Heredity of
elytral
colour in
of Poecilus
lepidus
Leske (Coleoptera, Carabidae) 185
L.the
Penev,
T. Erwin
&adults
T. Assmann
(eds.)
2008.
Back to the Roots and Back to the Future? Towards a New Synthesis between
Taxonomic, Ecological and Biogeographical Approaches in Carabidology
Proceedings of the XIII European Carabidologists Meeting, Blagoevgrad, August 20-24, 2007, pp. 1-4.
© Pensoft Publishers
Sofia–Moscow
Heredity of the elytral colour in adults of
Poecilus lepidus Leske (Coleoptera, Carabidae)
Wilfried Paarmann1, Wolfgang Rohe1, Ingrid Lüchtrath1,
Thorsten Assmann2 & Dietrich Mossakowski3
1
HAWK, Fakultät Ressourcenmanagement, Büsgenweg 1 A, D 37077 Göttingen.
E-mail: [email protected]
2
Institute of Ecology and Environmental Chemistry, Leuphana University Lüneburg,
Scharnhorststr. 1, D 21332 Lüneburg. E-mail: [email protected]
3
Institute for Ecology & Evolutionary Biology, University of Bremen,
P. O. Box 330440, D 28334 Bremen
ABSTRACT
The heredity of altogether seven colour morphs of the ground beetle species Poecilus lepidus
was studied by crossbreeding experiments: black, dark green and blue morphs found in
Italy (Lago Maggiore, Apennine), and red, yellowish green, bluish green and violet from
a population in Germany (Lüneburger Heide). In their natural populations the colour
morphs black, red, bluish green and blue are homozygous: ss (black), rr (red), gg (bluish
green), bb (blue). Other green colour morphs are heterozygous: dark green (Lago Maggiore
population) bs, yellowish green (Lüneburger Heide) gr. Crossbreeding experiments between beetles of the ‘German’ and the ‘Italien’ strains showed the red allele (R) is dominant
over black (s), and bluish green (G) dominant over blue (b). Heterozygous beetles with
the co-dominant alleles gs and br have a green colour. The dominant allele V (violet) was
only found in the ‘Lüneburger Heide’ population, where it seems to be very rare.
Violet colour morphs in the ‘Lüneburger Heide’ population should mainly have the
allele combination Vr, because the red colour morph is dominant there (82%). Beetles with
combination Vg are also violet, but the bluish green colour morph (gg) is comparable rare
(3%). Beetles homozygous for the allele V should be extremely rare in the ‘Lüneburger
Heide’ population. Only in the crossbreeding experiments homozygous females were
achieved. They were almost black with a small violet brim of the elytra.
The ecological significance of colour forms in Peocilus lepidus may be interpreted
partly as mimetic effects.
186 W. Paarmann et al.
Key words: Poecilus lepidus, colour forms, heredity.
1. INTRODUCTION
Poecilus lepidus has a Palaearctic distribution ranging from the Pyrenees to the Amur
region in Siberia and from Trondheim region in Norway to central Italy (Lindroth,
1949, Turin, 2000).
A number of colour morphs from P. lepidus of different regions are known: blue, green,
and copper-coloured (Erichson et al., 1860, Lindroth, 1949, Turin, 2000; for an overview of
the colour morphs of P. lepidus s.s tr, and P. lepidus gressorius see Schatzmayr, 1942-43). Erichson et al. (1860) and Turin (2000) additionally mention black morphs. From Kirschenhofer
(personal communication 1984) we have the information that in the Pyrenees uniformly
black populations exist (in the Bigorre-massive, Assmann found exclusively black Poecilus
lepidus, too). The Italian populations in the Apennine are uniformly blue (cf. Schatzmayr,
1942-43). This subspecies, P.l. gressorius, has females with shining elytra while the elytra of
the subspecies lepidus is dull. Kirschenhofer mentions a violet specimen in his collection.
Violet specimens were also recorded by Horion (1941). In Western Siberia also black, green
and cupper coloured beetles appear (Zinovyev, personal communication 2007).
P. lepidus is a species that can easily bred under laboratory conditions (Kegel, 1989,
Paarmann, 1990, De Vries, 2000). Laboratory studies on the environmental control of the life
cycle of P. lepidus (Paarmann, 1990) led to the idea to study the heredity of colour morphs.
