Exaggerated male eye span influences contest outcome in stalk-eyed flies (Diopsidae)

Evolution of male weapons or status signals has been hypothesized to precede evolution of female mating preferences for those traits. We used staged male fights among three species of Malaysian stalk-eyed flies (Diptera: Diopsidae) to determine if elongated eye span, which is preferred by females in two sexually dimorphic species, influences contest outcome. Extreme sexual dimorphism, with large males possessing longer eye span than females, is shared by Cyrtodiopsis whitei and C. dalmanni. In contrast, C. quinqueguttata exhibits a more ancestral condition – short, sexually monomorphic eye stalks. Videotape analysis of 20-min paired contests revealed that males with larger eye span and body size won more fights in the dimorphic, but not monomorphic, species. To determine if males from the dimorphic species use eye span directly to resolve contests, we competed male C. dalmanni from lines that had undergone artificial selection for 30 generations to increase or decrease eye span. We found that eye span, independently of body size, determines contest outcome in selected-line males. Furthermore, in both dimorphic species, the average encounter duration declined as the eye span difference between contestants increased, as expected if males use eye span to assess opponent size. The number of encounters also increased with age in dimorphic, but not monomorphic, species. Selected-line males did not differ from outbred males in either fight duration or number of encounters. We conclude that exaggerated male eye stalks evolved to influence both competitive interactions and female mating preferences in these spectacular flies. Received: 20 July 1998 / Received in revised form: 2 February 1999 / Accepted: 12 March 1999     

Patterns of parental provisioning covary with male morphology in a horned beetle (Onthophagus taurus) (Coleoptera: Scarabaeidae)

Male dung beetles, Onthophagus taurus, are dimorphic for a secondary sexual trait, head horns. Horned males participate in the production of brood masses while hornless male do not. Here we examine the reproductive performance of females mated with males exhibiting alternative horn morphologies. We found that exposure to males may be costly for females in that it reduced the total number of brood masses produced. However, females paired with horned males produced significantly larger brood masses than females paired with hornless males or females producing broods alone. We discuss the possible selection pressures that may underly horn evolution in this genus. Received: 22 August 1997 / Accepted after revision: 19 January 1998     

Breeding of Euterpina acutifrons,a harpacticid copepod,with special reference to dimorphic males

The breeding of Euterpina acutifrons, a marine harpacticid copepod, was studied both in the laboratory and in the natural population round Anglesey, England, with special reference to dimorphic males. As a result of successful rearing of the species under laboratory conditions, rates of development at different temperatures, and survival and maturation of all types of adults were studied. The dimorphic males developed faster than the females, and the small males had a shorter development time than the larger individuals. Laboratory observations clearly indicated that the males differ markedly in sexual behaviour, the small male being more potent, readily copulating with females, and always very active in breeding. In natural populations, temperature was found to have a limiting influence on the breeding period. Breeding began when the temperature reached 15° to 16°C, and ceased when it fell to 8°C. The most significant feature was the production of small males in nature; they increased in number with the onset of breeding, and always formed a high proportion of the actively breeding population. Both laboratory and field investigations have shown that the small male is a breeding form, and is apparently an adaptation of the species for carrying out successful breeding in the colder areas of its distribution.     

Sexual dimorphism and male mating success in the tentacled blenny, <Emphasis Type="Italic">Parablennius tentacularis</Emphasis> (Teleostei: Blenniidae)

Although external sexually dimorphic traits are commonly found in males of combtooth blenny species, little is known about the benefit they can convey to male mating success. Indeed, while female preferences for large males have been demonstrated in some species, the possible role played by dimorphic ornaments has been neglected. We now report on the tentacled blenny, Parablennius tentacularis, a species where males are characterized by bulb glands on the anal fin and both sexes exhibit a dark spot on the dorsal fin and orbital tentacles. Males are territorial, make nests in empty bivalve shells, and provide solitary parental care for the eggs. Using morphometric analysis and field collected data on male and female external features, nest characteristics and number of eggs in the nests, we have assessed the development of dimorphic traits in both sexes and male mating success. The results reveal that orbital tentacles of males are more developed and more variable in size than those of females. Larger males exhibit longer orbital tentacles and larger anal glands but do not necessarily occupy larger nests. Male mating success is significantly correlated with the inner nest surface area and with orbital tentacle size but not with body size. These results provide support for a primary role of male ornaments in enhancing blenny male mating success and are discussed in the context of mate choice for direct and indirect benefits.     

