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1.
The results of recent studies by our group have suggested that in social wasps of the genus Polistes the Dufour’s gland is involved in kin recognition. In fact, the same hydrocarbons occurring on the cuticle are found in the
gland secretion, and in P. dominulus the composition of the glandular secretion is more similar in foundresses belonging to the same colonies than in heterocolonial
foundresses. In this study, P. dominulus colonies were experimentally presented with previously treated dead conspecific females. These lures had been deprived of
their epicuticular lipids and coated with epicuticular or Dufour’s gland secretion extracts from females hetero- or homocolonial
with respect to the tested colonies. The behaviour of the colonies towards these lures indicates that, like the epicuticular
lipids, the Dufour’s gland secretion is involved in nestmate recognition.
Received: 14 August 1995/Accepted after revision: 5 November 1995 相似文献
2.
Klaus Peschke Peter Friedrich Uwe Kaiser Stephan Franke Wittko Francke 《Chemoecology》1999,9(2):47-54
Summary. In the rove beetle Aleochara curtula, a male specific sternal gland is described. Isopropyl (Z9)-hexadecenoate has been identified by GC/MS of surface and sternal gland extracts as a male specific compound. Its emission
in the air was demonstrated by closed-loopstripping-analyses. In field experiments, conspecific males and females were attracted
by the odor of carrion, caged males from laboratory cultures, and by the synthetic ester. Isopropyl (Z9)-hexadecenoate or live males, combined with fresh carrion attract more beetles than a fresh carcass alone. Isopropyl hexadecanoate
as a minor compound was not attractive. The term “aggregation” pheromone and the ecological significance of attracting females
to a fresh carcass allowing early copulation and egg-deposition are discussed.
Received 20 August 1998; accepted 15 January 1999. 相似文献
3.
Sexually deceptive orchids of the genus Ophrys attract male insects for pollination. Pollinator attraction is achieved by mimicking sex pheromones of virgin females of
their pollinators, mostly bee species. In earlier investigations, we showed that the phylogenetically distinct Ophrys species O.
chestermanii and O. normanii on Sardinia attract their pollinator, males of the cuckoo bumblebee B. vestalis, with the same bouquets of relatively polar volatile compounds. In this investigation, we studied the sex pheromone of virgin
females of B. vestalis with the aim of identifying male-attracting compounds and of comparing them with labellum extracts of the two orchids, which
were found to release male-attracting compounds in earlier investigations (G?gler et al. 2009). In bioassays, shock-frozen females, cuticle extracts and polar fractions of cuticle extracts of virgin females stimulated
mating behaviour in the males. Using gas chromatography coupled with electroantennography (GC-EAD) and gas chromatography
coupled with mass spectrometry (GC-MS), we detected in polar fractions of cuticle extracts of B. vestalis females the same electrophysiologically active compounds as in labellum extracts of both orchid species, including aldehydes,
esters, fatty acids and alcohols. Since statistical comparisons of the relative proportions of esters showed strong similarities
between virgin females and orchids, our results support the hypotheses that this highly specialized Ophrys–pollinator relationship represents another case of chemical mimicry and that esters play a key role in male attraction. 相似文献
4.
Alain Lenoir Am��lie Benoist Abraham Hefetz Wittko Francke Xim Cerd�� Rapha?l Boulay 《Chemoecology》2011,21(2):83-88
In the course of our studies on the chemical ecology of the widely distributed Mediterranean ant Aphaenogaster senilis, we found that trail following is triggered by extracts of the poison gland and Dufour’s gland. To assess the specificity
of the trail pheromone, we examined whether a cross-reaction exists between trails of A. senilis and the closely related species A. iberica. Specificity seemed to differ amongst these two species, because workers of A. senilis did not follow trails of A. iberica, whereas the latter followed trails made by both species. Chemical analyses of the glandular contents reveal that Dufour’s
glands of both species contain mainly alkanes and alkenes exhibiting species-specific profiles. However, differences in the
poison gland content of the two species were dramatic, with A. senilis showing high amounts of alkaloids that were completely absent in A. iberica. 相似文献
5.
