Lack of morphological coevolution between male forelegs and female wings in Themira (Sepsidae: Diptera: Insecta)

The complex, species-specific foreleg armature in males of the genus Themira (Diptera: Sepsidae) provides an ideal system for testing competing hypotheses for the evolution of sexually dimorphic character divergence. In sepsid flies, the male holds onto the female by clasping her wing base with his modified forelegs. In the present study, we document the male leg and the female wing morphology using scanning electron microscopy and confocal microscopy. We use a phylogenetic tree for Themira to reconstruct male foreleg and female wing evolution and demonstrate that the male legs have evolved elaborate structures with little or no corresponding changes in wing morphology. This lack of interspecific variation in female wings is not in agreement with the hypothesis of a morphological 'evolutionary arms race' between males and females. However, there is also no evidence for sex-specific wing differences in sensory organs on the wing base that may explain how females could assess males according to Eberhard's 'cryptic female choice' hypothesis. Finally, our study reveals the function of several novel morphological clasping structures and documents that the male foreleg characters in Themira are highly homoplastic. Male forelegs in two clades evolve considerably faster than in other species or clades. These two clades include Themira superba and Themira leachi , species that have some of the most dramatically modified forelegs known in Diptera.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 227–238.     

Grooming and pollen manipulation in bees (Apoidea): the nature and evolution of movements involving the foreleg

Three modes of self cleaning occur in insects: nibbling by the maxillae, scraping one structure by another in one direction only, and rubbing back and forth while the respective parts are in continuous contact. This paper describes a comprehensive and comparative account of this behaviour in bees, with special reference to the cleaning of or by the forelegs. Bees, like all Hymenoptera, clean various parts of the head, including the mouthparts and the antennae, with the forelegs. Lower Hymenoptera scrape each antenna with either foreleg; in the species of Aculeata that possess the antenna cleaner (strigil) on the foreleg, only the ipsilateral leg is used. The thoracic dorsum of most bees, as in many sphecoid wasps, is scraped in a forward direction by the middle leg; Triepeolus spp., however, use the hind leg, and the Anthophorinae the foreleg. Some beetles and lacewings clean their forelegs in the mouthparts by nibbling and scraping. Most higher Hymenoptera as a rule scrape the foreleg between the ipsilateral maxilla and the labium; bees, however, clamp the foreleg between the flexed ipsilateral middle leg and then scrape it. An evolution of this behaviour is postulated via several intermediate forms derived from original stepping movements. Halictidae and Andrenidae clamp the foreleg for scraping underneath the middle tibia, whereas all other bees nearly always clamp it underneath the middle basitarsus. Very similar movements are used in various species for transferring pollen, oil, or nest materials from the foreleg to the middle leg. It is argued that the original way of pollen carrying in bees must have been by filling the crop through direct eating or by scraping pollen off the foreleg between the ipsilateral maxilla and the labium. The latter movement is widespread among bees and is homologous to the normal foreleg cleaning in the mouthparts of most other Hymenoptera. The efficiency of this behaviour is enhanced in many lower bees by a comb on the galea, which is the homologue of a similar structure widespread among aculeate wasps. In higher bees, Apidae and Anthophoridae, the galeal comb is replaced by an equifunctional stipes comb. Many bees have neither of these types of maxillary combs.     

Bending for love: losses and gains of sexual dimorphisms are strictly correlated with changes in the mounting position of sepsid flies (Sepsidae: Diptera)

Background  

Sexually dimorphic structures ntribute the largest number of morphological differences between closely related insect species thus implying that these structures evolve fast and are involved in speciation. The current literature focuses on the selective forces that drive these changes, be it 'sexual conflict' or 'female choice'. However, there are only few studies examining the function of sexual dimorphisms and even fewer that investigate how functional changes influence dimorphisms. This is largely due to the paucity of taxa for which the morphology, behavior, and phylogenetic relationships for multiple species are known. Here we present such data for sepsid flies. Sepsids have starkly dimorphic forelegs whose function can be documented under laboratory conditions. We use data from 10 genes to reconstruct the phylogenetic relationships for 33 species and test whether mounting positions are correlated with the presence and absence of sexual dimorphisms in the forelegs.     

