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Narita S, Pereira RAS, Kjellberg F, Kageyama D. Gynandromorphs and intersexes: potential to understand the mechanism of sex determination in arthropods. Terrestrial Arthropod Reviews

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Abstract

Arthropods are sexually dimorphic. An arthropod individual usually diff erentiates into a male or a female. With very low frequencies, however, individuals with both male and female morphological characters have repeatedly been found in natural and laboratory populations of arthropods. Gynandromorphs (i.e., sexual mosaics) are genetically chimeric individuals consisting of male and female tissues. On the other hand, intersexes are genetically uniform (i.e., complete male, complete female or intermediate in every tissue) but all or some parts of their tissues have either a sexual phenotype opposite to their genetic sex or an intermediate sexual phenotype. Possible developmental processes (e.g., double fertilization of a binucleate egg, loss of a sex chromosome or upregulation/downregulation of sex-determining genes) and causal fac-tors (e.g., mutations, genetic incompatibilities, temperatures or endosymbionts) for the generation of gynandromorphs and intersexes are reviewed and discussed. © Koninklijke Brill NV, Leiden, 2010.
© Koninklijke Brill NV, Leiden, 2010 DOI 10.1163/187498310X496190
Terrestrial Arthropod Reviews 3 (2010) 63–96 brill.nl/tar
TAR
Gynandromorphs and intersexes: potential to understand the
mechanism of sex determination in arthropods
Satoko Narita 1 , Rodrigo A. S. Pereira 2 , Finn Kjellberg 3 and Daisuke Kageyama 1,*
1 National Institute of Agrobiological Sciences, Owashi 1-2, Tsukuba, Ibaraki 305-8634, Japan
2 Depto. de Biologia/FFCLRP-Universidade de São Paulo, Av. Bandeirantes, 3900,
14040-901, Ribeirão Preto, SP, Brazil
3 CNRS-CEFE, 1919 Route de Mende, 34293, Montpellier, Cédex 5, France
*Corresponding author; e-mail: kagymad@aff rc.go.jp
Received: 20 January 2010; accepted: 25 February 2010
Summary
Arthropods are sexually dimorphic. An arthropod individual usually diff erentiates into a male or a female.
With very low frequencies, however, individuals with both male and female morphological characters have
repeatedly been found in natural and laboratory populations of arthropods. Gynandromorphs (i.e., sexual
mosaics) are genetically chimeric individuals consisting of male and female tissues. On the other hand,
intersexes are genetically uniform (i.e., complete male, complete female or intermediate in every tissue)
but all or some parts of their tissues have either a sexual phenotype opposite to their genetic sex or an
intermediate sexual phenotype. Possible developmental processes (e.g., double fertilization of a binucleate
egg, loss of a sex chromosome or upregulation/downregulation of sex-determining genes) and causal fac-
tors (e.g., mutations, genetic incompatibilities, temperatures or endosymbionts) for the generation of
gynandromorphs and intersexes are reviewed and discussed.
© Koninklijke Brill NV, Leiden, 2010.
Keywords
Arthropod; gynandromorph; insect; intersex; sex determination; sexual mosaic
Introduction
Unlike in plants and some groups of animals, hermaphroditism is extremely rare in
arthropods (but see Hughes-Schrader and Monahan, 1966 , for the exceptional case of
coccids). In other words, an arthropod individual diff erentiates into a complete male
or a complete female and does not become an intermediate, i.e., arthropods are sexu-
ally dimorphic. However, developmental defects, which occur at low frequencies under
natural conditions, can lead to morphologically anomalous individuals with both
male and female traits. Owing to their strangeness and rarity, such individuals often
64 S. Narita et al. / Terrestrial Arthropod Reviews 3 (2010) 63–96
attract amateur insect collectors, and indeed, some of them are sold at surprising prices
(e.g., the website Insect-Sale).
To date, numerous papers have reported the occurrence of arthropod individuals
presenting phenotypically male and female parts.  e distribution patterns of the male
and female parts in such individuals are variable. For example, the male and female
parts can be clearly bilateral, patchily distributed or uniformly mixed. Individuals with
clear borders between the male and female parts are often referred to as gynandro-
morphs while those with ambiguous or no borders are usually referred to as intersexes.
Although this classifi cation based only on the external morphologies is quite common,
it does not refl ect the genetic and developmental processes.  e biological defi nition of
a gynandromorph (synonymous with a sexual mosaic) is a genetically chimeric indi-
vidual while that of an intersex is a genetically uniform individual (e.g., Goldschmidt,
1934 ; Laugé, 1985 ), which will be explained in the following sections.
In this paper, we review (1) the occurrences of gynandromorphs and intersexes in
arthropods, (2) the mechanisms of arthropod sex determination and (3) the possible
developmental processes for the generation of gynandromorphs and intersexes. Finally,
we discuss the signifi cance of sexually anomalous individuals for better understanding
the still unexplored sex-determining systems of arthropods.
Occurrence of male and female traits in a single individual
e occurrence of individuals consisting of phenotypically male and female parts has
been reported repeatedly in arthropods.  ey have been found in both natural and
laboratory populations of almost all orders of insects as well as non-insects (see Table 1).
In many cases, however, it remains to be clarifi ed whether they are gynandromorphs or
intersexes and they are arbitrarily called gynandromorphs or intersexes. Considering
the underlying processes, clearly bilateral individuals are likely to be gynandromorphs
(see below for details).
We should exercise caution regarding the point that heterogeneous individuals are
more likely to be detected in species in which the sexes are strikingly dimorphic than
species in which the sexes are less strikingly dimorphic. Consequently, estimating the
frequencies of occurrence merely through publication records can be misleading.
Moreover, owing to their conspicuous features, the occurrence of clearly bilateral
gynandromorphs may be published more frequently than the occurrence of ambiguous
forms of gynandromorphs and intersexes.
To discriminate gynandromorphs and intersexes, it is necessary to identify the
respective sexual genotypes of the tissues presenting male and female phenotypes. In
species for which molecular analyses are unavailable, conventional karyotyping is
widely used for sexual genotyping. In lepidopteran insects with a ZW-ZZ sex chromo-
some constitution, simple observation of the sex chromatin allows us to discriminate
the male (ZZ) and female (ZW) genotypes. Sex chromatin is a highly condensed W
chromosome that is visible in interphase nuclei (Traut and Marec, 1996 ). In insects for
which molecular data are available, diagnostic polymerase chain reaction amplifi ca-
tions using Y-specifi c or W-specifi c molecular markers also allow us to discriminate the
S. Narita et al. / Terrestrial Arthropod Reviews 3 (2010) 63–96 65
male and female genotypes. In addition, transcriptional analyses of genes expressed in
sex-specifi c manners may also be used to confi rm the male and female phenotypes in a
single individual.
Sex-determining systems of arthropods
In the majority of insects, sex is genetically determined. For example, dipteran insects
like the fruit fl y Drosophila melanogaster have a male-heterogametic sex chromosome
constitution (i.e., XX: female; XY: male). Lepidopteran insects like the silkworm
Bombyx mori have a female-heterogametic chromosomal constitution (i.e., ZZ: male;
ZW: female). Hymenopteran insects like the honeybee Apis mellifera have a haplodip-
loid sex determination system, in which fertilized (2n) eggs become females and unfer-
tilized (n) eggs develop into males (Bull, 1983 ; Werren and Beukeboom, 1998 ; Heimpel
and de Boer, 2008 ).  e molecular mechanisms underlying sex determination and dif-
ferentiation in the model insect D. melanogaster are well understood. At a very early
embryonic stage, each cell determines its sex independently, and once determined, the
sex of each cell is maintained during later development through a gene expression cas-
cade consisting of Sex-lethal ( Sxl ), transformer ( tra ), doublesex ( dsx ) and other genes, in
which sex-specifi c mRNA splicing plays an important role (Schutt and Nöthiger,
2000 ).