Summary Copulation in Ischnura graellsii may be divided into three stages, according to the movements and position of the male's abdomen. We measured sperm volumes in males and females interrupted at different phases of copulation in laboratory-reared and field specimens. The results showed that males remove sperm from the female during stage I, and do not transfer sperm until stage Il of the copulation. In the field females interrupted during stage I of copulation had less sperm than postcopula females, and the volume of sperm in laboratory females mated once or twice was similar. These results suggest that males can remove most of the sperm during stage I of copulation. Preparations of in-copula specimens showed the horns of the penis (used to remove sperm) inside the bursa copulatoox and the spermatheca. Therefore males can remove sperm from both organs, in contrast to the other Ischnura species so far studied, where males can empty only the bursa. The length of these horns is positively correlated with male body length, and there are significant differences in length between the left and right horns of individual males. This suggests great variability in the male's ability to remove sperm. On the other hand, ejaculate volume is positively related to male and female size, and negatively to male age. Males are likely to be able to detect the presence of sperm in females: if the effect of population density and time of start of copulation are taken into account, copulations are longer with mated than with virgin females. Using genetic markers, sperm precedence was studied by rearing the female offspring of 6 females mated with two males of different genotype. In 5 out of 6 crosses, the second male fertilized all the eggs laid by the female in her first clutch. On average, the following clutches were progressively more fertilized by the first male, but there were striking differences between crosses. These differences are probably due to the variability in the amount of sperm transferred and/or removed.
Offprint requests to: A. Cordero 相似文献
6.
Summary. This work validates a method for detecting potential semiochemicals in mouse urine samples with a volume as small as 10 μL. Using solid-phase microextraction, gas chromatography and mass spectrometry, we screened 2,3,5-trithiahexane, 2-sec-butyl-4,5-dihydrothiazole, geraniol, indole, trans-β-farnesene and farnesol in individual urine samples taken daily from mice housed under various social conditions. Excretion of 2-sec-butyl-4,5-dihydrothiazole by males did not occur when they were housed in rooms containing no females, but increased when raising the males to rooms containing females. Between-male differences in β-farnesene excretion were observed even in the absence of females. These results highlight the usefulness of the proposed analytical method for research in this area. 相似文献
7.
Mating in the red-sided garter snake,Thamnophis sirtalis parietalis: differential effects on male and female sexual behavior 总被引:1,自引:0,他引:1
Joan M. Whittier Robert T. Mason David Crews 《Behavioral ecology and sociobiology》1985,16(3):257-261
Summary Female red-sided garter snakes, Thamnophis sirtalis parietalis, become unattractive to most males after mating in the field and in the laboratory. Male red-sided garter snakes vary in their latencies to court attractive females following copulation, with courtship resuming in minutes to hours. Unsuccessful males in mating balls disperse from mating pairs, but are not residually inhibited from courting attractive females. These patterns of behavior indicate that males have evolved mechanisms to maximize opportunities for copulation with several females, while females mate only once per season. 相似文献
8.
Johannes Stökl Robert Twele Dirk H. Erdmann Wittko Francke Manfred Ayasse 《Chemoecology》2007,17(4):231-233
Summary.
Ophrys flowers mimic the female produced sex pheromone of their pollinator species to attract males for pollination. The males try
to copulate with the putative female and thereby pollinate the flower. Using electrophysiological and chemical analyses, floral
volatiles released by O. iricolor as well as the female sex pheromone of its pollinator species, Andrena morio are investigated. Overall, 38 peaks comprising 41 chemical compounds, were found to release reactions in the antennae of
male A. morio bees. Analyses using coupled gas chromatography-mass spectrometry revealed the presence of alkanes and alkenes with 20 to
29 carbon atoms, aldehydes (C9 to C24) and two esters. Almost all of those compounds were found in similar proportions in
both, the floral extracts of O. iricolor and cuticle surface extracts of A. morio females. The pattern of biologically active volatiles described here is very similar to that used by other Ophrys species pollinated by Andrena males. 相似文献
9.