On the sociobiology of four puerto rican parrotfishes (Scaridae)

Behavioral observations were made during SCUBA excursions out of the PRINUL underwater laboratory, off the west coast of Puerto Rico. Most of the work involved four species of parrotfishes, Sparisoma viride, Sparisoma aurofrenatum, Scarus croicensis and Scarus taeniopterus. The species of Sparisoma are relatively patrilocal, solitary, slowly and steadily feeding fishes; the terminal phase (termphase) males of S. aurofrenatum hold large territories that encompass the home ranges, possibly territories, of a number of females. The two species of Scarus are roving, aggregating fish that periodically descend to the bottom where they feed rapidly. Reproductive males are of two types, either highly dimorphic, brilliantly colored termphase males, or midphase males that resemble the females. At times, termphase males become territorial and are visited by females and midphase males. Females spawn with termphase and with midphase males. Aggression in all four species was directed largely to conspecific individuals. All species show the ability to change color rapidly, sometimes dramatically, in social interactions.     

Mating patterns and a test for female mate choice in Etheostoma spectabile (Pisces,Percidae)

Neither size nor breeding color correlated with spawning success of male orangethroat darters, Etheostoma spectabile (Pisces, Percidae), under natural field conditions. When females were presented experimentally with a simultaneous choice they spent no more time in proximity to large than small males, and were subsequently no more likely to spawn with large than with small males. Females also displayed no preference for bright versus dull males. Males and females did not differ significantly in size. Etheostoma spectabile may lack sexual size dimorphism as a result of the lack of female choice for size and the ineffectiveness of male attempts to monopolize females, or selection may be for increased size of females. Males are not dwarfs because of sperm competition. Contest competition among males appeared to be important in initiating spawnings but many males obtained spawnings by participating in ongoing spawning events. Etheostoma spectabile is an example of a sexually dimorphic species with no evident female preference for male size or color.     

Variance in the male reproductive success of western gorillas: acquiring females is just the beginning

Variance in male reproductive success is expected to be high in sexually dimorphic mammals, even when it is modulated by the costs and benefits of group living. Here, we investigate the variance in reproductive success of male western gorillas (Gorilla gorilla), a highly dimorphic primate with long-term male–female associations, using 12.5 years of data collected at Mbeli Bai in northern Congo. Access to mates and offspring survival were both major sources of variance in male reproductive success. Males with larger harems had lower offspring mortality with no apparent reduction in female fertility or observed tenure length, so the size of harems did not seem to be limited by female feeding competition or by the risk of takeovers and infanticide by outsider males. The lower mortality in larger harems may reflect improved vigilance against predators, and females may cluster around males that enhance offspring survival. Thus, this study illustrates how a detailed analysis of the components of male reproductive success can shed light on the interrelated social and ecological aspects that affect it.     

Evolution of male dimorphic allometry in a population of the Japanese horned beetle <Emphasis Type="Italic">Trypoxylus dichotomus septentrionalis</Emphasis>

I conducted a detailed morphological analysis of the Japanese horned beetle Trypoxylus dichotomus septentrionalis to clarify the allometric relationship between horn length and body size and examined its mating success and reproductive behaviour in the field. The relationship between horn and body size was not discontinuous at the switch point body size, but the slope of the linear relationship changed at the switch point. Shape of the allometric relationship was initially steep and became flatten around the switch point in both linear and log scales; that is, minor males showed a positive relationship and major males showed a negative one. Major males gained more mating success than minor males. Within major males, individuals with larger horn or body size had higher mating success than individuals with smaller ones. Within minor males there were no differences in horn and body size between mated and unmated individuals. Although sneak-like behaviours were exhibited by both morphs, it is likely that these behaviours rarely lead to direct benefit. These results suggest that dimorphic allometry of T. dichotomus is consistent with the hypothesis of a continuous reaction norm that meets a ceiling, which restrains further allometric growth.     

Comparative thermal metabolic patterns in Euterpina acutifrons dimorphic males

Thermal metabolic acclimation patterns have been determined for cold-and warm-acclimated dimorphic males of the copepod Euterpina acutifrons. The copepods were acclimated either to 15° or 25°C. Metabolic measurements were made at the two acclimation temperatures. At 25 °C, the small males had a higher rate than the large ones. At 15 °C, the large males had the higher rate. The metabolic pattern of thermal acclimation is also distinctive in the two forms. Small males showed metabolic adaptation at the lower acclimation temperature (15 °C), but not at the higher acclimation temperature (25 °C). In the large males the acclimation patterns were reversed, i. e., these males acclimated at the higher temperature, not at the lower. The acclimation patterns based on Precht's (1958) scheme of acclimation are entirely different in the two forms.Supported by PHS grant 5-SO5-FR-07070-02.     

