H[DPLQHG SHU VSHFLHV >X %:@ XQLW ERG\ ZHLJKW  0HDQV IROORZHG E\ GLIIHUHQW OHWWHUV DUH VLJQLILFDQWO\ GLI IHUHQW 3   HVWLPDWHG E\ WWHVWV RQ SDLUZLVH GLIIHUHQFHV EHWZHHQ SDUDPHWHU HVWLPDWHV LQ WKH *HQHUDO /LQHDUL]HG 0RGHO   6LJ QLILFDQW GLIIHUHQFHV EHWZHHQ WKH IDW FRQWHQW RI VRPD DQG HJJ ORDG IRU D SDUWLFXODU VDZIO\ VSHFLHV DW 3   DUH LQGLFDWHG E\ 6SHFLHV >PJ@
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