Although the molecular mechanisms of sex determination in non- Drosophila sys-
tems are not well understood, all the sex-determining mechanisms in insects are pro-
posed to be variations of a single model consisting of a master regulator gene (like
Sex-lethal in D. melanogaster ) at the top of the cascade and the highly conserved dou-
blesex gene at the bottom of the cascade (Figure 1) (Nöthiger and Steinmann-Zwicky,
1985 ; Hoy, 2003 ). Sex determination in a cell-autonomous manner is also believed to
be widespread among insects on the basis that sexually mosaic individuals often occur
in a diverse array of insects (Laugé, 1985 ).
Like insects, some of the non-insect arthropods are also considered to have geneti-
cally based sex determination. However, they diff er from insects as sexual diff erentia-
tion is deeply aff ected by sex hormones that are secreted by particular organs (e.g., the
androgenic gland in crustaceans). It is known that some crustaceans are subject to
environmental sex determination. In the shrimp Gammarus duebeni , sex is deter-
mined post-conception in response to photoperiod cues and is then fi xed for life
(Legrand et al., 1987 ).
Gynandromorphs
Gynandromorphs are chimeric individuals consisting of genetically male and geneti-
cally female tissues. In each cell of a gynandromorph, the genetic sex (e.g., sex chromo-
some constitution) is consistent with the sexual phenotype. In other words, the
sex-specifi c expression or alternative splicing of sex-determining genes should be
consistent with the genetic sex (Figure 2a). Gynandromorphs allow us to carry out
fate mapping, which reveals the developmental history of each cell in the body of an
66 S. Narita et al. / Terrestrial Arthropod Reviews 3 (2010) 63–96
organism (Garcia-Bellido and Merriam, 1969 ; Hotta and Benzer, 1972 ; Mori and
Perondini, 1984 ; Milne and Rothenburler, 1983a ; Myohara, 1994 ).  e underlying
causes of the generation of gynandromorphs are described below.
Gynandromorphs generated by loss or damage of a sex chromosome
In some insects like D. melanogaster , the number of X chromosomes (or Z chromo-
somes) relative to the number of autosomes is the initial key factor for sex determina-
tion. Loss of one of the X chromosomes during mitosis results in individuals with XX
and X0 cells (Figure 3a). If the loss occurs during the fi rst mitotic division (cleavage),
individuals with 50% male tissues and 50% female tissues appear. Furthermore, the
later the X chromosome is lost, the smaller the male part in the gynandromorph will
be. In D. melanogaster , a mutant strain with an unstable ring-X chromosome often
produces gynandromorphs (Hinton, 1955 ). In this strain, an X chromosome is easily
lost during early cleavage, resulting in the frequent appearance of individual that are
gynandromorphic at various degrees.  e generation of gynandromorphs owing to loss
lFl Ml
Maternal genes
Female zygote Male zygote
Zygotic genes
ON OFF
Master regulator gene
dsx-like gene
??
Male-splice form
Female-splice form
Female
differentiation
genes
male
differentiation
genes
Female
differentiation
genes
male
differentiation
genes
ON OFF OFF ON
Female phenotype Male phenotype
Figure 1. A proposed general model of sex determination in insects.  is model assumes that the actions
of both maternal genes and zygotic genes aff ect the expression of a master regulator gene, which corre-
sponds to Sex-lethal in D. melanogaster . e expression of the master regulator gene activates or suppresses
the expression of subsequent genes (downstream genes). At the end of the hierarchical gene expression
cascade, a highly conserved doublesex -like gene is subjected to alternative RNA splicing and produces the
male-specifi c protein DSX
M or female-specifi c protein DSX
F
. e sex-specifi c DSX proteins activate and
suppress a series of sex-specifi c diff erentiation genes, leading to either the female phenotype or the male
phenotype.
S. Narita et al. / Terrestrial Arthropod Reviews 3 (2010) 63–96 67
(a) (b) (c) (d)
Gynandromorph Intersex
Genotype XX XX XX XX
XX
XY XY XY XY XXY XXYXXY
Phenotype
Phenotypically mosaic Phenotypically
uniform
Figure 2. Relationships between the genotype and the phenotype of individual cells in gynandromorphs
and intersexes.  e squares indicate single cells (maleness: black; femaleness: white; intermediate: gray).
e dotted lines indicate single individuals. For simplicity, a single individual is assumed to be composed
of three cells.  e genotype and phenotype are connected by arrows. (a) A gynandromorph (i.e., a sexual
mosaic). (b) An intersex caused by partial feminization of a genetic male. (c) An intersex caused by partial
masculinization of a genetic female. (d) An intersex with an intermediate genotype between male and
female (e.g., a triploid intersex is shown).
of an X chromosome has also been reported in the fl y Sciara ocellaris (Mori and
Perondini, 1980 ).
Gynandromorphs generated from binucleate eggs
Double fertilization of a binucleate egg by X and Y sperms can also give rise to a gynan-
dromorph in Drosophila (Hollingsworth, 1955 ). In lepidopteran insects such as
B. mori , double fertilization of a binucleate egg (ZW) by Z sperms is thought to be the
major cause of the generation of gynandromorphs (Goldschmidt and Katsuki, 1927 ).
Since the female karyotype of B. mori is ZW, the meiotic division separates Z and W
chromosomes, so that one remains in the egg and the other is discarded with the polar
body. If the polar body is accidentally retained in the egg, both nuclei can be fertilized
by Z sperms, resulting in the development of an individual that has male (ZZ) and
female (ZW) tissues (Figure 3b). In some strains of B. mori , the polar bodies are prone
to being retained in the eggs, leading to a high frequency of gynandromorphs
(Goldschmidt and Katsuki, 1931 ). In the hymenopteran insect A. mellifera , fertilized
eggs (2n) become females and unfertilized eggs (n) develop into males (Bull, 1983 ;
Werren and Beukeboom, 1998 ). Accidental production of binucleate eggs (2n) fol-
lowed by fertilization of one of the nuclei can result in the development of an indi-
vidual that has male (n) and female (2n) tissues (Figure 3c).
Gynandromorphs generated by symbionts
Bacteria belonging to the genus Wolbachia (alpha subdivision of the phylum
Proteobacteria) are ubiquitous endosymbionts of insects. In some hymenopteran
insects, Wolbachia -infected females produce unfertilized eggs that parthenogeneti-
cally develop as females. Both antibiotic curing and high temperatures result in male
production with elimination of the bacteria (Legner, 1985 ; Stouthamer and Luck,
68 S. Narita et al. / Terrestrial Arthropod Reviews 3 (2010) 63–96
XX
n
n
(a) (b) (c) (d)
Partial suppression of
Wolbachia-induced
diploidization
Binucleation
In eggs
Loss of a sex chromosome
during cleavage
Binucleation
In eggs followed by
double fertilization
Wolbachia
ZWnnn
X
XX
XX
XX
XX
XX
XX XX
XX
n
nn
nn
ZZn
X
X
XX
ZW
ZZ 2n
n2n 2n 2n
2n
n
X
Honeybee
Apis mellifera
(Hymenoptera)
Wasp
Trichogramma
(Hymenoptera)
Silkmoth
Bombyx mori
(Lepidoptera)
Fruitfly
Drosophila melanoaster
(Diptera)
Figure 3. Possible examples of the developmental processes of gynandromorphs. (a) Loss of an X chromosome during mitosis in Drosophila melanogaster . (b) Double
fertilization of a binucleate egg in Bombyx mori . (c) Fertilization of one of the nuclei in a binucleate egg in haplodiploid insects such as Apis mellifera . (d) Partial sup-
pression of Wolbachia -induced diploidization during mitosis after antibiotic or heat treatments in haplodiploid wasps such as Trichogramma species. See the text for
details.