Kathryn B. McNamara Therésa M. Jones Mark A. Elgar 《Behavioral ecology and sociobiology》2007,61(8):1177-1184
Male copulation experience may have a profound impact on female reproductive success if male reproductive investment declines
over consecutive copulations and if females are unlikely to re-mate. Male reproductive investment is particularly interesting
in lepidopterans because males produce dimorphic sperm: a fertilizing (eupyrene) and a non-fertilising (apyrene) sperm. In
two experiments, we explored the lifetime reproductive investment of male almond moths, Cadra cautella (also known as Ephestia cautella) and examined its influence on female reproductive success. In the almond moth, females re-mate infrequently and males transfer
sperm in a spermatophore. Attached to the spermatophore is a large chitinous process, the function of which is unknown. One
group of males were permitted consecutive copulations with virgin females and the amount of sperm and size of the spermatophore
transferred were compared for all females. We found that the number of both eupyrene and apyrene sperm per ejaculate decreased
with his increased mating frequency, while the size of the spermatophore process decreased dramatically after the male’s first
copulation. In a second experiment, we allowed males to mate with females throughout their lives and then compared female
fecundity and fertilisation success. We found no obvious decrease in female fecundity and fertilisation success with increased
male copulation experience, despite the likely reduction in male gametic investment. We discuss potential explanations for
the development of this enlarged and elaborate first spermatophore of male almond moths given that it confers no clear fitness
advantage to females. 相似文献
10.
Gas chromatography with simultaneous flame ionisation and electroantennographic detection (GC–EAD) and gas chromatography
with mass spectrometry analysis (GC–MS) of abdominal extracts of adult male Dermestes haemorrhoidalis Kuster (Coleoptera: Dermestidae) revealed the presence of electrophysiologically and behaviourally active compounds to its
conspecific males and females. Isopropyl dodecanoate (3), isopropyl (Z)-9-tetradecenoate (5), isopropyl tetradecanoate (6), isopropyl (Z)-9-hexadecenoate (7) and isopropyl hexadecanoate (8) were detected in male abdominal extracts only. Analysis of collected
male headspace volatiles revealed the presence of six EAD-active compounds (3), (5), (6) and isopropyl tridecanoate (4) plus
two unidentified compounds (1) and (9). Synthetic compounds (3), (4), (5), (6) and (7) showed EAD activity with antennae of
both sexes in contrast to synthetic (8) which showed EAD activity with female antennae only. Male and female antennae of D.
haemorrhoidalis reacted with high receptor potentials to isopropyl (Z)-9-dodecenoate (2), although this compound itself was detected in neither male nor female abdominal extracts or headspace
volatiles. Petri dish bioassays indicated that male abdominal extracts and compounds (2), (3), (5) and (6) aroused and attracted
conspecific male and female beetles significantly (P < 0.05) compared to female extracts. These results suggested the presence of a male-produced aggregation pheromone in D. haemorrhoidalis. Field assays with any of the described compounds, however, did not result in attraction of this beetle in significant numbers. 相似文献
11.
Summary. The basic chemical structure of the sex
pheromone of the pine sawfly Microdiprion pallipes
(Fallén) has earlier been identified as the propionate
ester of (2S,3S,7R/S,11R/S)/(2R,3R,7R/S,11R/S)-3,7,11-
trimethyl-2-tridecanol. We now report the results from
further investigations on the male response to individual
stereoisomers and to blends of stereoisomers, both in electroantennographic
(EAG) recordings and in field trapping
experiments. We also present our attempts to determine the
stereochemistry of the compounds present in females of
M. pallipes. By comparing gas chromatograms and mass
spectra obtained from natural extracts with those from synthetic
compounds it was found that the females contain one
or more of the four (2S,3S,7R/S,11R/S)-3,7,11-trimethyl-2-
tridecanol isomers (SS++-1). The active pheromone component
is the corresponding propionate ester 2. In EAG
experiments, males responded most strongly to five propionate
ester samples, namely two four-isomer blends: SS++-2
and SR++-2, and three individual stereoisomers: SSSR-,
SRRR- and SRSR-2. In a series of field trapping experiments
it was found that males were attracted to the SR++-2
four-isomer blend and to the individual isomer SSSR-2.
Based on the EAG-recordings and field responses of males
and the stereoisomers found in the females, we suggest that
the propionate ester of (2S,3S,7S,11R)-3,7,11-trimethyl-2-
tridecanol (SSSR-2) is used as a main component of the sex
pheromone in M. pallipes. Apparently the males react to
other stereoisomers in addition to that or those produced by
the females. 相似文献
12.
