Allometric Relationship Between Genitalic Size and Body Size in Two Species of

Mordellid Beetles (Coleoptera: Mordellidae)

Author(s): Gregor Schmitz, Klaus Reinhold, Peter Wagner
Source: Annals of the Entomological Society of America, 93(3):637-639.
Published By: Entomological Society of America
DOI: http://dx.doi.org/10.1603/0013-8746(2000)093[0637:ARBGSA]2.0.CO;2
URL: http://www.bioone.org/doi/full/10.1603/0013-8746%282000%29093%5B0637%3AARBGSA
%5D2.0.CO%3B2

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 MORPHOLOGY, HISTOLOGY, AND FINE STRUCTURE

Allometric Relationship Between Genitalic Size and Body Size in Two

Species of Mordellid Beetles (Coleoptera: Mordellidae)
GREGOR SCHMITZ, KLAUS REINHOLD, AND PETER WAGNER
Institut für Evolutionsbiologie und Ökologie, An der Immenburg 1, D - 53121 Bonn, Germany

Ann. Entomol. Soc. Am. 93(3): 637Ð639 (2000)

ABSTRACT Although there is often an isometric relationship between the size of nongenital parts
and body size, data on insects and spiders indicate that male genitalic size is only slightly inßuenced
by variation in body size. Eberhard et al. (1998) interpreted the low inßuence of body size on
genitalic size as evidence of stabilizing selection based on cryptic female choice. Data on this aspect
of sexual selection are rare. We compared the variation in genitalic size with the variation in elytra
length as an indicator for body size in the males of two mordellid beetle species, Mordellistena weisei
Schilsky and M. bicoloripilosa Ermisch (Coleoptera: Mordellidae), to Þnd additional indications of
stabilizing selection on genital traits. The slopes of the allometric regressions between paramere size
and elytra length were ⬍1 for left and right parameres in both species, and signiÞcantly less in three
of the four comparisons. There was no signiÞcant correlation between elytra length and genitalic
size in either species and the coefÞcient of variation was similar for paramere size and elytra length.
Although there is no unequivocal proof for the existence of cryptic female choice, the results
reinforce the evidence in favor of stabilizing selection on genitalic size.

KEY WORDS Mordellidae, allometry, genital structure, cryptic female choice, body size,
parameres

WHEN ANIMALS VARY substantially in body size, the size
of their body parts often differs as well. An isometric
relationship exists when the size of body parts is pro-
portional to body size. When the size of body parts
differs relative to body size, an allometric relationship
is better suited to describe the variation in the size of
body parts. A speciÞc allometric relationship is known
for weapons and other traits inßuenced by intersexual
selection. In this case, larger animals have body parts
of large relative size, which leads to a slope greater
than one in double logarithmic regression. In a coreid
bug (Eberhard 1998) that uses its legs in combat, large
individuals have hind legs of larger relative size. A
similar relationship is also known for the horns of dung
beetles (Scarabaeidae: Emlen 1994), stag beetle man-
dibles (Lucanidae: Clark 1977), and the eye-stalk of
stalk-eyed ßies (Diopsidae: Burkhardt et al. 1994). The
inverse relationshipÑa slope less than oneÑwas re-
cently documented for genitalic size in a sample of
spiders and insects, where larger males have relatively
smaller genitalia than smaller males. This weak inßu-
ence of body size also leads to smaller variation in
genitalic size than in body size, and is interpreted as
the result of stabilizing selection on genitalic size
(Eberhard et al. 1998).
In the framework of comparative taxonomic studies
(Schmitz and Wagner 2000), we measured paramere
length as an indicator of genitalic size and elytra length
as an indicator of body size (Eberhard et al. 1998) in
the males of two mordellid beetles species. To deter-
mine whether stabilizing selection on genitalic size
also occurs in mordellid beetles, we tested whether
the allometric regression between these two para-
meters yields a slope of ⬍1. In addition, we tested
whether the coefÞcient of variation is smaller for
paramere size than for elytra length.
Materials and Methods
Species. We examined 22 males of Mordellistena
bicoloripilosa Ermisch and 36 males of M. weisei Schil-
sky, two of the 67 species of the genus occurring in
Germany (Köhler and Klausnitzer 1998). Within the
subgenus Mordellistena, these two species belong to
the weisei Schilsky group, which is characterized by
elytra trichomes of two different colors (Ermisch
1967). The two species can easily be distinguished by
the shape of their parameres (Schmitz and Wagner
2000). M. bicoloripilosa and M. weisei are oligophagous
stem miners of the Compositae and are most fre-
quently found in mugwort, Artemisia vulgaris L. The
males were reared from mugwort stems collected in
the area of Bonn (western Germany) (Schmitz 1995).
The imagines hatched mainly between mid-June and
the beginning of August, often from the same plant
stands and sometimes even from the same individual
stems (Schmitz 1995, 1996). Additional specimens
from other sites in Germany were also included
(Schmitz and Wagner 2000).
Methods of Measurement. As in other mordellids,
the parameres are asymmetric in the two examined
species. Therefore, both the left and right ones were

0013-8746/00/0637Ð0639$02.00/0 䉷 2000 Entomological Society of America

638 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 93, no. 3

surement accuracy: 10 ␮m). Paramere length is de-

Þned as longest distance from the base to the top of the
dorsal appendix (i.e. the trichome-bearing appendix)
measured along the longitudinal axis (Fig. 1). Elytra
length was measured from the fore margin of the
“shoulder” of the right elytra to its apex (stereomi-
croscope scale: 25⫻, measurement accuracy: 0.1 mm).
Statistics. The measured values (mm) were trans-
formed into their natural logarithms and the inßuence
of body size (as elytra length) on the size of the
parameres was estimated by linear regression. An F-
test was used to compare the variance of paramere
length between the two species. We estimated the
relationship between left and right parameres with
Spearman rank correlation. As with all other statistics,
the slope between paramere length and elytra length
was estimated using Statistical Program for Social Sci-
ences (SPSS 1999).