S. Narita et al. / Terrestrial Arthropod Reviews 3 (2010) 63–96 69
1991 ; Zchori-Fein et al., 1992 ). Treatment with temperatures slightly higher than nor-
mal temperatures can give rise to gynandromorphs at high frequencies (Wilson and
Woolcock, 1960 ; Bowen and Stern, 1966 ; Cabello and Vargas, 1985 ).
e cytogenetic mechanisms of Wolbachia -induced parthenogenesis have been stud-
ied in Trichogramma spp. and Muscidifurax uniraptor (Stouthamer and Kazmer, 1994 ;
Legner, 1985 ). Meiosis is normal. In the fi rst mitotic division, the chromosomes con-
dense properly in prophase but fail to segregate in metaphase, resulting in diploidiza-
tion of the nucleus and homozygosity at all loci.
It is considered that slightly high temperatures incompletely suppress the eff ects of
Wolbachia . Under such conditions, diploidization may occur during the second cleav-
age or a later cleavage instead of the fi rst cleavage and some nuclei escape diploidization
and remain haploid (Figure 3d) (Stouthamer, 1997 ).
Intersexes
In contrast to gynandromorphs, intersexes are genetically uniform (Figure 2b-d). Some
intersexes are genetically intermediate between the typical male genotype and the typi-
cal female genotype (e.g., XXY) and the sex-determining genes in every cell follow
their own genetic signal (e.g., X:A ratio in Drosophila ) (Figure 2d). On the other hand,
other intersexes are genetically purely male or female, but some parts of their bodies
have a sexual phenotype that is opposite to their genetic sex, i.e., partial feminization
(Figure 2b) or partial masculinization (Figure 2c). However, all intersexes should be
caused by defects in the processes of sex determination and diff erentiation, and thus
strict classifi cation of intersexes is of little importance. Here, we arbitrarily classify the
intersexes according to the causal factors.
Intersexes generated by chromosomal aberrations
Genetic intersexes are individuals that are genetically intermediate between male and
female. Triploid intersexes are one such example. In Drosophila , the initial signal for sex
determination is the ratio of the X chromosome number to the autosome (A) number,
in which XX;AA individuals (X/A = 1) become female and XY;AA individuals (X/A =
0.5) become male. In a triploid intersex (XXY;AAA) with an X/A ratio of 0.66, each
cell diff erentiates into an intermediate sexual phenotype (Figure 2d). Triploid inter-
sexes are also known in the moth Solenobia triquetrella (Seiler et al., 1958 ). Other types
of intersexes are genetically purely male or female and some parts of their bodies
are feminized or masculinized during the processes of sex determination and/or
diff erentiation.
Intersexes generated by loss-of-function mutations
In D. melanogaster , loss-of-function mutations such as Sex-lethal ( Sxl ), transformer
( tra ), intersex ( ix ) and doublesex ( dsx ) exhibit sex reversal in their phenotypes.  ese are
examples of phenotypic intersexes. Epistatic interactions between these mutations
revealed by classical genetic experiments have allowed understanding the hierarchical
70 S. Narita et al. / Terrestrial Arthropod Reviews 3 (2010) 63–96
gene expression cascade controlling sex determination in Drosophila (Schutt and
Nöthiger, 2000 ). Furthermore, molecular genetic experiments have revealed that sex-
specifi c mRNA splicing of the Sxl , tra and dsx genes plays a crucial role for sex deter-
mination in Drosophila (Schutt and Nöthiger, 2000 ).
Intersexes generated by crosses between diff erent strains
In the moth Lymantria dispar , crosses between diff erent geographic strains result in the
generation of intersexes that exhibit a uniformly intermediate phenotype (Goldschmidt,
1934 ; Mosbacher, 1973 ). Since the karyotype of these individuals shows the female
genotype (ZW), the cause of the intersexual phenotype is considered to be incomplete
masculinization of genetic females. Likewise, hybridization between the closely related
moths Smerinthus ocellata and Smerinthus populi results in the generation of intersexual
off spring (Morgan, 1909 ). A particular genetic combination may aff ect the normal
expression of the sex-determining genes in L. dispar and the Smerinthus species.  e
generation of intersexes through crosses between diff erent strains or between closely
related species may result from diverging sex-determining systems. It is generally con-
sidered that the sex-determining systems of arthropods are evolving rapidly (Werren
and Beukeboom, 1998 ).
Intersexes generated by epigenetic factors
Environment
Exposure to high temperatures can induce partial feminization of genetic males in the
mosquito Culex stimulans (Brust, 1966 , 1968 ; Brust and Horsfall, 1965 ; Horsfall et al.,
1964 ; Horsfall and Anderson, 1961 , 1965 ; Craig, 1965 ). On the other hand, high
temperatures induce masculinization in the bagworm S. triquetrella (Seiler, 1935 ) and
stick insect Carausius morosus (Bergerard, 1958 , 1961 ). Upregulation or downregula-
tion of sex-determining genes induced by unusual temperatures may be possible causes
of intersexual development. In the shrimp G. duebeni , photoperiods after mating
determine the sex of the off spring (Legrand et al., 1987 ).
Sex hormone
In the fi refl y Lampyris noctiluca , transplantation of the larval male gonad into female
larvae was reported to result in masculinization, suggesting the presence of a sex hor-
mone in these insects (Naisse, 1966a , 1966b ).  is is an exceptional case among insects
as sex hormones are generally considered to be absent. However, similar experiments
carried out recently did not alter the sexual phenotype of these insects (Maas and
Dorn, 2005 ). erefore, the authenticity of hormone-associated masculinization in
these insects and its relationship with the sex-determining genes remain unclear.
Symbionts
In some amphipods such as G. duebeni and Orchestia gammarellus , in which sex
is determined environmentally, males can be partially or completely feminized by
unicellular eukaryotes such as the microsporidian parasites Octosporea eff eminans and
Nosema granulosis , and the haplosporidian parasite Paramartelia orchestiae , leading to
S. Narita et al. / Terrestrial Arthropod Reviews 3 (2010) 63–96 71
intersexual development (Bulnheim and Vavra, 1968 ; Bulnheim, 1977 , 1978 ; Kelly
et al., 2004 ; Ginsburger-Vogel, et al., 1980). In some isopods such as Armadillidium
vulgare , sex is genetically determined but genetic males are known to be completely
feminized by the endosymbiotic bacteria Wolbachia (Figure 2b) (Rigaud et al., 1997).
In A. vulgare , heat treatment attenuates the eff ects of Wolbachia and leads to the gen-
eration of intersexes (Juchault et al., 1980 ). On the other hand, a virus is known to
maculinize some of the tissues of A. vulgare genetic females, leading to the formation
of intersexes (Figure 2c) (Juchault et al., 1991 ).
Wolbachia also have a feminizing ability of insect hosts. In the butterfl y Eurema
mandarina (former name Eurema hecabe , yellow-type), genetic males are completely
feminized by Wolbachia . Antibiotic treatment of E. mandarina infected with feminiz-
ing Wolbachia results in the generation of intersexes (Narita et al., 2007a ). On the
other hand, in the moths Ostrinia scapulalis and Ostrinia furnacalis , naturally occurring
Wolbachia kill genetic males. However, genetic males can be rescued and develop as
intersexes when their mothers are treated with antibiotics or high temperatures
(Kageyama and Traut, 2004 ; Sakamoto et al., 2008 ).  ese observations imply that
Wolbachia have a feminizing eff ect that is lethal for Ostrinia males. It is unknown how
Wolbachia interfere with the sex-determining system of insects. However, epigenetic
modifi cation is suggested to play an important role in the leafhopper Zyginidia pullula ,
in which naturally-occurring Wolbachia cause partial feminization of genetically male
individuals. In Z. pullula , DNA methylation of the feminized males exhibits a similar
pattern with normal females (Negri et al., 2006 , 2009 ).