Contextual determinants of female-female mounting in laboratory rats

Female-female mounting is widespread among mammalian species, but little is known about the proximal function of this behaviour. While such mounting is often regarded as a 'masculine' trait, its widespread occurrence may indicate that it serves specific functions within the context of female-female social behaviour. We valuated female mounting behaviour in Long-Evans rats in standard observation chambers and in a seminatural enclosure. Under these conditions, we examined a number of potential factors that might influence mounting, including the oestrous cycle, social hierarchy, familiarity and male presence. The female's mounting was not influenced by her own oestrous cycle, but did vary with the oestrous cycle of the stimulus female. Socially dominant females mounted significantly more than subordinate females, and mounting by the dominant female was most frequent when the subordinate female was sexually receptive. Females mounted (and fought with) unfamiliar females significantly more than they did with familiar cagemates. Female-female mounting was dramatically reduced when males were present. Further testing showed that female mounting did not affect the induction of the progestational state of pregnancy, suggesting that female mounting does not function as a pseudomale behaviour that can substitute for genital stimulation provided by the male. Based on these data, female mounting does not appear to function as a sexual behaviour per se, but may serve as a form of female social behaviour related to maintenance of the female's social status within female groups. In this regard, the results of this study suggest that female mounting is part of the normal female's complex behavioural repertoire and does not necessarily reflect masculinization of some underlying neural substrate. Copyright 1999 The Association for the Study of Animal Behaviour.     

OIL FLOWERS AND OIL BEES: FURTHER EVIDENCE FOR POLLINATOR ADAPTATION

We examined foreleg length and body size variation in two species of oil-collecting bees (Rediviva; Melittidae) in southern Africa. Oil-collecting bees harvest oil from host flowers by rubbing their forelegs against oil-secreting trichomes. Significant differences in foreleg length occur among populations of both species. Rediviva “pallidula” populations vary significantly in mean foreleg length (11.34 ± 0.42 mm to 12.67 ± 0.36 mm), but not in body length (10.59 ± 0.74 to 10.80 ± 0.64), and foreleg length and body size are not significantly correlated. Instead, foreleg variation appears to be a function of host plant spur length. Ninety-two percent of the variance in foreleg length of R. “pallidula” is explained by mean Diascia spur length. Rediviva rufocincta populations vary significantly in mean foreleg length (10.12 ± 0.70 mm to 12.34 ± 0.68 mm) and in body length (9.03 ± 0.26 mm to 10.56 ± 0.24 mm). Foreleg length scales allometrically with body size in this species as 90.5% of the variance in foreleg length can be explained as a function of body length. Body size appears to be constrained by the morphology of the oil-secreting host plant. Both bees collect floral oil with specially modified setae on the tarsi of their forelegs. The length of the disti- + mediotarsus (refered to here as “tarsus”) in relation to the entire foreleg is shorter in R. rufocincta and does not increase as rapidly with increasing foreleg length as for R. “pallidula.” These differences in variation can be attributed to differences in position of oil within the flowers of the respective host plants. Rediviva “pallidula” collects oil from Diascia species that have the oil deeply situated in narrow floral spurs of varying length, while R. rufocincta collects oil from the broadly saccate flowers of Bowkeria verticillata and B. citrina.     

Territoriality and fighting in a colonial thrips, Hoplothrips pedicularius, and sexual dimorphism in Thysanoptera

ABSTRACT.

• 1 Hoplothrips pedicularius (Haliday) (Thysanoptera: Phlaeothripidae), a tubuliferan thrips in which males possess greatly enlarged forelegs, lives in colonies on Stereum fungus.
• 2 Females oviposit onto communal egg masses, and males fight by grasping and stabbing with their forelegs in territorial defence of oviposition areas. Prolonged escalated fights occur between males who are of similar size.
• 3 Larger males usually win fights and become dominant at the oviposition area. Dominant males secure 80% of matings, and mate most frequently during oviposition periods, with an ovipositing female.
• 4 Smaller, subordinate males avoid fights and attempt to 'sneak’copulations. However, they occasionally challenge the dominant male. Challenges tend to follow copulations by the subordinate male and occur more frequently between males who are of similar size.
• 5 Subordinate males who eventually leave the oviposition area are larger than those who remain, have frequently challenged the dominant male, and have more frequently been stabbed.
• 6 Sexual dimorphism in thrips is associated with gregariousness, claustral habitats, female-biased sex ratios, and male winglessness. In thrips genera in which males exhibit foreleg armature, males are larger relative to females. The ecological circumstances promoting sexual dimorphism and male fighting in spatially-structured populations are discussed.