Mixed reproductive strategy and mate guarding in a semi-colonial passerine,the swallow Hirundo rustica 总被引:6,自引:0,他引:6
Anders Pape Møller 《Behavioral ecology and sociobiology》1985,17(4):401-408
Summary Both male and female swallows Hirundo rustica have a mixed reproductive strategy (parental care for offspring and extra-pair couplations). Mate guarding protects females from male harassment and male swallows from being cuckolded. Females hide their fertile period by copulating successfully with their mates for an extended period during first clutches. Males guard in the pre-fertile period, when many unpaired males are present. Early breeding swallows guard more than late breeders since more sexual chases of females by non-mate males take place in the early period. Solitarily breeding females experience few chases by strange males; copulation frequency, length of copulation period and mate guarding is adjusted to a lower level than among colonial birds. Male guarding activity is more intense in the fertile than in the pre-fertile period. Guarding reduces success of extra-pair copulation attempts.Three female swallows each paired and copulated with a single male and later changed to a new male before starting to breed. Extra-pair copulations most often take place between neighbouring swallows in the fertile period of the female. Many old, early breeding males and many young, late breeding females participate in extra-pair couplations. Successful extra-pair copulations peak in the fertile period contrary to success of pair copulations which does not change during the copulation period. 相似文献
13.
Manfred Ayasse Wolf Engels Gunter Lübke Timo Taghizadeh Wittko Francke 《Behavioral ecology and sociobiology》1999,45(2):95-106
The present paper reports on behavioral experiments and gas chromatographic analysis of chemical communication in the mating
biology of the primitively eusocial sweat bee Lasioglossum (Evylaeus) malachurum. In a dual-choice experiment, a female made odorless was significantly less attractive than an untreated one. Attraction in
L. (Evylaeus) malachurum is therefore mediated by a female-produced sex pheromone. Further bioassays showed that unmated gynes are more attractive
to males than mated ones. Males are able to differentiate between the two groups of females as little as 3 h after mating.
Biotests with different samples obtained from attractive gynes showed surface extracts to be most attractive. Behavioral tests
with synthetic copies of the compounds identified as cuticular constituents of virgin gynes were highly attractive to males;
the volatile bouquets consisting of n-alkanes, n-alkenes and iso- pentenyl esters of unsaturated fatty acids were the most
attractive samples. Isopentenyl esters of unsaturated fatty acids were the key compounds in inducing male inspections as well
as stimulating pounces and copulatory attempts. Virgin and nesting gynes differed clearly in the relative and absolute amounts
of the volatiles on the cuticle. The total amount of volatiles was significantly higher in virgin gynes and decreased in breeding
queens. Hydrocarbons were the dominant group of compounds in both groups of females. The relative amounts of the wax-type
ester, hexadecyl oleate, iso- pentenyl esters, and a hitherto unidentified steroid were higher in attractive virgin gynes,
while the relative proportions of hydrocarbons and lactones dominated in nesting queens. The site of sex pheromone production
in attractive young L. (Evylaeus) malachurum gynes remains unknown. Head glands or Dufour's gland secretions may be involved. Another possible source of the `active principle'
found among the cuticular lipids could be glandular cells of the epidermis. The significance of modulation of female sex pheromone
composition is discussed in terms of a reduction in mating expenditures.
Received: 30 April 1998 / Accepted after revision: 24 July 1998 相似文献
14.
Extrapair paternity in birds has been the focus of an increasing number of studies over the last decades. Nevertheless, there is little knowledge about extrapair copulation (EPC) behavior. In this study, we investigate whether female bluethroats (Luscinia s. svecica) with no extrapair offspring (EPO) in their broods really have been sexually monogamous or if they have copulated with extrapair males. We used an experimental approach to prevent transfer of sperm from the social male during copulation by fitting a rubber tube around the cloaca of randomly chosen males. If the females mated to these males do not participate in EPCs, they will produce infertile eggs, whereas females that copulate with extrapair males will only produce EPO. We found that 87% (n=15) of the experimental pairs compared to only 36% (n=51) of unmanipulated pairs produced EPO. Our result therefore suggests that females having no EPO may still have copulated with extrapair males and that promiscuity may be an obligate sexual strategy among bluethroat females.Communicated by S. Pruett-Jones 相似文献
15.