Not only microorganisms but also nematodes, trematodes and turbellarians can
have signifi cant eff ects on gonad development and/or secondary sexual characters of
various arthropods (reviewed by Baudoin, 1975 and references therein). It has been
clearly shown that the infection by the mermithid nematode Gasteromermis sp. is
known to feminize genetic males of the mayfl y host Baetis bicaudatus morphologically
and behaviorally (Vance, 1996 ).
As described above, various factors are responsible for the generation of intersexes
but the individual factors are likely to aff ect particular steps of a single process of sex
determination (Figure 1). Future studies will allow us to reconcile the seemingly vari-
ous types of intersexes in the context of sex determination and diff erentiation.
Conclusions
Ever since ancient times, people have been eager to know about the diff erences between
males and females. In some societies, animals with both male and female organs were
sanctifi ed and admired. At the present time, such phenomena occurring in insects
attract both insect collectors and researchers.  e high degree of attention can be
inferred from the number of published papers describing gynandromorphism and
intersexuality in arthropod species.
However, most of these papers only describe the external morphology of the anoma-
lous individuals. From the scientifi c perspective, it would be most useful to elucidate
the mechanisms that generate gynandromorphs and intersexes in non- Drosophila
72 S. Narita et al. / Terrestrial Arthropod Reviews 3 (2010) 63–96
systems, in which the mechanisms of sex determination and diff erentiation are poorly
understood. Indeed, as mentioned above, the great progress made toward elucidating
the detailed molecular mechanism of sex determination in D. melanogaster is primarily
based on the presence of numerous mutant stocks that exhibit intersexual phenotypes
(Schutt and Nöthiger, 2000 ). Since such mutant strains are absent or very rare in other
insects, current studies on sex determination basically rely on the cloning of homolo-
gous sequences to the sex-determining genes of D. melanogaster (e.g., Sievert et al.,
1997 , Oliveira et al., 2009 ; Suzuki et al., 2003 ; but see Beye et al., 2003 ). In particular,
sex reversals (i.e., intersexes) induced by symbiotic microorganisms are often repeatable
and controllable under laboratory conditions (Rigaud, 1997 ; Kageyama and Traut,
2004 ; Narita et al., 2007a ). Elucidation of the molecular mechanisms of such phe-
nomena may allow us to reveal the common mechanism of sex determination among
arthropods.
SUBPHYLUM (CLASS)
ORDER
Family Species Reference
HEXAPODA INSECTA
EPHEMEROPTERA
Baetidae Baetis bicaudatus Vance, 1996
Polymitarcyidae Ephoron leukon McCaff erty & Bloodgood, 1986
ODONATA
Libellulidae Calopteryx virgo Ris, 1929
Leucorrhinia intermedia Asahina, 1979
Nannophya pygmaea Inoue & Yoshida, 2008
Sympetrum eroticum Kagaya & Ukai, 1994
Sympetrum maculatum Wada, 2007
Sympetrum striolatum Torralba-Burrial & Ocharan, 2009
PLECOPTERA
Capniidae Capnia sequoia Nelson & Baumann, 1987
Leuctridae Leuctra digitata Klotzek, 1971
Leuctra fusca Klotzek, 1971; Aubert, 1958
Leuctra prima Aubert, 1958
Paraleuctra Ricker, 1965
Nemouridae Nemoura besametsa Nebeker & Gaufi n, 1966
Nemoura cinctipes Nebeker & Gaufi n, 1966
Prostoia besametsa Stark et al., 1986
Protonemura Sanchez-Ortega, 1992
Zapada cinctipes Stark et al., 1986
Notonemouridae Austronemoura chilena Illies, 1961
ORTHOPTERA
Acrididae Anacridium moestrum Potter, 1940
Camnula pellucida Paul, 1941; Friauf, 1947
(Continued )
Table 1. Occurrence of individuals having male and female morphologies in arthro-
pods
S. Narita et al. / Terrestrial Arthropod Reviews 3 (2010) 63–96 73
Chorthippus biguttulus Ebner, 1951; Oschmann, 1971
Chorthippus longicornis Karaman, 1959; Oschmann, 1971
Chorthippus montanus Bednarz, 1970
Chorthippus rammei Ebner, 1940
Chrysochraon dispar Brisout de Barneville, 1847, 1848
Euchorthippus pulvinatus Descamp, 1968
Locusta migratoria Joly, 1959; Verdier, 1960
Melanoplus adelogyrus Hubbell, 1932
Melanoplus diff erentialis Slifer, 1966; Slifer & King, 1967
Melanoplus fasciatus White & Rock, 1945
Melanoplus mexicanus Severin, 1943, 1955
Oedaleonotus phryneicus Hebard, 1919
Oedaleus inormatus Ritchie, 1978
Oxya velox Kimura, 1951
Pardalophora phoenicoptera Friauf, 1947
Podisma pedestris Baccetti, 1954; Dirsh, 1957
Podisma sapporoensis Natori, 1931
Schistocerca gregaria Dirsh, 1957; Pener, 1964;
Maeno & Tanaka, 2007
Schistocerca paranensis Morales Agacino, 1957
Sphingonotus oaerulans Dirsh, 1957
Trimerotropis oitrina x T. maritima Carothers, 1939
Valanga irregularis White, 1968
Ephippigeridae Ephippiger ephippiger Dumortier, 1962
Gryllidae ( Gryllus argentinus x capitatus ) x
G. capitatus
Cousin, 1967
( Gryllus bimaculatus x campestris ) x
G. bimaculatus
Cousin, 1935, 1937
Gryllus bimaculatus Johnstone, 1975
Gryllus bimaculatus x capitatus Cousin, 1963
Gryllus lineaticeps Chopard, 1955
Homoeogryllus japonicus Ohmachi, 1929, 1932; Suzuki,
1933, 1934
Phaneropteridae Isophya pyrenaea Dumontier & Paly, 1971
Rhaphidophoridae Dolichopoda linderi Boudou-Saltet, 1968
Tetrigidae Paratettix texanus Robertson, 1936
Tettigoniidae Amblycorypha oblongifolia Pearson, 1927, 1929
Amblycorypha rotundifolia Pearson, 1927, 1929
Aoridopeza reticulata Agar, 1940
Barbistes conetrictus Chladek, 1968
Barbistes yersini Brunner von Wattenwyl, 1876
Decticus albifrons Boudou-Saltet, 1975
SUBPHYLUM (CLASS)
ORDER
Family Species Reference
(Continued )
Table 1. (Cont.,)
74 S. Narita et al. / Terrestrial Arthropod Reviews 3 (2010) 63–96
Decticus verrucivorus Ramme, 1951
Ephippiger ephippiger Dumortier, 1962
Ephippiger terrestris Kheil, 1914
Ephippiger vitium Pantel & de Sinety, 1908
Insara elegans Rehn & Hebard, 1914
Isophya modesta Kiss, 1960
Isophya pavelii Brunner von Wattenwyl, 1876
Isophya pyrenaea Dumortier & Paly, 1971
Leptophyes punctatissima Cappe de Baillon, 1924, 1932;
Carothers, 1939; Chopard, 1955
Leptophyes punotatissima Cappe de Baillon, 1924, 1932
Metrioptera brachyptera Cappe de Baillon, 1924; Ebner,
1940; Harz, 1960
Metrioptera brachyptera Cappe de Baillon, 1924
Microcentrum retinerve Nickle, 1983
Odontura sp. Chadima, 1872
Poecilimon elegans Ramme, 1926
Poecilimon orbeliscus Harz, 1967
Pycnogaster graellsi Pantel & de Sinety, 1908
Pycnogaster inermis Barranco et al., 1995