Up to now only very few studies on the potential value of these different colour morphs
for an adaptation to different habitat conditions are already done. Mossakowski (1980) stated
differences in reduction of solar radiation absorbance and heat gain for green versus red
morphs of Cicindela campestris. Schultz (1986) demonstrated the role of structural colours in
predator avoidance by tiger beetles of the genus Cicindela. Van Natto & Freitag (1986) got
differences in the amount of reflectance in a comparison of carabids with structural colours
and black pigmentation. Terrell-Nield (1990) found a positive relationship between ground
temperature and percentage of black colour morphs of the legs in Pterostichus madidus.
For most of these studies the heredity of the colours is a precondition for further purpose (e.g. to determine selection coefficients). Poecilus lepidus is a suitable model organism
because of its different habitats (see discussion), many colour morphs and the possibility
to rear it under laboratory conditions, which is essential for crossbreeding experiments.
2. MATERIAL AND METHODS
For our crossbreeding experiments we used three strains of different origin:
1. North Italy - Lago Maggiore (fig. 1, LM). This strain was founded in 1979 with
only three specimens of the colour morphs black, blue and green each. It is still
in culture in the 34th generation now.
Heredity of the elytral colour in adults of Poecilus lepidus Leske (Coleoptera, Carabidae) 187
2. Central Italy - Apennine (fig. 1, Plg). Only one pregnant blue female founded
this strain.
3. Germany - Lüneburger Heide (fig. 1, LH). Founded by many specimens of the
colour morphs red (copper coloured), yellowish green, bluish green and violet.
Some of the data were collected during the studies dealing with the life cycle control.
For breeding conditions see Paarmann (1990). During the following studies we simulated
winter conditions by keeping the adults in 8 °C and short day conditions (8h light) or
in continuous darkness at 4 ºC. To reach maturation, we transferred them into 18 or 20
°C and long day conditions (16h light). Larvae were either collected directly from the
culture vessel or the females were separated on fine moisten sand. After a few days the
sand was washed through a sieve to extract the eggs. The eggs were kept on moisten sand,
separated by pieces of plastic drinking pipes, pressed into the sand. After hatching, the
larvae were transferred into glass tubes of a height of 7.5 cm and a diameter of 2.5 cm.
They were filled with moist peat moss up to a height of 5 cm. The larvae were fed with
pieces of mealworms during their whole development.
The determination of the colour morph was done with beetles older than one month
(hardened beetles). In some beetles a colour change was observed during the first four
Figure 1. Distribution of Poecilus lepidus in Europe. Map from Turin (2000). Places were the
parental generations of our laboratory strains were collected are marked: LH = Lüneburger Heide
(Germany), LM = Lago Maggiore, Plg = Apennine (subspecies gressorius), both places in Italy.
188 W. Paarmann et al.
weeks of their life. Especially in the red (copper-coloured) morph some beetles appeared
to be green during the first weeks of their life, but then at least the elytra changed to
red. Such colour changes after moulting are due to the post-ecdysal development of
the colour producing layers as shown for Cicindelids by Schultz & Rankin (1985), who
described a gradual increase of the thickness of the layers.
A series of 176 P. lepidus beetles, collected with pit fall traps in the ‘Lüneburger
Heide’ during a period from 11.04. to 14.10.81 (see Mossakowski et al., 1990), were
sorted for their colour morphs.
All crossing experiments were done with virgin females, mostly in groups between
four and ten females and a similar number of males. Females of P. lepidus have a spermatheca and are able to store sperm from multiple mating (De Vries, 2000). In consequence,
the numbers of mating could not be counted and the offspring cold not be separated in
our breeding experiments. There is only one exception: the females, homozygous for the
allele violet (V) (see Tab. 4).
We use the chi-square test of Exel (Chitest) to determine whether there was a significant difference between observed and expected frequencies (95% confidence level).
3. RESULTS
3.1. Heredity of the colour morphs from the Italian strain
The colour morphs blue and black are homozygous while the colour morph green is heterozygous: carrying one allele of the blue and one of the black morph. In the heterozygous
type the gene expression is intermediate (green). The heredity follows the Mendelian
rules. The deviations from the expected values are not significant.