Fig. 1. Parameres of M. weisei (M. w.) and M. bicoloripi-
losa (M. b.): their natural position within the genitals (sche-
matic), structural detail (ventral view), and the method of
measurement (ae, aedeagus; ep, epimere of the phallobase;
da, dorsal appendix; lp, left paramere; phb, phallobase; rp,
right paramere; va, ventral appendix).
measured. After macerating the caudal parts of the
abdomen, the parameres were prepared and Þxed on
clear plastic plates. The total length of the parameres
was measured using a microscope scale (100⫻, mea-
Results
In both Mordellistena species studied here, the
length of the left and right parameres do not correlate
with elytra length (P ⬎ 0.05 for all four correlations).
The regression slopes range between 0.01 and 0.57.
They are not signiÞcantly different from 0 in any case,
whereas they are signiÞcantly different from one in
three cases (Table 1). The coefÞcients of variation for
elytra length (CVelytra: M. bicoloripilosa ⫽ 0.085, M.
weisei ⫽ 0.045) and genitalic size (see values in Table
1) are similar. The length of the left and right
parameres correlate signiÞcantly in both species
(Spearman correlation coefÞcients: RspM.b. ⫽ 0.66,
P ⬍ 0.001, RspM.w. ⫽ 0.66, P ⫽ 0.001).
Mordellistena bicoloripilosa (n ⫽ 36) show a signif-
icantly larger variance in the length of the left
parameres than M. weisei (n ⫽ 22). VarM.b. ⫽ 7.2 ⫻
10⫺4, VarM.w. ⫽ 3.0 ⫻ 10⫺4, F ⫽ 2.4; df ⫽ 35, 21; P ⬍
0.05. A difference in the variance is not observed in the
right parameres (VaM.b. ⫽ 8.1 ⫻ 10⫺4, VarM.w. ⫽ 6.7 ⫻
10⫺4, F ⫽ 1.2; df 35, 21; not signiÞcant).
Discussion
In accordance with the general conclusion of Eber-
hard et al. (1998), the following applies to both beetle
species: (1) the slopes of the allometric regression
were ⬍1 for both genitalic traits, and (2) there was no
overall signiÞcant correlation between body size and
genitalic size. These results show that genitalic size is

Table 1. Linear regression between paramere length and elytra length in M. weisei (M.w.) and M. bicoloripilosa (M.b.): mean paramere
length, coefficients of variation, slopes of log-log regression with standard error (SE), significance of the deviation of the slope from one,
regression coefficients (r), and sample size (n)

Species and paramere Mean, mm CV Slope ⫾ SE P (slope ⫽ 1) r n

M. w. left paramere 0.33 0.052 0.26 ⫾ 0.25 ⬍ 0.01 0.23 22
M. w. right paramere 0.34 0.075 0.57 ⫾ 0.37 NS 0.32 22
M. b. left paramere 0.36 0.075 0.01 ⫾ 0.15 ⬍ 0.001 0.01 36
M. b. right paramere 0.31 0.091 0.08 ⫾ 0.18 ⬍ 0.001 0.08 36

M. w., mean of elytra length ⫽ 2.61 mm; M. b., mean of elytra length ⫽ 2.47 mm.
May 2000 SCHMITZ ET AL.: ALLOMETRIC RELATIONSHIPS IN MORDELLIDAE
at most only weakly inßuenced by body size and thus
reinforce the evidence in favor of stabilizing selection
on genitalic size. Given that genitalic size is inßuenced
by stabilizing selection one would expect a lower co-
efÞcient of variance for genitalic size than for body
size. However, in line with the results of Eberhard et
al. (1998), the coefÞcients of variation for genitalic
size and body size were similar; a result that may be
caused by the greater measurement error for the
smaller size of the genitalic parts.
In principle, several selection regimes can cause
stabilizing selection on genitalic size. Eberhard et al.
(1998) proposed that a mechanical lock-and-key
mechanism is an unlikely explanation for the low vari-
ation observed in genitalic size because of the fact that
similar allometric values were measured for male geni-
talic structures that contact rigid female genitalic
structures, and for those that contact soft female geni-
talic structures during copulation. Thus, they inter-
preted allometric values as evidence for cryptic female
choice. When there is also variation in female genitalic
size, it is assumed that males that can best stimulate
average females have an advantage and are able to
adequately stimulate a large proportion of females.
However, it is generally difÞcult to separate the
effects of male variation in sperm competition and
cryptic female choice (Telford and Jennions 1998). In
our opinion, the low allometric values observed can
also be explained apart from cryptic female choice
when sperm competition strategies cause stabilizing
selection on male genitalic size. Male genitalic size
might be optimized for sperm removal, quick sperm
transfer, or sperm displacement (Birkhead and Møller
1998). In all of these cases, males may have an advan-
tage if their genitalia Þt best to the most typical female.
Stabilizing selection on male genitalic size through
selection on sperm competition capability is thus
equally as possible as stabilizing selection through
cryptic female choice. Only careful experiments sim-
ilar to those conducted by Arnqvist and Danielsson
(1999) can resolve the inßuence of cryptic female
choice and male sperm competition capability on vari-
ation in male genitalic size.
639
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