Tettigonia viridissima Klapalek, 1897
manotrizon fallax Ramme, 1913
PHASMATODEA
Bacillidae Clonopsis gallica Chopard, 1918
Diapheromeridae Carausius morosus Bergerard, 1958; Pijnacker &
Ferwerda, 1980
Phasmatidae Extatosoma tiaratum Rumbucher, 1974; Calberg, 1981
Phyllidae Heteropteryx dilatata Brock, 1989; Seow-Choen, 1995
Phyllium bioculatum Ziegler, 1989, 1995
Phyllium celebicum Grosser, 2003
BLATTODEA
Blaberidae Byrsotria fumigata Barth & Bell, 1971
Gromphadorhina portentosa Graves et al., 1986
Blattellidae Blattella germanica Ross & Cochran, 1967
HEMIPTERA
Cicadellidae Zyginidia pullula Negri et al., 2006
Cicadidae Melampsalta muta Dugdale & Fleming, 1966
COLEOPTERA
Carabidae Akimerus schaeff eri Auvray & Auvray, 1998
SUBPHYLUM (CLASS)
ORDER
Family Species Reference
(Continued )
Table 1. (Cont.,)
S. Narita et al. / Terrestrial Arthropod Reviews 3 (2010) 63–96 75
Carabus nemoralis Hartkorn, 1982
Pterostichus musahiensis Kashara & Karube, 1995
Cerambycidae Ergates faber Balazuc, 1952
Leptura rubra Weber, 1913
Rhagium mordax Starzyk, 1984
Curculionidae Euplatypus hintzi Beaver, 2000
Dynastidae Megasoma elephas Dechambre, 1987;
Blackaller-Bages &
Delgado-Castillo, 1990
Galerucidae Cerotoma facialis Ruppel, 1971
Lucanidae Lucanus elaphus Wickham, 1903
Melolonthidae Polyphylla fullo Vasko, 2008
Salpingidae Boros discicollis Spilman, 1953
Scarabaeidae Cotinis mutabilis Deuve, 1992
Dasylepida ishigakiensis Tanaka et al., 2006
Dicranocephalus wallichii Mizunuma, 2002
Goliathus cacicus Ture, 2001
Golofa tersander Ratcliff e, 1989
Polyphylla adspersa Vasko, 2008
Polyphylla fullo Vasko, 2008
Polyphylla olivieri Vasko, 2008
Polyphylla tridentata Vasko, 2008
Tenebrionidae Tribolium castaneum Sokoloff & Hoy, 1968
HYMENOPTERA
Agaonidae Blastophaga psenes Pereira et al., 2003
Pegoscapus tonduzi Pereira et al., 2003
Tetrapus sp. R.A.S. Pereira, unpublished data
Andrenidae Andrena chengtehensis Xu & Cui, 2007
Apidae Apis mellifera Rothenbuhler et al., 1952;
Witherell, 1971;
Milne & Rothenbuhler, 1983b;
Brockmann & Brückner, 1999
Nomada sp. Tsuneki, 1975
Xylocopa brasilianorum Gordh & Gulmahamad, 1975
Xylocopa nigrocincta Lucia et al., 2009
Chalcididae Hockeria rubra Halstead, 1992
Diprionidae Diprion similis Mertins & Coppel, 1971
Neodiprion sertifer Heliövaara et al., 1992
Encyrtidae Microterys ishiii Zhang & Zhu, 2007
Formicidae Cardiocondyla emeryi Heinze & Trenkle, 1997
Cardiocondyla kagutsuchi Yoshizawa et al., 2009
SUBPHYLUM (CLASS)
ORDER
Family Species Reference
(Continued )
Table 1. (Cont.,)
76 S. Narita et al. / Terrestrial Arthropod Reviews 3 (2010) 63–96
Monomorium pharaonis Berndt & Kremer, 1982
Pheidole dentata Jones & Phillips, 1985
Vollenhovia emeryi Kinomura & Yamauchi, 1994
Halictidae Evylaeus albipes Nilsson, 1987
Megalopta genalis Wcislo et al., 2004
Megachilidae Dianthidium sayi Hicks, 1926
Osmia pentstemonis Sandhouse, 1923
Pteromalidae Nasonia vitripennis Kamping et al., 2007
Sphecidae Psenulus concolor Schneider & Feitz, 2003
Psenulus laevigatus Schneider & Feitz, 2003
Tenthredinidae Pachynematus clitellatus Baker, 1996
Pteronidea ribesii Peacock, 1925
Trichogrammatidae Trichogramma cordubensis Pintureau et al., 1999
Trichogramma pretiosum Beserra et al., 2003
DIPTERA
Anthomyiidae Pseudonupedia intersecta Blackith & Blackith, 1991
Asilidae Nannocyrtopogon minutus Cooper, 1990
Ceratopogonidae Culicoides crepuscularis Smith & Perry, 1967
Culicoides haematopotus Smith & Perry, 1967
Culicoides lailae Naval, 1969
Culicoides stellifer Smith & Perry, 1967
Chironomidae Chironomus decorus Hubschman & Stack, 1992
Chironomus tentans Martin, 1995
Culicidae Aedes aegypti Antunes & Forattini, 1960
Aedes albopictus Barreto et al., 2008
Aedes canadensis Roth, 1948
Aedes cantans Campbell & Service, 1987
Aedes craggi Huang, 1974
Aedes detrius Roth, 1948
Aedes dorsalis Blakeslee et al., 1966
Aedes fi tchii Kardatzke, 1978
Aedes hendersoni Grimstad & DeFoliart, 1974
Aedes implacabilis Roth, 1948
Aedes mcintoshi Gargan et al. 1989
Aedes punctor Roth, 1948
Aedes taeniorhynchus Lum, 1960; Pratt & Sudia, 1964
Aedes togoi Chellappan, 1965; Takai & Tadano,
1995
Aedes triseriatus Ezenwa & Venard, 1973
Aedes vexans Minson, 1969; Oemick, 1976
Anopheles gambiae Mason, 1980
Armigerus subalbatus Reena & Ramakrishna, 1996
Culex erythrothorax Blakeslee & Rigby, 1965
Culex fuscocephalus Aslamkhan, 1970
SUBPHYLUM (CLASS)
ORDER
Family Species Reference
(Continued )
Table 1. (Cont.,)
S. Narita et al. / Terrestrial Arthropod Reviews 3 (2010) 63–96 77
Culex molestus Roth, 1948
Culex nigripalpus Roth, 1948; Meadows, 1966;
Barreto et al., 2008
Culex pedroi Barreto et al., 2008
Culex pipiens Roth, 1948;
Blazquez & Maier, 1951;
Ausat & Koshi, 1955;
Meadows, 1966; Seal, 1966;
Ahmad et al., 1985;
Ali & Rasheed, 2008
Culex pseudovishnui Aslamkhan & Reisen, 1979
Culex salinarius Roth, 1948; Meadows, 1966;
Hall, 1990
Culex tarsalis Rigby, 1966; Rosay, 1968;
Mitchell & Hughes, 1969;
Harmston, 1971
Culex tritaeniorhynchus Aslamkha & Baker, 1969
Culiseta inornata Benge, 1970
Culiseta melanura Zimmerman & Morris, 1978
Culiseta morsitans Howard et al., 2007
Culiseta novaezealandiae Dobrotworsky, 1972
Mansonia dyari Slaff & Nemjo, 1984
Orthopodomyia signifera Roth, 1948
Taeniorhynchus uniformis Laurence, 1959
eobaldia annulata Classey, 1942
eobaldia annulata Roth, 1948
Trichoprosopon digitatum Lee, 1967
Drosophilidae Drosophila melanogaster Mavor, 1924; Dobzhansky, 1930;
Gowen, 1942;
Bonnier et al., 1949;
Goldschmidt, 1949;
Bonnier & Lüning, 1951;
Cook, 1978
Drosophila pseudoobscura Dobzhansky & Spassky, 1941
Drosophila subobscura Hollingsworth,1955; 1960
Drosophila virilis Lebedeff , 1939
Limoniidae Dicranomyia mitis Geiger, 1983
Muscidae Hydrotaea zao Iwasa & Shinonaga, 1982
Musca autumnalis Cilek & Knapp, 1994
Musca domestica Rubini et al., 1980
Polietina orbitalis Nihei & Carvalho, 2002
Mycetophilidae Sciara ocellaris Mori & Perondini, 1980, 1984
Psychodidae Lutzomyia davisi de Souza et al., 2008
Lutzomyia rorotaensis Geoff rey, 1984
Phlebotomus perniciosus Gállego et al., 1994
SUBPHYLUM (CLASS)
ORDER
Family Species Reference
(Continued )
Table 1. (Cont.,)
78 S. Narita et al. / Terrestrial Arthropod Reviews 3 (2010) 63–96
Sergentomyia minuta Gallego et al., 1991
Sergentomyia minuta parroti Harrat et al., 1993; Addadi &
Dedet, 1977
Sarcophagidae Parasarcophaga harpax Kurahashi, 1977
Simuliidae Austrosimulium dumbletoni Craig & Crosby, 2008.