Table 1. Heredity of the colour morphs blue (bb), black (ss), and green (bs) from the Italian strain
of Poecilus lepidus. Genotypes in brackets.
coupled colour morphs
blue (bb) x blue (bb)
resulting colour morphs
blue (bb)
F1 (n)
429*
F1 (%)
100
black (ss) x black (ss)
black (ss )
50
100
blue (bb) x black (ss)
green (bs)
47
100
blue (bb) x green (bs)
green (bs)
blue (bb)
green (bs)
black (ss)
green (bs)
blue (bb)
black (ss)
36
32*
20
13
60
29
34
54
46
61
39
49
24
28
black (ss) x green (bs)
green (bs) x green (bs)
* including P. l. gressorius
Heredity of the elytral colour in adults of Poecilus lepidus Leske (Coleoptera, Carabidae) 189
3.2. Heredity of the colour morphs from the German strain
The following distribution of colour morphs was found in the catch series from 1981 in
the ‘Lüneburger Heide’ (n=176):
red
144 (82%)
yellowish green 17 (10%)
bluish green
6 ( 3%)
black
9 ( 5%)
No specimen of the colour morph violet was among them.
Red (copper coloured), yellowish green, bluish green
The heredity of the colour morphs red, yellowish green and bluish green is summarized
in Table 2. It is similar to the heredity of the Italian colour morphs black, green, and blue.
Bluish green and red are homozygous while yellowish green is heterozygous for one red
and one bluish green allele with an intermediate gene expression. No specimens of the
German black colour morph were available for the crossbreeding experiments.
Table 2. Heredity of the colour morphs bluish green (gg), red = copper coloured (rr), and yellowish
green (gr) from a German strain of Poecilus lepidus. Genotypes in brackets.
coupled colour morphs
red (rr) x red (rr)
resulting colour morphs
red = copper coloured (rr)
F1 (n)
49
F1 (%)
100
bluish green (gg) x bluish green (gg)
bluish green (gg) x red (rr)
bluish green (gg)
yellowish green (gr)
17
46
100
100
red (rr) x yellowish green (gr)
yellowish green (gr)
red (rr)
23
20
53
47
bluish green (gg) x yellowish green (gr)
yellowish green (gr)
bluish green (gg)
24
31
44
56
yellowish green (gr) x yellowish green (gr)
yellowish green (gr)
bluish green (gg)
red (rr)
19
12
12
44
28
28
The deviations from the expected values are not significant.
Violet
During the crossbreeding experiments some unexpected results occurred which made it
necessary to differentiate an additional allele V for the colour violet.
190 W. Paarmann et al.
Violet beetles are heterozygous for the dominant allele V and a recessive allele:
either red (r) or bluish green (g) (Tab. 3)
Table 3. Heredity of the colour morph violet (V) in the ‘Lüneburger Heide’ strain (Germany).
Genotypes in brackets
coupled colour morphs
violet (Vr) x red (rr)
resulting colour morphs
violet (Vr)
red (rr)
violet (Vr) x violet (Vr)
violet (Vr + VV?)
red (rr)
28
12
70
30
3:1
violet (Vr) x violet (Vr)
(F1 see Tab. 4)
violet (Vr) x bluish green (gg)
violet (Vr +VV?)
red (rr)
violet (Vg)
yellowish green (gr)
violet (Vr + Vg)
yellowish green (gr)
red (rr)
40
20
14
23
36
8
3
67
33
38
62
77
17
6
3:1
violet (Vr) x yellowish green (gr)
F1 (n)
21
38
F1 (%)
36
64
expected
1:1
1:1
2:1:1
The deviations from the expected values (Vr x rr: p-value = 0.023, Vr x gr: p-value
= 0,01) may be partly due to the relative low numbers. The occurrence of homozygote
violet specimens could not be verified. The relationship of about 2:1 in the results instead
of the expected value 3:1 may be due to a deficiency of this genotype.
In the ‘Lüneburger Heide’ population V is the most rare allele. That means that
under natural conditions violet beetles are mostly heterozygotes. Under breeding conditions in the laboratory we got some females with nearly black elytra. Only the brim of
the elytra was violet. When we crossbred them with red males (rr) all resulting beetles of
the F1 were violet (n = 28). In this experiment 4 females were included and we did not
know if we got the offspring from one or more females. Therefore we separated them in
a second breeding experiment but the offspring of all females was violet (female 1 - n =
26, female 2 - n = 42, female 3 - n = 50, female 4 - n = 31). The results of both experiments are summarized in Table 4.
Table 4. Heredity of the colour morph black with a violet brim of the elytra in the ‘Lüneburger
Heide’ strain (Germany). Genotypes in brackets.
coupled colour morphs
black with a violet brim(VV) x red (rr)
resulting colour morphs
violet (Vr)
F1 (n)
177
F1 (%)
100
Homozygous females therefore do not show the typical violet colour. They look
almost black with only a small violet brim around the elytra. Up to now we did not
detected any homozygous males for the variant V.