The morphology and behavior of dimorphic males in Perdita portalis (Hymenoptera : Andrenidae)

Summary In Perdita portalis, a ground nesting, communal bee, males are clearly dimorphic. The two male morphs are easily distinguished based on head size and shape into (1) a flight-capable, small-headed (SH) morph that resembles the males of other closely related species and (2) a flightless, large-headed (LH) morph that possesses numerous derived traits, such as reduced compound eyes, enlarged facial foveae and fully atrophied indirect flight muscles. The SH morph occurs exclusively on flowers while the LH morph is found only in nests with females. While on flowers, SH males are aggressive, fighting with conspecific males and heterospecific male and female bees, and they mate frequently with foraging females. Using artificial observation nests placed in the field, I observed the behavior of females and LH males within their subterranean nests. LH males are aggressive fighters; males attacked each other with mandibles agape, and male-male fights always ended in the death of one male. LH males are highly attentive to the reproductive behavior of females; they spend increasing amounts of time near open cells during cell provisioning, and mating only takes place immediately prior to oviposition when females are forming the accumulated pollen and nectar into a ball. Based on larvae reared to adulthood in the laboratory, the two male morphs occur in equal proportions. The behavior of males in closely related species, especially P. texana, and the origin and maintenance of male dimorphism are discussed.     

The cost of success: reproductive effort in male southern elephant seals (<Emphasis Type="Italic">Mirounga leonina</Emphasis>)

Reproductive effort is a key parameter of life history because it measures the resources allocated to reproduction at the expense of growth and maintenance. Male reproductive effort always had a minor role with respect to female effort both in the development of theories and in field research. Elephant seals are an ideal subject for reproductive effort studies because they fast during the breeding season, splitting the phase of energy acquisition from the phase of energy use for breeding. In this paper, we present results on male reproductive effort (weight loss estimated by photogrammetry) in southern elephant seals (Mirounga leonina), the most dimorphic and polygynous of all mammal species. We show that total reproductive effort increases with age, with no sign of late decrease or senescence. Male reproductive effort in this species depends mostly on behavioral factors, i.e., the success in competition with other males, and the intensity of interaction with females. A large effort results in large gains in both mating success and fertilizations. The large reproductive success that a few males are able to achieve come at a big cost in terms of energy expenditure, but this cost does not seem to affect the likelihood of survival to the following breeding season.     

Correlates of male mating success in great bustard leks: the effects of age, weight, and display effort

We examined how mating success varied in relation to age, weight, body size, and display behavior among great bustard Otis tarda males. The estimated mating success was strongly skewed, with 45% of adult males being involved in copulation attempts and only 9.7% actually seen copulating successfully. Unlike most birds, body size continued increasing in great bustards several years after reaching sexual maturity. Age, weight, and display effort were all significant and independent predictors of male mating success. The higher display effort involved performing longer full-display bouts. Older males could detach from the male flock earlier in the season as well as on each day and spend longer seasonal and daily periods displaying as solitary birds, which contributed to increase their mating success. In contrast, males weighing more did not invest more in display, which suggests that they could be recognized as dominants by other males and selected by females through assessment of their plumage sexual traits. In contrast to most other bird species, the system described for great bustards resembles that found in some lek-mating ungulates, where social rank is a complex trait determined by both age and mass, and as in these mammals, it suggests that sexual selection continues to favor a high male weight in this extremely sexually dimorphic species.     

Is <Emphasis Type="Italic">Hippolyte williamsi</Emphasis> gonochoric or hermaphroditic? A multi-approach study and a review of sexual systems in <Emphasis Type="Italic">Hippolyte</Emphasis> shrimps

Sexual systems vary considerably among caridean shrimps and while most species are gonochoric, others are described as sequential protandric hermaphrodites or simultaneous hermaphrodites with an early male phase. At present, there is confusion about the sexual system exhibited by several species mostly because those studies attempting to reveal their sexual system draw inferences solely from the distribution of the sexes across size classes. Here we investigated the sexual system of the shrimp Hippolyte williamsi from Chile to determine if the species is protandric or gonochoric with sexual dimorphism (males smaller than females). Morphological identification and size frequency distributions indicated that the population comprised small males, small immature females, and large mature females, which was confirmed by dissections. No transitional individuals were found. Males maintained in the laboratory molted 1–8 times, and many grew up to reach sizes observed in only a small fraction of males in the field. No indication of sex change was recorded. Our results indicate that H. williamsi is a sexually dimorphic gonochoric species and emphasizes the importance of using several kinds of evidence (size measurements, growth experiments, morphological dissections, and histological studies) to reveal the sexual system of Hippolyte species. Whether the observed size dimorphism between males and females in many species of Hippolyte is expression of contrasting sexual and natural selection, and whether divergent sexual fitness functions can contribute to the evolution of hermaphroditism remains to be revealed in future studies.     