     

Foreleg Autotomy Reduces Mating Success of Male <Emphasis Type="Italic">Schizocosa ocreata</Emphasis> Wolf Spiders

Autotomy is a taxonomically widespread antipredator tactic that allows animals to escape life-threatening situations. Opposing the benefits of survival, animals that have autotomized appendages may later suffer reduced ability in important determinants of fitness. Male Schizocosa ocreata wolf spiders use their forelegs during courtship for visual displays, for tactile courtship, and to defend against attacks by females. In nature they are often found missing one, and sometimes both, forelegs. We found that autotomy of one foreleg has little effect on male ability to mate with virgin females, but that autotomy of both forelegs causes a significant reduction in mating success. Among males that mated, autotomy of one or both forelegs did not influence latency until mating, period spent mounted, probability that his mate would accept a subsequent suitor, or probability that his mate would kill a subsequent suitor.     

Weaponry, color, and contest success in the jumping spider Lyssomanes viridis

Weaponry and color badges are commonly theorized to function as visual signals of aggressiveness or fighting ability. However, few studies have supported a signaling function of weaponry, and the role of color in invertebrate competitive interactions remains virtually unexplored. Jumping spiders (Salticidae) make excellent invertebrate models for studying weaponry and color because males of many species are colorful and possess exaggerated chelicerae, which are used as weapons in escalated contests. To determine whether color or weaponry might function as visual signals in male-male competitions, we investigated relationships between contest success, cheliceral length, and red coloration in Lyssomanes viridis. Males having longer chelicerae than their opponents were significantly more likely to win (p=0.0008). Males who won, despite being smaller than their opponents, had significantly less red chelicerae than their opponents (p=0.01). Male and female cheliceral length, as well as foreleg length, correlated tightly with body size. Cheliceral and foreleg length showed significantly stronger positive allometry in males than in females. We conclude that male chelicerae and forelegs are under strong positive selection for their use in physical fights and/or as visual signals of fighting ability.     

The functional morphology of species-specific clasping structures on the front legs of male Archisepsis and Palaeosepsis flies (Diptera, Sepsidae)

Several possible explanations for the elaborate species-specific morphology of male front leg clasping organs were tested by comparing six species of Archisepsis, Palaeosepsis and Microsepsis flies. The only previously published hypothesis regarding these clasping organs was refuted by the finding that species-specific portions of the male femur and tibia consistently meshed tightly with prominent veins and folds in the female's wing, rather than meshing with each other. Female wing morphology in the region grasped by the male was relatively uniform and in general did not vary in ways that would prevent non-conspecific males from grasping them, arguing in all but one species against both simple lock-and-key and male-female conflict of interests hypotheses based on morphology. Interspecific differences in male front leg morphology generally represent alternative ways to accomplish the same basic mechanical function of holding tightly onto the relatively invariant female. Despite the fact that female resistance behaviour indicates that male-female conflict over male mounting is common, only one female wing structure in one species resembled an anti-clasper device, giving a second reason to doubt the morphological male-female conflict of interest hypothesis, at least for five of the six species. The positions of probable sensory structures on the wings of females were relatively similar in different species and did not correspond in any obvious way to species-specific features of male clasping structures. This, plus the intraspecific variation in both the positions of these sensilla and the exact site where the male grasped the female's wing, argued against simple 'sensory lock-and-key' ideas about male front leg function. By a process of elimination, it appears that generalized female receptors are able to sense species-specific differences in male front legs. This idea was supported by increased female rejection behaviour in cross-specific pairs.     

Five additions to the list of Sepsidae Diptera for Vietnam: Perochaeta cuirassa sp. n., Perochaeta lobo sp. n., Sepsis spura sp. n., Sepsis sepsi Ozerov, 2003 and Sepsis monostigma Thompson, 1869

A recent collecting trip to Vietnam yielded three new species and two new records of Sepsidae (Diptera) for the country. Here we describe two new species in the species-poor genus Perochaeta (Perochaeta cuirassasp. n. andPerochaeta lobosp. n.) and one to the largest sepsid genus Sepsis (Sepsis spurasp. n.) which is also found in Sumatra and Sulawesi. Two additional Sepsis species are new records for Vietnam (Sepsis sepsi Ozerov, 2003; Sepsis monostigma Thompson, 1869). We conclude with a discussion of the distribution of Perochaeta and the three Sepsis species.     

Copulatory Courtship in Drosophila birchii and D. serrata,Species Recognition and Sexual Selection

Two endemic Australian Drosophila species, D. birchii and D. serrata, have a copulatory courtship, i.e., the males court the female mainly during copulation. In the present study we found the males of both species to mount their prospective mating partners selectively, exhibiting both sex and species recognition. The males began to sing after mounting the female, and they often exhibited also postcopulatory displays typical to copulatory courtship. D. birchii and D. serrata females discriminated against males which did not sing during mounting/copulation, which suggests that the females utilize cryptic female choice. Our findings raise the question of how widespread a phenomenon cryptic female choice is in Drosophila species.     

