Sympatry and allopatry of song in North American tanagers

Summary The possibility that the hypotheses of character convergence and character displacement might explain interspecific variation in song features among North American tanagers is examined. Scarlet and summer tanagers (Piranga olivacea and P. rubra) are the most sympatric among the four species and have the largest differences in song characteristics. However, songs of scarlet tanagers do not differ significantly between sympatry and allopatry. Songs of summer tanagers more resemble those of scarlet tanagers in allopatry, but this geographical variation has been explained by variation in habitat characteristics. Both species countersing in sympatry, and react aggressively to each other's songs in sympatry as well as in allopatry, probably due to similarities in the song. The limited data from the narrow sympatric zone between western and hepatic tanagers (P. ludoviciana and P. flava) indicate that their songs differ to a great extent. Therefore, the results do not support the hypothesis of character convergence. Character displacement might explain partially interspecific differences in song features between scarlet and summer tanagers but cannot be tested further.     

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.     

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     

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.     

Geographical patterns in reproductive biology of the Pan-American sandy beach isopod<Emphasis Type="Italic"> Excirolana braziliensis</Emphasis>

Eleven populations of the Pan-American sandy beach isopod Excirolana braziliensis, distributed from tropical (9°N) to temperate (39°S) sandy beaches in Atlantic and Pacific Oceans, were analyzed to determine latitudinal variations in: breeding and recruitment patterns, sex ratios, size of ovigerous females and of juveniles and length–fecundity relationships. E. braziliensis exhibited strong latitudinal patterns in all reproductive traits throughout its distribution range. Breeding and recruitment shifted from continuous to seasonal from tropical to temperate beaches, having a predominance of females at higher latitudes. In agreement with the latitudinal gradient hypothesis, ovigerous females inhabiting tropical sandy beaches (low latitudes) were smaller, became sexually mature at smaller sizes and had lower individual fecundity than on temperate beaches. Juveniles were also smallest at low latitudes. Between-ocean comparisons showed very similar reproductive characteristics for roughly the same latitude. These linked reproductive parameters suggest that the intensity of breeding effort is associated with the duration of the breeding season and geographically size-related characteristics of the species. Geographic variations in the breeding and recruitment seasons, as well as in individual fecundity, size structure of mature females and sex ratios, are proposed to have major consequences in explaining local variations in population demography. Our paper also reinforces the notion that sandy beach animals are highly plastic.Communicated by O. Kinne, Oldendorf/Luhe     

Grassland songbird survival and recruitment in agricultural landscapes: implications for source-sink demography

Population growth and decline are particularly sensitive to changes in three key life-history parameters: annual productivity, juvenile survival, and adult survival. However, for many species these parameters remain unknown. For example, although grassland songbirds are imperiled throughout North America, for this guild, only a small number of studies have assessed these parameters. From 2002 to 2006, in the agricultural landscape of the Champlain Valley of Vermont and New York, USA, we studied Savannah Sparrow (Passerculus sandwichensis) and Bobolink (Dolichonyx oryzivorus) demography on four grassland treatments: (1) early-hayed fields cut before 11 June and again in early- to mid-July; (2) middle-hayed fields cut once between 21 June and 10 July; (3) late-hayed fields cut after 1 August; and (4) rotationally grazed pastures. We assessed whether these treatments affected adult apparent survival (phi) and recruitment (f), how sensitive these parameters were to the presence of nonbreeders and local dispersal, and the populations' ability to persist in these four habitats. On average, birds using late-hayed fields had > 25% higher apparent survival than those on the more intensively managed early-hayed, middle-hayed, and grazed fields. Overall male phi was 35% higher than female phi, and Savannah Sparrow phi was 44% higher than Bobolink phi. Across all analyses and treatments, apparent survival estimates were 0.58-0.85 for male and 0.48-0.71 for female Savannah Sparrows, and 0.52-0.70 for male and 0.19-0.55 for female Bobolinks. For males of both species, potential nonbreeders decreased the precision of and lowered apparent survival estimates by 25%; female estimates showed little variation with the inclusion of nonbreeders. Inclusion of local dispersal observations increased apparent survival estimates and, in many cases, increased precision, though the effect was stronger for Savannah Sparrows than for Bobolinks, and also stronger for males than for females. High Savannah Sparrow apparent survival rates resulted in stable or near stable populations (lambda approximately 1), particularly in late-hayed and grazed fields, while low Bobolink apparent survival rates resulted in strongly declining populations (lambda < 1) in all treatments.     

Consumer resource matching in urbanizing landscapes: are synanthropic species over-matching?