An experimental study of competition for food between male and female nestlings of the red-winged blackbird

Summary Ten experimental broods of red-winged black-bird (Agelaius phoeniceus) nestlings, experimentally manipulated to contain two males and two females of similar age, were observed with the aid of video cameras to determine (a) which nestling characteristics were most important in influencing its chances of being fed and (b) if males and females differed with respect to these characteristics. Nestlings that could reach highest while begging were most successful at obtaining food from parents during individual feeding bouts. However, while there was a tendency for nestlings that begged earlier to be offered food first, a nestling's position in the nest during these bouts did not influence its feeding success. Males begged more than often females and were offered more food by parents. Males were also more likely to be fed when they begged. This was most likely because males, being larger than females, were able to reach higher while begging. Neither sex was more likely to beg sooner or occupy certain positions in the nest. However, while males that had been transferred from other nests received less food than natal males, there were no differences in food acquisition between transferred and natal females. Differences in the relative success of males and females under different conditions may help explain differences in fledgling sex ratios observed in many dimorphic species.     

Population dynamics of Euterpina acutifrons (Copepoda: Harpacticoida) from North Inlet,South Carolina,with reference to Dimorphic Males

The population dynamics of Euterpina acutifrons (Dana), a pelagic, harpacticoid copepod, are summarized in a life table based on field data. Highest mortality occurred in the last naupliar stage (NVI) and the first copepodite stage (CI). Overall survival in the field was 0.06% from the first naupliar stage (NI) to adult (CVI). The net reproductive rate (R _o=55.590) and intrinsic rate of increase (I _m=0.28) were sufficiently high to maintain a population with such a low survival rate in nature. E. acutifrons was present and breeding in the field from April through December. Low temperatures limited breeding, which began when the temperature reached 16.5°C and ceased when it fell to 11°C. Optimum temperature for North Inlet E. acutifrons was 25°C, with a maximum laboratory survival of 15.3% and a generation time of 10.3 days. Generation time in the field (20°C) was 14 days. Temperature also affected the abundance of dimorphic males. Small males were always most abundant, but peaked during the coldest month; large males became equal in abundance only during the varmest months.Contribution No. 298 from the Belle W. Baruch Institute for Marine Biology and Coastal Research.     

Shouting the odds: vocalization signals status in a lizard

Many species possess multiple sexually dimorphic traits, which incorporate different sensory modalities (e.g., acoustic, olfactory and visual), although their relative roles in sexual selection and in determining reproductive success are still poorly understood for most taxa. We assessed the role of multiple male traits, including one acoustic (dominant call frequency) and one visual (yellow throat patch) trait, in residency advertisement, contest behavior, and breeding success in barking geckos (Ptenopus garrulus garrulus). We show that male barking geckos maintain largely exclusive home ranges, with a trend for larger males to maintain larger home ranges. We also show that larger males have a lower dominant calling frequency. When aggressive behavior was elicited in the field using a recorded call of average frequency, resident males with low frequency calls were more likely to respond aggressively and charge the speaker compared to males with high frequency calls. However, body size and small relative throat patch size, rather than call frequency, were the best predictors of overall aggressiveness. Body size was also the best predictor of whether males bred. We suggest that call frequency in this crepuscular species constitutes an effective long-range signal of body size, used by males for remote rival assessment and to advertise home range boundaries in low-light environments.     

Resource quality or harem size: what influences male tenure at refuge sites in tree weta (Orthoptera: Anostostomatidae)?