L. L. Wolf E. C. Waltz K. Wakeley D. Klockowski 《Behavioral ecology and sociobiology》1989,24(1):63-68
Summary In many odonates, females mate with more than one male while laying a single clutch of eggs. We studied paternity of eggs laid by remated females of Leucorrhinia intacta, a small libellulid dragonfly, at a pond near Syracuse, NY, USA. The probability of a female remating is a function of male density on the pond. The length of copulations differs considerably among males active on the study pond at the same time. Much of this variation was correlated with differences in mating tactics of the males; copulations by males that stayed on their territories during copulation were shorter than those by other males (Fig. 2). Eggs collected from females mated to irradiated, sterile males and to free-living, fertile males indicated that the average paternity expectation was higher for long than for short copulations, and that the variance in paternity expectation was lower for long than for short copulations. Some possible causes of the high variation in paternity at low copulation durations and possible reasons for differences in copulation duration between male mating tactics are discussed. 相似文献
16.
Sexual conflict in the snake den 总被引:3,自引:0,他引:3
Red-sided garter snakes (Thamnophis sirtalis parietalis) court and mate in spring, soon after they emerge from large communal overwintering dens in south-central Manitoba. Because
of a massive bias in the operational sex ratio, every female attracts intense courtship from dozens to hundreds of males.
We suggest that this courtship constitutes significant ”harassment,” because it delays the females’ dispersal from the den
and hence increases their vulnerability to predation. Small females may face the greatest costs, because they are less able
to escape from amorous males (who court all females, even juvenile animals). Our measurements show that males are stronger
and faster than females. Experimental trials confirm that the locomotor ability of females (especially small females) is greatly
reduced by the weight of a courting male. Arena trials show that intense courtship stimulates females to attempt to escape.
Remarkably, some females that are too small to produce offspring may nonetheless copulate. This precocious sexual receptivity
may benefit juvenile females because copulation renders them unattractive to males, and thus allows them to escape more easily
from the den. Female ”tactics” to escape male harassment may explain other puzzling aspects of garter snake biology including
size-assortative mating, temporal patterns in dispersal from the den, avoidance of communal dens by young-of-the-year snakes,
and female mimicry. Hence, sexual conflict may have influenced important features of the mating system and behavioral ecology
of these animals.
Received: 8 May 2000 / Revised: 28 July 2000 / Accepted: 30 July 2000 相似文献
17.
Summary. Male locust borers, Megacyllene robiniae
(Förster), responded to females only after contacting them
with their antennae, indicating that mate recognition was
mediated by a contact sex pheromone. GC-MS analyses of
whole-body extracts of males and females determined that the
profiles of compounds in the extracts were qualitatively similar,
but differed considerably in the ratios of compounds
between sexes. Biological activities of reconstructed blends
of the most abundant straight-chain (nC23, nC24, nC25, nC26),
methyl-branched (3me-C23, 3me-C25), and unsaturated
(Z9:C23, Z9:C25, Z9:C27 compounds in extracts from females
were assessed in arena bioassays, assessing four distinct steps
in the mating behavior sequence of males (orientation, arrestment,
body alignment, mounting and attempting to couple the
genitalia). Males were unresponsive to freeze-killed, solventwashed
females treated with blends of straight-chain and
methyl-branched alkanes, but responded strongly to females
treated with the blend of alkenes. Further trials determined
that the complete sequence of mating behaviors, up to and
including coupling the genitalia, was elicited by Z9:C25 alone.
Z9:C25 comprised 16.4 ± 1.3% of the total hydrocarbons in
whole-body hexane extracts of females and was co-dominant
with two other hydrocarbons that were not active. In contrast,
in solid phase microextraction (SPME) wipe samples from
several areas of the cuticle, Z9:C25 appeared as the single
dominant peak, comprising 34.6 – 37.8% of the sampled
hydrocarbons. Our data indicate that Z9:C25 is a contact sex
pheromone of M. robiniae, being the most abundant hydrocarbon
on the surface of the cuticular wax layer of females
where it is readily accessible to the antennae of males. 相似文献
18.