Heredity of the elytral colour in adults of Poecilus lepidus Leske (Coleoptera, Carabidae) 191
3.3 Crossbreeding of colour morphs from the German and Italian strains
From the crossbreeding experiment we can conclude that the red allele (R - German
strain) is dominant over the allele black (s - Italian strain) in heterozygous beetles.
Similarly bluish green (G - German strain) is dominant over blue (b - Italian strain).
Heterozygous beetles of the allele combinations gs and br are green like the combinations bs (Italian strain) and gr (German strain). They show a similar intermediate colour
and therefore a co-dominant allele expression. We did not try to divide the green colour
morphs of these crossbreeding experiments into a German type (yellow green) or an
Italian type (darker green) (see figure 2 and 3) due to the fact that these phenotypes are
sometimes difficult to distinguish in given specimens (the subsuming of similar alleles is
not an argument against the heredity of the colour morphs). The single crossing experiments are summarized in Table 5.
Figure 2. Colour morphs of the Lago Maggiore strain: a) black (ss), b) blue (bb) and c) dark
green (bs). Alleles in brackets.
Figure 3. Colour morphs of the Lüneburger Heide strain: a) red (rr), b) bluish green (gg), c)
yellowish green (gr) and d) violet (Vr).
192 W. Paarmann et al.
Table 5. Crossbreeding experiments of German and Italian colour morphs of Poecilus lepidus.
Allele combinations in brackets, I - Italian, G - German colour morph; hybrids are not marked
with I or G.
coupled colour morphs
black (ss) I x red (RR) G
red (Rs) x red (Rs)
resulting colour morphs
red (Rs)
red (RR, Rs)
black (ss)
F1 (n)
37
30
8
F1 %
100
79
21
blue (bb) I x red (rr) G
green (br)
57
100
black (ss) I x yellowish green (gr) G
green (gs)
red (Rs)
green (gs)
bluish green (Gb)
19
16
13
17
54
46
43
57
bluish green (GG, Gb)
blue (bb)
bluish green (Gb)
blue (bb)
green (gs + br)
bluish green (Gb)
red (Rs)
8
5
7
6
21
16
9
62
38
54
46
46
35
20
green (bs) I x bluish green (gg) G
bluish green (Gb) x bluish green (Gb)
bluish green (Gb) x blue (bb)I
green (bs) I x yellowish green (gr) G
The deviations from the expected values are not significant.We did not yet crossbred the colour morph violet from the German strain with the colour morphs from the
Italian strains.
4. DISCUSSION
We have to conclude that there are at least two alleles (on one locus) from the Italian
strain and three alleles (on one locus) from the German strain, which are controlling in
a dominant, co-dominant or recessive way the colour of the elytra. Few rare phenotypes,
e.g., black from the German strain could not be incorporated into the experiments. Crossbreeding between specimens with alleles from both origins proof, that some of the alleles
are different (despite the fact that the phenotypes are very similar) and that the alleles
controlling the colour are localized on the same locus. The high number of colour morphs
of Poecilus lepidus is therefore controlled by one gene locus with multiple alleles.
Besides Poecilus lepidus, some other polychromous carabid beetles occur in heath lands,
peat bogs and other open habitats, e.g., P. versicolor, P kugelanni, Carabus nitens, Cicindela
campestris (cf. Schatzmayr, 1942-43, Mossakowski, 1980, Assmann & Forman, 1981),
Agonum ericeti (Främbs et al., 2002), and Agonum sexpunctatum (own observation).
The heredity of colour morphs of polychromous species has been only rarely studied
in carabid beetles. Puisségur (1964) used crossbreeding experiments to show that blue
and green colour morphs of Carabus solieri are controlled by two co-dominant alleles.
Heredity of the elytral colour in adults of Poecilus lepidus Leske (Coleoptera, Carabidae) 193
The blue or green coloured elytra in Carabus auronitens specimens is controlled by two
alleles (the genetic variant for green is dominant over the one for the blue elytra) (Puisségur, 1964). Liebherr (1983) found in his study on the morphs of Agonum decorum
independent alleles producing the red and hirsute phenotype which were dominant to
those producing the green and glabrous conditions.