Rather than seeking females directly, males in many animal species locate and defend sites that contain spatially limited resources essential for female survival and reproduction. In these cases, resident males successfully repelling conspecific rivals will mate with sexually receptive females that seek to use the resident’s resources. Theory predicts that if resources are limiting in nature, are costly to procure, and if residency at the resource site increases male reproductive success, each site should be monopolised by a single adult male. Moreover, if females aggregate at these sites, males should be sedentary and monopolise harems. This predicts that males should reside at resource sites longer than females and male tenure should be positively correlated with harem size. I address these hypotheses using a wild population of the Wellington tree weta, Hemideina crassidens, a sexually dimorphic insect in which males use mandibular weapons in fights to control resource sites (galleries in trees) required by females for diurnal refuge. In a longitudinal study using artificial galleries, I show that male tenure in a gallery is positively related to harem size and, contrary to prediction, not gallery size per se. Moreover, contrary to prediction, females are more sedentary than males: males appear to move frequently between galleries to search for novel mates. I show experimentally that galleries represent mating sites to males: males prefer galleries housing adult females whereas females prefer unoccupied galleries. Females likely avoid male-occupied galleries because they incur injuries when interacting with males.     

Alternative mating tactics in dimorphic males of the harvestman <Emphasis Type="Italic">Longiperna concolor</Emphasis> (Arachnida: Opiliones)

Intense male–male competition for females may drive the evolution of male morphological dimorphism, which is frequently associated with alternative mating tactics. Using modern techniques for the detection of discontinuous allometries, we describe male dimorphism in the Neotropical harvestman Longiperna concolor, the males of which use their elongated, sexually dimorphic legs IV in fights for the possession of territories where females lay eggs. We also tested three predictions related to the existence of alternative mating tactics: (1) if individuals with relatively longer legs IV (majors) are more likely to monopolize access to reproductive resources, they are expected to remain close to stable groups of females more than individuals with relatively shorter legs IV (minors) do; (2) if minors achieve fertilization by moving between territories, they are expected to be less faithful to specific sites; and (3) majors should be observed in aggressive interactions more often. We individually marked all the individuals from a population of Longiperna during the reproductive season and recorded the location of each sighting for males and females as well as the identity of males involved in fights. Majors were more likely to have harems, and large majors were even more likely to do so. Majors were more philopatric and all males involved in fights belonged to this morph. These results strongly suggest that the mating tactic of the majors is based on resource defense whereas that of the minors probably relies on sneaking into the territories of the majors and furtively copulating with females.     

Copulation duration and sperm precedence in the stalk-eyed fly Cyrtodiopsis whitei (Diptera : Diopsidae)

Summary By means of field observations and laboratory experiments on the Malaysian stalk-eyed fly Cyrtodiopsis whitei we examined the consequences of variation in copulation duration for sperm competition. In this sexually dimorphic species over 90% of all copulations occur in nocturnal aggregations with from one to four males and up to 24 females. Copulation duration observed in both the field and the laboratory exhibited a bimodal distribution with peaks at 10 and 50 s. In the field short copulations less than 30 s long occurred frequently when more than one male was present in an aggregation but most were not the direct result of male interference. Sperm counts from female spermathecae after artificial interruptions indicated sperm are not transferred during the first 40 s of a copulation. When solitary males mated up to five times in succession to virgin females, short copulations did not occur, nor was the number of sperm transferred reduced. However, short copulations did occur when we mated isolated females within 6 min of a previous copulation. By mating irradiated and non-irradiated males in reciprocal pairs we discovered that C. whitei exhibits both first-male sperm precedence and sperm mixing. More than half of the females mated first to sterile and then to fertile males failed to produce offspring. Such variation in copulation duration and sperm precedence is consistent with male placement and detection of a spermatophore that acts as a temporary mating plug. Our data suggest that those male C. whitei which successfully defend large aggregations of females reduce sperm waste and competition by preferentially transferring sperm to females that have not mated recently. Correspondence to: G.S. Wilkinson     

Facultative sex ratio manipulation in American kestrels

Summary For animals that are sexually dimorphic in size, the larger sex is expected to be more costly to raise to independence. Manipulating offspring sex ratios may thus be one means by which parents can fine-tune their reproductive effort to resource availability. Parents in poor physical condition or during poor food years should produce more of the cheaper (smaller) sex. We examined the sex ratios of 259 broods of American kestrels (Falco sparverius) between 1988 and 1990 in relation to food abundance (small mammals) and various attributes to the parents. The proportion of males at hatching increased as the food supply declined, and both male and female parents in poor physical condition were more likely to have male-biased broods than those in good condition. The mortality of eggs and young did not appear to be responsible for the biased sex ratios. The sex ratio was independent of the laying date; however, it was correlated with female body size. Small females produced more sons, perhaps because small size is more detrimental for females than males. Offprint requests to: G.R. Bortolotti