Population responses of synanthropic species to urbanization may be explained by the resource-matching rule, which postulates that individuals should distribute themselves according to resource availability. According to the resource-matching rule, urban habitats will contain greater densities if they provide better resources than rural habitats. However, because resource availability is density dependent, individuals in urban areas would ultimately achieve fitness levels comparable to, but no better than, individuals in less urban areas. Some ecologists suggest that synanthropic birds may not conform to the resource-matching rule and may instead overmatch (i.e., overexploit) in urban habitats, ultimately leading to lower fitness despite greater resource levels. Using the Northern Cardinal (Cardinalis cardinalis) as a focal species, we evaluated if Cardinal populations in urban and rural habitats were consistent with predictions of consumer resource matching. During 2003-2006 we documented population density, adult body condition, apparent survival, and annual reproductive productivity of Cardinals in riparian forest stands within urban (n = 8 stands) and rural (n = 6 stands) landscapes in Ohio, USA. Density of Cardinals in urban forests was four times that found in more rural forests. Mark-resight data from 147 males and 125 females over four years indicated that apparent survival rates were similar between urban and rural landscapes (phi = 0.64, SE = 0.039 for males and phi = 0.57, SE = 0.04 for females). Similarly, body condition indices of 168 males, 142 females, and 118 nestlings did not differ significantly between landscapes. Annual reproductive productivity (mean number of fledglings per pair over breeding season) of 294 pairs was comparable for urban (2.4 +/- 0.18 [mean +/- SE] and rural (2.1 +/- 0.18) young birds. Thus, contrary to recent suggestions, we find that high densities of certain synanthropic species in urban landscapes are consistent with expectations of consumer resource matching.     

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.     

Shift in foraging grounds and diet broadening during ontogeny in southern elephant seals from Kerguelen Islands

Highly dimorphic species like southern elephant seals (Mirounga leonina, SES hereafter) frequently exhibit resource partitioning according to sex and/or age classes. We measured carbon and nitrogen stable isotopes (δ¹³C and δ¹⁵N) of 404 blood samples (136 males and 268 females from Kerguelen Islands, 49°21′S, 70°18′E) from 2004 to 2011. Assuming that the distribution of carbon isotopes (δ¹³C value) reflects the two main foraging grounds (Polar Frontal and Antarctic Zones), we quantified the proportion of SES foraging within each zone in relation with size, a proxy for their age. We found a clear shift from Polar Frontal to Antarctic waters as male SES aged, but no relation as far as females is concerned. We also observed a widening range of nitrogen isotopic (δ¹⁵N) values, suggesting that both males and females expanded their diet spectra with age. Whereas males increased their trophic level, females remained constant on average, with some adult females feeding both at lower and at higher trophic levels than juveniles.     

Life history trade-offs in tropical trees and lianas

It has been hypothesized that tropical trees partition forest light environments through a life history trade-off between juvenile growth and survival; however, the generality of this trade-off across life stages and functional groups has been questioned. We quantified trade-offs between growth and survival for trees and lianas on Barro Colorado Island (BCI), Panama using first-year seedlings of 22 liana and 31 tree species and saplings (10 mm < dbh < 39 mm) of 30 tree species. Lianas showed trade-offs similar to those of trees, with both groups exhibiting broadly overlapping ranges in survival and relative growth rates as seedlings. Life history strategies at the seedling stage were highly correlated with those at the sapling stage among tree species, with all species showing an increase in survival with size. Only one of 30 tree species demonstrated a statistically significant ontogenetic shift, having a relatively lower survival rate at the sapling stage than expected. Our results indicate that similar life history trade-offs apply across two functional groups (lianas and trees), and that life history strategies are largely conserved across seedling and sapling life-stages for most tropical tree species.     

The economics of harem maintenance in the sac-winged bat, Saccopteryx bilineata (Emballonuridae)

Saccopteryx bilineata has a polygynous mating system in which males defend females in a harem territory. Harem defense and courtship include energetically costly flight maneuvers and hovering displays. We tested if (1) harem males have a greater field metabolic rate than non-harem males or females and if (2) the field metabolic rate of harem males is correlated with the number of females in a harem territory. We measured the energy budget in 32 S. bilineata with the doubly labeled water method and compared these estimates with behavioral observations in the daytime roost. Among adult bats, field metabolic rate varied with body mass by an exponent of approximately two. We found no significant difference in field metabolic rate or mass-specific field metabolic rate between harem and non-harem males. The mass-specific field metabolic rate of harem-males increased with harem size. The latter finding supports the hypothesis that the energy costs of courtship display and territorial defense influence the energy budget of harem males. Overall, field metabolic rates of S. bilineata were lower than those of similarly sized bats of the temperate zone and only 2.3 times above the basal metabolic rate recorded for this species. We suggest that male S. bilineata did not take advantage of their metabolic capacity because a prudent allocation of energy to activities of harem maintenance is an adaptive strategy for males in this mating system.     