Volatiles from male hindgut extracts of a newly described spruce bark beetle, Ips shangrila Cognato and Sun, from different attack phases were analyzed by GC–MS/FID with both polar and enantioselective columns. The
GC–MS/FID analyses showed that unmated males (Phase 1) or males mated with <3 females (Phases 2–4) produced 2-methyl-3-buten-2-ol
and 99%-(+)-ipsdienol as major components, and (−)-cis-verbenol, (−)-trans-verbenol, myrtenol and 2-phenyl ethanol as minor or trace components. The release of these male-produced compounds was confirmed
by the analysis on aeration sample of an I. shangrila infested wind-thrown spruce trunk. The quantities of 2-methyl-3-buten-2-ol, cis-verbenol and trans-verbenol from male hindgut extracts were almost unchanged or even slightly increased during gallery development, while ipsdienol
decreased dramatically after mating with three females. No obvious Ips-related aggregation pheromone components were detected in the female hindgut extract. A field trapping bioassay in Qinghai,
China, showed that the ternary blends containing 2-methyl-3-buten-2-ol, (−)-cis-verbenol and 97%-(+)-ipsdienol or (±)-ipsdienol, caught significantly more I. shangrila (♂:♀ = 1:2.14) than did the unbaited control. Replacing 97%-(+)-ipsdienol (close to the naturally produced enantiomeric ratio)
with (±)-ipsdienol in the ternary blend seemed to reduce trap catches by 50%, but the difference was not statistically significant.
Surprisingly, addition of (−)-trans-verbenol (at 0.2 mg/day) to the active ternary blends significantly reduced traps catches to the level not different from
the unbaited control. Our results suggest that the two major components, 2-methyl-3-buten-2-ol and 99%-(+)-ipsdienol, plus
a minor component, (−)-cis-verbenol, produced by fed males, are likely the aggregation pheromone components of I. shangrila. 相似文献
19.
Nina Wedell 《Behavioral ecology and sociobiology》2010,64(9):1385-1391
A cost of mating is common to both sexes but has predominantly been examined in females. In species where males provide resources
to females at copulation, male mating costs are expected to be high as nutrient provisioning enhancing female fecundity is
assumed to carry costs. In addition, males frequently court females prior to mating, which is known to carry survival costs
to both sexes. However, the magnitude and basis of variation in males’ mating costs remains largely unknown. Here, I examine
the effect of nutrient provisioning and courtship on male longevity across full-sib families in the paternally investing green-veined
white butterfly, Pieris napi. Copulating males suffered a survival cost as did courting males prevented from copulating, indicating the courtship component
of mating is costly. Male P. napi release aphrodisiacs during courtship to promote mating, indicating that these compounds may also be costly to produce. Contrary
to expectation, nutrient provisioning was not associated with reduced survival relative to males only allowed to court females,
although it is possible that this could be masked by the potentially elevated courtship rates of courting males relative to
mating males. Families differed in magnitude of reduced male survivorship, indicating a likely genetic basis to variation
in costs of courtship and copulation. Male weight was unrelated to longevity and mating success, whereas longevity strongly
influenced male mating success, indicating lifespan is an important male fitness trait in this species. 相似文献
20.
X. Fu J. Tabata T. Takanashi S. Ohno S. Tatsuki Y. Ishikawa Y. Huang H. Honda 《Chemoecology》2004,14(3-4):175-180
Summary. The sex pheromone of Ostrinia orientalis
(Lepidoptera: Crambidae) was analyzed by gas chromatography–electroantennographic detection (GC–EAD), GC–mass spectrometry and a series of bioassays. Three EAD-active
compounds were detected in the female sex pheromone
gland extract, and identified as tetradecyl acetate (14:OAc),
(Z)-11-tetradecenyl acetate (Z11-14:OAc) and (E)-11-tetradecenyl acetate (E11-14:OAc). The titers (ratio) of 14:OAc, Z11-14:OAc and
E11-14:OAc in 3-day-old virgin
females were 0.49 ng (10), 4.86 ng (98) and 0.10 ng (2),
respectively. In a wind-tunnel bioassay, the 98:2 blend of
Z11- and E11-14:OAc, but not Z11-14:OAc alone, elicited
the same male behavioral responses as virgin females and
crude gland extracts. 14:OAc was inactive by itself, and did
not show any synergistic effect on the binary blend.
Field trapping experiments also confirmed the attractiveness
of the binary blend to O. orientalis
males. Based on
these results, we concluded that the sex pheromone of
O. orientalis
is a 98:2 mixture of Z11-14:OAc and
E11-14:OAc. This sex pheromone is very similar to that of the Z-type European corn borer,
O. nubilalis. The present
finding raises the question of whether O. orientalis
, which is indistinguishable from O. nubilalis
based on external morphology,
is a biologically distinct species independent from
O. nubilalis. 相似文献