Austrosimulium australense Crosby 1973
Simulium aokii Hadi & Takaoka, 1993;
Hadi et al., 1994
Simulium arakawae Hadi & Takaoka, 1993
Simulium asakoae Fukuda et al., 2004
Simulium auricoma Grenier & Bertrand, 1950.
Simulium bidentatum Hadi & Takaoka, 1993;
Hadi et al., 1994
Simulium damnosum Cheke & Garms, 1985
Simulium soubrense Dang & Peterson, 1979
SIPHONAPTERA
Ceratophyllidae Nosopsyllus henleyi Beaucournu & Launay, 1987
LEPIDOPTERA
Adelidae Nemophora rubrofascia Hirowatari, 2005
Lemyra imparilis Yamauchi, 2000
Spilosoma mendica Bocklet, 1909
Bombycidae Bombyx mori Goldschmidt & Katsuki, 1927,
1931
Crambidae Hedylepta accepta Riotte, 1979
Ostrinia furnacalis Sakamoto et al., 2007
Ostrinia scapulalis Kageyama et al., 2003
Geometridae Bupalus piniarius Kusnezov, 1926
Paleacrita vernata Muller, 1968
Phaeoura mexicanaria Blanchard, 1969
yrinteina arnobia Bernardino et al., 2007
Lasiocampidae Malacosoma neutria Kusnezov, 1926
Lycaenidae Lycaeides argyrognomon Sashida, 1977
Lycaena argus Kusnezov, 1926
Neozephyrus taxila Tochikura, 1971
Strymon bazochii Riotte, 1979
Lymantriidae Lymantria dispar Goldschmidt, 1934
Noctuidae Agrotis ipsilon Gemeno et al, 1998
Agrotis segetum Blair, 1976
Helicoverpa armigera Josephrajkumar et al., 1998
Nymphalidae Argynnis paphia Tennent, 2006
Epinephele tithonus Bocklet, 1908
Hyponephele narica Ivinskis & Saldaitis, 2001
Ladoga camilla Beccaloni, 1988
SUBPHYLUM (CLASS)
ORDER
Family Species Reference
(Continued )
Table 1. (Cont.,)
S. Narita et al. / Terrestrial Arthropod Reviews 3 (2010) 63–96 79
Papilionidae Ornithoptera croesus Parrott & Schmid, 1984
Ornithoptera priumus Schmid, 1973
Ornithoptera victoriae Schmid, 1973
Papilio bianor Waki, 1976
Papilio glaucus Scriber & Evans, 1988
Papilio memnon Nimura & Otani, 2006
Papilio polyxenes Blau, 1978
Troides rhadamantusrhadamantus Yamada, 1979
Pieridae Colias erate poliographus Saitoh, 1955
Eurema mandarina Narita et al., 2007a,b
Gonepteryx cleopatra Winokur, 2002
Gonopteryx rhamni Kusnezov, 1926
Pieris melete Takahashi, 1981
Pieris napi Kamei, 1970
Pyralidae Galleria mellonella Smith, 1968
Saturniidae Aglia japonica Kobayashi, 1997
Antheraea mylitta Sen & Jolly, 1967
Antheraea yamamai Kawazoe, 1984
Arsenura armida Motta, 2000
Automeris io Manley, 1971
Hyalophora cecropia Bridgehouse, 2000
Periga circumstans Moraes, 2005
Satyridae Maniola jurtina Albrecht, 1993
Sphingidae Smerinthus ocellata Morgan, 1909
Smerinthus populi Morgan, 1909
Tortricidae Acleris celiana Hodges & Brown, 2007
Yponomeutidae Yponomeuta cagnagellus Kuijten, 1973
TRICHOPTERA
Hydroptilidae Hydroptila angustata Dia & Botosaneanu, 1982
Leptoceridae Oecetis ochracea Bochert & Bochert, 2005
Limnephilidae Anabolia furcata Mey, 1982
Psychomyiidae Psychomyia ctenophora Botosaneanu, 1995
CRUSTACEA (MALACOSTRACA)
AMPHIPODA
Gammaridae Gammarus chevreuxi Sexton, 1924
Gammarus duebeni Dunn et al., 1990; Kelly et al., 2004
Orchestia gammarellus Ginsburger-Vogel, 1975
DECAPODA
Coenobitidae Coenobita rugosus Gusev & Zabotin, 2007
Parastacidae Samastacus spinifrons Rudolph, 1995, 2002
Potamidae Geothelphusa dehaani Araki & Matsuura, 1995
Potamon fl uviatile Micheli, 1991
SUBPHYLUM (CLASS)
ORDER
Family Species Reference
(Continued )
Table 1. (Cont.,)
80 S. Narita et al. / Terrestrial Arthropod Reviews 3 (2010) 63–96
SUBPHYLUM (CLASS)
ORDER
Family Species Reference
Table 1. (Cont.,)
Sergestidae Acetes sibogae Hanamura & Ohtsuka, 2003
alassinidea Upogebia stellata Pinn et al., 2001
ISOPODA
Armadillidiidae Armadillidiun vulgare Juchault et al., 1980, 1991
Asellidae Asellus communis Smith, 1967
Idoteidae Idotea balthica Mocquard et al., 1978
MYSIDA
Mysidae Siriella japonica Ohtsuka et al., 2003
CRUSTACEA BRANCHIOPODA
ANOSTRACA
Artemiidae Artemia franciscana Campos-Rasmos et al., 2006
Artemia parthenogenetica Campos-Rasmos et al., 2006
Artemia salina Bowen & Hanson, 1962
Chirocephalidae Branchinecta lindahli Sassaman & Fugate, 1997
CLADOCERA
Daphniidae Daphnia longispina Bykova & Markevich, 1979
Daphnia magna Mitchell, 2001; Olmstead &
LeBlanc, 2007
CHELICERATA (ARACHNIDA)
ACARI
Ixodidae Ixodes pacifi cus Keirans & Lane, 1997
Amblyomma cajennense Labruna et al., 2002
Amblyomma hebraeum Clarke & Rechav, 1992
Amblyomma neumanni Aguirre et al., 1999
Amblyomma oblongoguttatum Labruna et al., 2000
Amblyomma variegatum Stampfl i, 1985; Balde &
Konstantinov, 1994
Aponomma hydrosauri Chilton & Sharrad, 1992
Dermacentor andersoni Dergousoff & Chilton, 2007;
Homsher & Yunker, 1981
Dermacentor occidentalis Oliver & Delfi n, 1967
Hyalomma truncatum Kostrzewski et al., 1986; Clarke &
Rechav, 1993
Rhipicephalus appendiculatus Mwase et al., 1987
Rhipicephalus evertsi Clarke, 1991
Rhipicephalus sanguineus Labruna et al., 2002
Phytoseiidae Metaseiulus arboreus McMurtry & Show, 2007.