Complex tibial organs in fore-, mid-, and hindlegs of the bushcricket Gampsocleis gratiosa (Tettigoniidae): Comparison of morphology of the organs

The structure of the complex tibial organs in the fore-, mid-, and hindlegs of the East Asian bushcricket Gampsocleis gratiosa (Tettigoniidae, Decticinae) is described comparatively. In each leg the tibial organs consist of three scolopale organs: the subgenual organ, the intermediate organ, and the crista acoustica. Only in the forelegs are the tibial organs differentiated as tympanal organs, and sound transmitting structures (acoustic trachea, tympana, and tympanal covers) are present. The morphology of the tracheae in the mid- and hindlegs is significantly different from that found in the forelegs. The number of scolopidia in the subgenual organ is highest in the midleg and lowest in the foreleg; in the intermediate organ the number is also highest in the midleg, and the fore- and hindleg contain 40% fewer scolopidia. In the crista acoustica, the number of scolopidia decreases from, the fore- to the mid- and hindlegs. The morphology and the dimensions of the scolopidia and the attachment structures within the crista acoustica of the mid- and hindlegs differ strongly from those in the foreleg. The results indicate that, in addition to the presence of a sound transmitting system, the specific differentiations within the crista acoustica are important for the high auditory sensitivity of the tibial organs in the forelegs. © 1994 Wiley-Liss, Inc.     

Male versus female mate searching in fiddler crabs: a comparative analysis

We present a comparative analysis of mate searching in fiddlercrabs, genus Uca. Several ecological factors determine whichsex will search for mates and how complex male signaling willbe. Female searching is most tightly correlated with matingin male burrows. Female searching is associated with high burrowdensity, small body size, and large soil size. These factors explain variation in a female's need for male-defended incubationsites. Female searching also is correlated with short eyestalks.In species in which females search for mates, males use a morecomplex mate attraction signal than in species in which malessearch.     

Digging Behaviour while Foraging by the European Badger,Meles meles,in a Mediterranean Habitat

Digging behaviour while foraging by the European badger (Meles meles) was studied in the Maremma Natural Park, central Italy, during late winter and spring of 1984 and 1985. Badgers digging for Lamellicornia larvae were successful in approximately 77% of the observed digging attempts. Digging success was similar among individuals and months and was positively correlated with the depth of the hole. A change of position while digging was associated with successful digging. Badgers always started their digging activity using only a foreleg at a constant rate and at a later stage used both forelegs alternately at a higher rate. The adoption of such technique is regarded as adaptive since it maximizes the chance of capturing the prey while reducing the energy expenditure due to digging. The reduced availability of those resources to be collected on the ground surface during late winter and spring is suggested as the possible cause of the energetically expensive digging behaviour.     

小翅稻蝗的精子竞争及交配行为的适应意义

许多昆虫有多次交配行为 ,因多次交配而引起不同雄虫的精子竞争 ,提供雌虫一种有效的性选择方式。小翅稻蝗 (Oxyayezoensis Shiraki)具多次交配行为 ,雌雄交配时间长 ,且交配后常伴有长时间的抱对行为。利用近缘种的种间交配 ,对小翅稻蝗的精子竞争、交配后抱对行为的适应意义进行了探讨。结果表明 ,小翅稻蝗的 P2 值 (最后交配雄虫子代的比例 )达 94 .3%±5 .3% ,说明最后交配雄虫的精子优先用于卵子的受精 ,交配时存在着精子置换。长时间的交配后抱对行为是为了阻止雌虫与不同雄虫个体的再交配 ,保护精子免被置换。     

A comparison of the courtship and mating behavior of three species ofNasonia (Hymenoptera: Pteromalidae)

The parasitic waspNasonia vitripennis (Hymenoptera: Pteromalidae) is the subject of numerous genetic, evolutionary, ecological, and ethological studies, particularly relating to sex-ratio evolution, non-Mendelian inheritance, courtship behavior, and speciation. Here we describe the courtship behavior of two sibling species(N. longicornis andN. giraulti) and compare their courtship to that ofN. vitripennis. Courtship behavior ofNasonia males includes mounting of females, orientation to the female's head, and a repeated series of stereotypic movements, including head nods, mouthpart extrusions, antennal sweeps, and wing vibrations. Females signal receptivity by lowering their antennae in synchrony with opening their genital orifice. After copulation, males engage in brief postcopulatory displays. All three species have the basic components described above. However, although the three species are quite similar morphologically, there are distinct differences in courtship displays. Notable differences include the length of courtship cycles, the absence of a ritualized antennal sweep prior to the first cycle invitripennis, additional head nods and ritualized foreleg movements inlongicornis, and a dramatic increase in numbers of head nods ingiraulti. Results show that closely related species can often be distinguished based on courtship differences, although many of these differences may not contribute to a reproductive barrier.     