Moreover, Puisségur (1964) was able to show that the genetic basis of the coloration
of elytra and pronotum are independent from each other (interspecific crossbreeding
with C. rutilans and C. hispanus). From our study on Poecilus lepidus we believe that the
genetic basis for the colour of the pronotum and the elytra seems to be controlled by
two different loci. This aspect should be kept into mind if studying the selection pressure
on colour morphs in the field.
The pigment within the cuticle of all colour morphs of P. lepidus is black. This is
a very common pigment colour among ground dwelling beetles like Carabidae and
especially darkling beetles (Tenebrionidae). Black colour might be useful for body temperature regulation. Especially beetles in arid environments raise their body temperature
above air temperature by sun basking. Earlier activity in the morning helps to save body
water (less evaporation). It can also be used to reach maturity earlier in the season, as
shown by the desert dwelling ground beetle Thermophilum sexmaculatum (Erbeling &
Paarmann, 1985, 1986). Forest dwelling and nocturnal ground beetle of the temperate
zone are more often black than beetles in open landscape and with preferred diurnal
activity (Lauterbach, 1964, Thiele & Weber, 1968, Luff, 1978, Löser, 1980, Desender et
al., 1984, Kegel, 1990). Dark colour may be a good camouflage in darker environments
like leaf litter in the forest. For night active beetles there is no need to have other colours
than black, while it may be of selective advantage to have other colours if the beetles are
day active: for example green in a green environment of grasses and herbs at a meadow.
It could lower the chances to be caught by a day active bird, compared to other colour
morphs living in the same habitat. Such diurnal bird species seem to play a big roll as
ground beetle predators (Thiele, 1977). Red colour morphs, dominant in heath land, may
be better camouflaged in this habitat than the other colour morphs.
Schultz (1986) pointed out that background matching of colour patterns is present in
many American Cicindela species. He also described the occurrence of a few other species with
highly reflective colours. When they fly into deep shadow the beetles disappear abruptly.
Along riversides of the Apennine only the blue morph of P. l. gressorius appears. The
whole population is homozygous for the allele blue (b). In other habitats with differing
surface colours or denser green vegetation, other morphs might be favoured by selection.
Deviations from Hardy-Weinberg-equilibrium (HWE) can give first indications for a
selection against any genotypes. The data from the Lüneburger Heide (cf. above) give
only weak (and not highly significant) evidence because of more homozygotes of the
variant g than expected after HWE. In habitats were the heterozygous morph green has
an higher adaptive value, the homozygous morphs blue (bb) or black (ss) as well as red
(rr) or bluish green (gg) cannot vanish. One forth of the offspring of green parents will
be black or red, and one fourth blue or bluish green.
194 W. Paarmann et al.
Reasons for rarity of the violet morph, caused by the dominant allele V, may be
due to a developmental depression of the homozygote. Field studies are necessary to
get more information about the percentage of beetles of this colour in the ‘Lüneburger
Heide’ population. No specimen was found in a series of 176 beetles.
The studied colours, with the exception of black, are structural colours, caused
through light reflection and interference by thin layers in the upper cuticle in front of the
black background. So our study of the heredity of colour morphs was a study of the heredity
of the thickness of light reflecting layers. For details see Mossakowski et al. (2008).
Wilmer & Unwin (1981) studied heat gain and loss in relation to weight and reflectance of insects. They emphasized size and reflectance as an important feature, which
regulate body temperature. For moderate or large sized diurnal species it should be a
great advantage to be highly reflective. They can thus avoid overheating by radiation. To
be coloured may have two advantages to diurnal Carabidae: protection against birds and
overheating (Mossakowski, 1980, Schultz, 1986).
It is still not known if there are differences in the daily activity of the different
colour morphs. May be, that darker morphs tend to be more nocturnal than brightly
coloured ones. Life span of colour morphs of the P. lepidus Lago Magiore strain differs
under long day (16h light) and short day (8h light) at 20 °C (Paarmann, 1990). In the
first case life span is significantly shorter for green males compared to blue and black
males, while under short day condition it is significantly shorter in black males than in
blue or green.
ACKNOWLEDGEMENTS
Collecting the P generation of the P. lepidus gressorius strain was possible only because
of a detailed description of the population site in the Apennine mountains by E. Contarini (Bagnacavallo Ravenna) and an invitation to Italy by P. Brandmayr (Universita
della Callabria, Arcavavcata di Rende). S. Albrecht bred and crossed P lepidus beetles
with great enthusiasm.
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