ARANEAE
Linyphiidae Centromerus prudens Roberts, 1976
Lycosidae Schizocosa sp. Stratton, 1995
eridiidae Episinus nubilus Kumada, 1989
(Continued )
S. Narita et al. / Terrestrial Arthropod Reviews 3 (2010) 63–96 81
SUBPHYLUM (CLASS)
ORDER
Family Species Reference
Table 1. (Cont.,)
Note: Not all the published records are shown.
OPILIONES
Gagrellidae Gagrellula montana Tsurusaki, 1982
Melanopa grandis Tsurusaki, 1982
Nemastomatidae Nemastoma triste aler, 2004
CHELICERATA (PYCNOGONIDA)
PANTOPODA
Ammotheidae Cilunculus armatus Miyazaki & Makioka, 1993
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... The distribution of male versus female attributes within the insect can be striking and clear, such as the expression of different sexual traits along a particular axis of the body, such as bilaterally asymmetrical individuals (e.g., Wcislo et al., 2004;Michez et al., 2009;Hinojosa-Díaz et al., 2012). In the few systems where such a phenomenon has been investigated, they are the result of chimeric intermingling of genetically male and female tissues within a single individual (Narita et al., 2010). This may result from a variety of developmental aberrations or, in some hymenopteran species with Wolbachiainduced parthenogenesis, higher temperatures partially suppress the action of the bacterial symbiont and result in the formation of gynandromorphs (e.g., Bowen and Stern, 1966;Cabello-García and Vargas-Piqueras, 1985). ...
... Indeed, in the cases of those individuals reported as mixed gynandromorphs, whereby the distribution of male versus female traits are not clearly separated along some axis and are instead intermingled, it remains to be determined whether or not they are genetically intersexes rather than true gynandromorphs. Nonetheless, gynandromorphs have been reported in 15 Orders of insects with dramatically different underlying sex-determination systems, namely: Ephemeroptera, Odonata, Plecoptera, Orthoptera, Phasmatoe, Dermaptera, Blattaria, Hemiptera, Psocoptera, Coleoptera, Hymenoptera, Trichoptera, Lepidoptera, Diptera, Siphonaptera (Cui and Cai, 2003;Narita et al., 2010) and now Zoraptera, the 16th Order. ...
... Gynandromorphs have the potential to inform us about the underlying sex determining mechanisms in certain insects (Narita et al., 2010), and can also be used to refine hypotheses of homology between sexually dimorphic structures such as the genitalia (e.g., Michener, 1943;Engel, 2007). ...
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The first gynandromorph of the insect order Zoraptera is reported. A gynandromorph of Zorotypus brasiliensis Silvestri is described from a likely parthenogenetic population in the Atlantic Forest around the border of the Brazilian states of Minas Gerais, Rio de Janeiro, and São Paulo. As has been previously surmised for this species, the sample consisted of only females, tending to support the hypothesis of thelytokous parthenogenesis for this population. The gynandromorph is largely female in outward appearance, but exhibits male secondary sexual traits on the left side of the apical abdominal segments. The left (male) antenna consists of antennomeres of different proportions to those of the right (female) antenna. The internal genitalia and associated sclerites, however, are female although some are augmented in their form relative to normal-type females. Comments are made on possible factors resulting in the development of the gynandromorph. This is the 16th insect order with gynandromorphism reported.
... For instance, populations of the house fly Musca domestica carry several male-determining chromosomes and an additional female sex-determination locus (Dübendorfer et al. 2002;Hediger et al. 2010). Intersex individuals that express both male and female traits spontaneously arise at low frequencies in natural populations of certain arthropod species (Narita et al. 2010). In isopods, the appearance of intersex individuals is often linked to infection of Wolbachia, probably one of the most widespread invertebrate-associated bacteria (Bouchon et al. 1998(Bouchon et al. , 2009Engelstädter and Hurst 2009;Cordaux et al. 2011Cordaux et al. , 2012. ...
... Intersex individuals are rarely observed in natural populations of arthropods (Narita et al. 2010). The intersex phenotype of the current study appears to be present at an extremely low frequency in natural populations of P. scaber. ...
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Intersex phenotypes are rarely observed in natural isopod populations and their expression is typically associated with infection of Wolbachia , a reproductive parasite that manipulates arthropod reproduction. During an intensive sampling effort of a natural population of the isopod Porcellio scaber , an adult individual was isolated that expressed both male and female traits. The intersex individual exhibited clearly developed external male genitalia and carried multiple eggs in its brood pouch. No Wolbachia infection could be identified in this individual, a result that needs to be approached with caution due to suboptimal DNA preservation for diagnostic PCR assays. Wolbachia were, however, detected in two adult females of the same population, and appear closely related to isolates that infect other terrestrial isopod species. This is the first demonstration that intersex phenotypes can arise under natural conditions in P. scaber .
... " 3 Mitchell used the terms "intersex" and "gynandromorph" to refer to different anomalies, and this distinction is still in place today. The difference is now defined genetically, such that intersexes are genetically uniform but express some physical characteristics of the other sex, while gynandromorphs have genetically distinct male and female tissues (Mitchell 1928, Narita et al. 2010). Mitchell described and interpreted these specimens for his dissertation and in a 1928 paper, "Sex Anomalies in the Genus Megachile." ...
... Although this possibility has never been confirmed, it remains a plausible hypothesis. Along with other genetic and environmental factors, parasitism is one of the known causes of intersexual phenotypes in other arthropods (Narita et al. 2010). ...
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“ Each species must be known by one name only, throughout the entire extent of its range. This systematic study is the foundation upon which all the other work of seemingly more practical importance is based ” [T. B. Mitchell (1946)][1] [1]: #ref-10
... In separate-sex animals, abnormal individuals with a mix of phenotypically male and phenotypically female anatomical parts are known as gynandromorphs. These are usually described as genetic or chimeric mosaics that consist of both male and female cells (e.g., [1,2]). Gynandromorphism can occur when sexual differentiation is not regulated at a systemic level and, depending on the species' developmental cell lineages and the stage of development in which the anomaly occurs, the distribution of "male tissues" and "female tissues" can exhibit some kind of symmetry (e.g., bilateral) or produce a patchwork throughout the body. ...
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Gynandromorphs, i.e., individuals with a mix of male and female traits, are common in the wild bees of the genus Megachile (Hymenoptera, Apoidea). We described new transverse gynandromorphs in Megachile pilidens Alfkeen, 1924 and analyze the spatial distribution of body parts with male vs. female phenotype hitherto recorded in the transverse gynandromorphs of the genus Megachile. We identified 10 different arrangements, nine of which are minor variants of a very general pattern, with a combination of male and female traits largely shared by the gynandromorphs recorded in 20 out of 21 Megachile species in our dataset. Based on the recurrence of the same gynandromorph pattern, the current knowledge on sex determination and sex differentiation in the honey bee, and the results of recent gene-knockdown experiments in these insects, we suggest that these composite phenotypes are possibly epigenetic, rather than genetic, mosaics, with individual body parts of either male or female phenotype according to the locally expressed product of the alternative splicing of sex-determining gene transcripts.