Female reproductive decisions and parasite burden in a calopterygid damselfly (Insecta: Odonata)

There is currently a gap in sexual selection theory about how much the environment drives female mating decisions. We present field data that suggest that female sexual behaviour in the damselfly Calopteryx haemorrhoidalis is influenced by parasite burden. Male wing pigmentation in Calopteryx is a sexually selected trait that signals a male's ability to cope with eugregarine parasites (an intestinal parasite that feeds on the adult's ingested food). Because adult C. haemorrhoidalis females also show wing pigmentation, we examined whether this trait is similarly influenced by parasite burden and whether it may signal the female's reproductive value. MaleC. haemorrhoidalis defend riverine substrates that females use for oviposition. After copulation and during oviposition, females are guarded by the copulating male against intruder males. Alternatively, females may avoid mating and ‘steal’ an oviposition site within a male's territory. In the present study, we found that the amount of female wing pigmentation was negatively correlated with the number of eugregarines present. Females with more parasites produced fewer eggs, survived fewer days, spent less time during courtship, ‘inspected’ fewer males before mating, had a lower mating success, were guarded for less time during oviposition and engaged in fewer ‘stealing’ events during oviposition. The reduced egg production and survival of heavily infected females may result from eugregarine depletion of the females' consumed food reserves. Thus, to offset reduced longevity, heavily infected females may accept a mating more rapidly and mate with fewer males. ‘Stealing’ behaviour may be related to the female's differential use of sperm from some males, particularly high-quality males. Interestingly, males that mated with low-pigmented females showed greater variance in wing pigmentation than did males that mated with high-pigmented females. Possibly, female wing pigmentation may signal a female's reproductive value, which provides females with longer mate-guarding episodes and reduced interference from intruder males. This study points out one possible constraint, intestine parasites, that females may face during mating decisions. Because females in bad condition mate with males in both good and bad condition, this constraint may be pervasive enough to weaken the intensity of selection for a male sexually selected trait, wing pigmentation, and help to maintain its variation in phenotypic expression. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Swarming and mating behavior in Ephemera orientalis Mclachlan, 1875 (Ephemeroptera: Ephemeridae) with morphological analyses

Swarming and mating behaviors of a mayfly species, Ephemera orientalis Mclachlan, 1875 were observed in 2015, 2016, and 2018 at a river bank of the Asahi River, Japan. Males started to make swarms between late April and middle May in 2016 and 2018. The numbers of mated pairs in a swarm correlated with the numbers of flying males in a swarm in 2016 and 2018. Swarms were formed during a limited period at dusk most probably because that interval is free from natural enemies. Males competed with each other to copulate with females in swarms. We clarified the function of the forelegs of males, which are significantly longer than those of females. Males used their forelegs to hold up a female from below. Besides forelegs, males have longer tails than females. We will discuss why sexual differences are found in these traits. Our results represent the first observation of swarm mating behavior in E. orientalis.     

Homosexual behaviour and female-male mounting in mammals—a first survey

Information from the literature is given on the presence or absence of homosexual behaviour and female-male mounting in 125 species of mammals, both captive and wild. Such behaviour occurs in the male and often female young of many species soon after their birth. It is more common in young, often in play, than in adults. Adult homosexual behaviour is widespread in male and female mammals (recorded in 63 and 71 species respectively), but common in few species. In males it is most likely to be correlated with dominance and thus to occur in species with hierarchies such as terrestrial monkeys and members of the sheep and goat tribe. In females it is often correlated with sexual condition; a female in heat most often mounted another female, and one in heat was next most likely to be mounted by another female. Anoestral females rarely mounted other anoestral females. Females of 43 species mounted males, which often excited them sexually. Captive mammals tended to mount animals of the same sex more often than did wild ones when comparative data were available. Domestic animals also mounted more man did wild ones, with several exceptions. Some phylogenetic groups of animals displayed similar degrees of homosexual mounting, but there was often considerable variation between closely related species. Nor could homosexual mounting be always correlated with the social structure of a group. The four reasons for, or contexts of, homosexual and female-male mountings were social play (in 34 species), aggression (19 species), sexual excitement (36 species), and physical contact—non-play (30 species). This last category included a state of tension, getting attention, greeting, grooming, caressing, reassurance and appeasement. There was some overlap between categories. Homosexual pair-bonds occur in captive mammals and have been observed throughout the year in non-captive female Japanese monkeys.