... Il est à noter qu'aucune étude génétique, physiologique ou cytologique n'a été effectuée sur le spécimen afin d'en comprendre sa génèse et de déterminer à coup sûr s'il s'agit d'un gynandromorphe au sens strict ou d'un intersexué au sens de Narita et al. (2010). Toutefois, un tel individu présentant des caractères à la fois mâle et femelle ne nous semble pas avoir été signalé dans la famille des Trictenotomidae. ...
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We present here the first case of gynandromorphism in the beetles family Trictenotomidae. The specimen belongs to the species Autocrates vitalisi Vuillet, 1912 and demonstrates male and female mixed characteristics distributed in a complex manner, not as in the case of a strict bilateral gynandromorphism. Therefore, this individual could be better defined as an example of mosaic gynandromorphism.
... Sexual anomalies were first reported for bees in 1857, when Sichel described a gynandromorphic form of Bombus lapidarius (Linnaeus, 1758) ( Wcislo et al., 2004;Michez et al., 2009;Silveira et al., 2012). Gynandromorphs are individuals that have male and female traits simultaneously in the body, and the causes of this anomaly are discussed in relation to problems in fertilization, chromosome damages or loss, and association with symbionts and parasites (Wcislo et al., 2004;Michez et al., 2009;Narita et al 2016). More recently, only three categories started to be used by some mellitologists: bilateral, transverse and mosaic ( Michez et al., 2009;Giangarelli & Sofia, 2011;Vivallo, 2015;Ugajin et al., 2016). ...
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Gynandromorphism is the most common case of sexual anomaly reported in bees and is characterized by individuals that show male and female traits simultaneously in the body. Gynandromorphic cases have been reported for 140 species of bees, an underestimated number comparing to the twenty thousand bee species described nowadays. Here we describe and illustrate the first case of a gynandromorphic Xylocopa darwini Cockerell, 1926 and the fourth case of Xylocopa varipuncta Patton, 1879. The specimens show a mixed form of gynandromorphism with predominantly female features and with all its male traits concentrated in one side of the body, right side in X. darwini and left side in X. varipuncta. The gynanders of X. darwini and X. varipuncta were collected on Isabela Island Galapagos – Ecuador) and Riverside (California – USA), and were deposited in Smithsonian Collection and California Academy of Sciences, respectively. Including this work, eighteen cases of gynandromorphism were reported to Xylocopa and twelve were recorded from Neoxylocopa subgenus.
... Some intersexes are genetically intermediate between the typical male genotype and the typical female genotype. Other intersexes are genetically purely male or female, but some parts of their bodies having a sexual phenotype that is opposite to their genetic sex (Wigglesworth 1972, Narita et al. 2010, Yang & Abouheif 2011). In this paper, we describe and illustrate three new cases of bilateral gynandromorphism within the three ant species Leptothorax kutteri Buschinger, 1965, Polyergus rufescens (Latreille, 1798) and Formica lugubris Zetterstedt, 1838. ...
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2016. Records of bilateral gynandromorphism in three species of ants (Hymenoptera, Formicidae) in Norway. Norwegian Journal of Entomology 63, 65–70. This paper describes and illustrates three new cases of gynandromorphism in three ant species, Leptothorax kutteri Buschinger, 1965, Polyergus rufescens (Latreille, 1798) and Formica lugubris Zetterstedt, 1838. All specimens described have bilateral asymmetry, in which one side is male and one is female. Gynandromorphism is in general a very rare phenomenon. The specimens of P. rufescens and F. lugubris were collected from exceptionally warm microhabitats, a fact that might affect the probability for gynandromorphism to appear.
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A gynandromorph is an organism combining both female- and male-type tissues. While the vast majority of studies have described the morphology of gynandromorphs, their sexual behaviours remain under-explored. We studied a predominantly bilateral gynandromorph specimen of the predatory wasp Discoelius dufourii (Hymenoptera: Vespidae: Zethinae) reared from a trap-nest. In addition to describing the morphology, we explored the gynandromorph’s response to a conspecific female, comparing it with the behavioural interactions between normal males and females of D. dufourii and the phylogenetically related D. zonalis . For most specific behavioural elements, males of both Discoelius species exhibited a greater frequency and a higher probability of occurrences of the behaviour than did females. However, aggression and plopping behaviours were prevalent in females. The gynandromorphic specimen demonstrated behavioural activities similar to those of a male, including a relatively frequent mate body antennation and an attempt to mount a female. However, it approached the female less frequently in comparison to conspecific normal males, and it did not demonstrate substrate antennation and jumping, typical of Discoelius male mating behaviour. Based on the parameters of the nest cell, the probability of the development of the gynandromorphic specimen from an unfertilized (male) egg was higher than the probability of development from a fertilized (female) egg.
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Two new records of gynandromorphs in Xylocopa Latreille, 1802 (Hymenoptera, Apidae). Gynandromorphs are deviant morphological individuals with genetically distinct male and female tissues. Records of sex anomalies seems to be important to better understand the mechanisms regulating phenotypic expression. Herein, two new cases of gynandromorphs in carpenter bee species of Xylocopa from Brazil are described and figured: a mixed gynandromorph of the X. (Neoxylocopa) brasilianorum (Linnaeus, 1767) from São Paulo and a bilateral gynandromorph of the X. (Neoxylocopa) ordinaria Smith, 1874 from Sergipe.
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Gynandromorphy, which is characterized by the coexistence of male and female tissues in a single individual, is known in insects. Gynandromorphs exhibit diverse levels of defects in sexual behavior. The distribution pattern of both sexes within the nervous systems could be responsible for these differences in behavioral traits; however, most studies have mainly focused on the unique external morphological traits of gynandromorphs, and little attention has been paid to the evaluation of nervous systems, at least in Hymenoptera. In this study, we evaluated a gynandromorphic individual of bumblebee (Bombus ignitus) with a bilaterally dimorphic head and thorax (left side, male; right side, female) and a uniformly masculine gaster. The gynandromorph exhibited abnormal sexual behavior; it normally approached and touched virgin queens but rarely made subsequent copulation attempt. Dissection of the gaster indicated that it possessed a set of male reproductive organs. We analyzed the encephalic and antennal expression patterns of the fruitless (fru) gene, which shows sex-specific alternative splicing conserved in various insect orders and has been thought to act as the master regulator of sexual behavior in male nervous systems. The gynandromorph showed left-side-biased expression of male-type fru transcripts. We discuss a possible mechanism for the observed unusual interruption of sexual behavior of the gynandromorph by reference to the fru expression pattern and our recent findings on the sexually dimorphic response to female-derived pheromones.
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Reproductive behaviour was studied in 192 gynandromorphs with female genitalia and reproductive system, produced by ring-X chromosome loss. Male and female behaviour patterns were frequently found to coexist in the same individuals, and male courtship behaviour, when it occurred, retained its characteristic hierarchical organisation. Sexually receptive individuals were found to be an almost perfect subset of those ovipositing, and the control of both of these behaviours mapped to the head, as did male orientation (courtship). High rates of wing flicking, a response of males to courtship, mapped rather to the thorax, although a quantitative analysis demonstrated that the frequency of flicking behaviour was also influenced by male tissue in the head. In non-ovipositing individuals mature oocytes were retained in the ovary. An egg held in the uterus is not deposited by a fly without female tissue in the head and all but one sexually receptive individuals laid eggs. It is therefore concluded that both of these behaviours depend upon closely related neural circuitry operating the genital musculature under control from the brain.
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Six gynandromorphic individuals of Neodiprion sertifer (Geoffrey) are described from Finland. They were bilaterally gynandromorphic, mosaic-like gynandromorphic or only showed such characters in their head or abdomen.