A Contemporary,Sex‐Limited Change in Body Size of an Estuarine Turtle in Response to Commercial Fishing

Abstract: Juvenile growth rate and adult body size are important components of life‐history strategies because of their direct impact on fitness. The diamondback terrapin (Malaclemys terrapin) is a sexually dimorphic, long‐lived turtle inhabiting brackish waters throughout the Atlantic and Gulf coasts of the United States. In parts of its range, terrapins face anthropogenically imposed mortality: juveniles of both sexes inadvertently enter commercial crab traps and drown. For adult females, the carapace eventually grows large enough that they cannot enter traps, whereas males almost never reach that critical size. We compared age structure, carapace dimensions, growth curves, and indices of sexual dimorphism for a Chesapeake Bay population of terrapins (where mortality of turtles is high due to crab traps) with contemporary terrapins from Long Island Sound and museum specimens from Chesapeake Bay (neither group subject to commercial crab traps). We also calculated the allochronic and synchronic rates of evolutionary change (haldanes) for males and females to measure the rate of trait change in a population or between populations, respectively. We found a dramatic shift to a younger male age structure, a decrease in the length of time to terminal female carapace size, a 15% increase in female carapace width, and an increase in sexual dimorphism in Chesapeake Bay. In a new twist, our results implicate a fishery in the selective increase in size of a reptilian bycatch species. These sex‐specific changes in life history and demography have implications for population viability that need to be considered when addressing conservation of this threatened turtle.     

Territory defence in tropical birds: are females as aggressive as males?

Much of our knowledge concerning the functions of territorial behaviour and how territories are defended by individuals comes from research on birds. The vast majority of this work has focused on temperate zone breeding territoriality in which territories are defended most obviously by males. Our understanding of the female role in territory defence is limited because they are less conspicuous and much harder to observe. We studied sex roles in territory maintenance and defence in a duetting, resident neotropical passerine, the white-bellied antbird (Myrmeciza longipes). This species maintains territories and pair bonds year round and both sexes sing and actively participate in territory defence. We performed a series of playback experiments throughout the dry (non-breeding) and wet (breeding) seasons. We exposed territorial pairs to three types of stimuli including: (1) single sex, male only songs, (2) single sex, female only songs, and (3) both sex songs/duets. Contrary to findings for most other tropical species, individuals defended their territories with equal levels of aggression regardless of stimuli. Furthermore, sex roles were very different, with males responding more aggressively than females to all stimuli throughout both seasons. Both males and females consistently responded more aggressively to territorial intrusions during the dry season than during the wet season, likely because food abundance is low in the dry season and territory value is high. Our analysis of duetting behaviour suggests that duets do not serve a significant role in mate guarding, or territory defence.     

Parental care and mate attraction in the Florida flagfish, <Emphasis Type="Italic">Jordanella floridae</Emphasis>

Theory suggests that there is a trade-off between mate attraction and investment in parental care. Yet, under some circumstances, such as in species with uniparental male care, it is conceivable that males who provide more care also attract more females, and hence avoid this cost. We tested these two alternative hypotheses using the Florida flagfish, Jordanella floridae, which is a highly sexually dimorphic species with male parental care. We predicted that if components of male care behavior primarily serve offspring survival, they should display these behaviors more in the presence of eggs than without eggs. We also predicted that males should show a trade-off between display, and other activities directed to other fish, and care behaviors. Alternatively, if care has a positive effect on mating success, we expected males to show care even in the absence of eggs, and that they should display more care in the presence of females. We found that care behavior mostly depended on the presence of eggs, males increased care when they guarded eggs, and there was a negative relationship between rates of interaction and care behavior. Most care was exhibited in the absence of other fish while the presence of males and females both had a negative effect on care behavior. There was no difference in care behavior with males or females present. Only nest-visitation rates were highest in the presence of females, suggesting that this behavior is used as part of the courtship. These results support the idea that male care in this species is primarily influenced by natural selection. Males did, however, display fanning and other egg-directed behaviors even in the absence of eggs. The level of these behaviors was constant and independent of display behavior. Hence, it seems that care may be used in mate attraction before the male has received his first clutches.     

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.     

Female adaptation to developmental plasticity in male calling behavior

The katydid Neoconocephalus triops exhibits in North America substantial developmental plasticity of male mating calls. The AM rate of the summer calls is significantly faster than that of the winter calls at the same temperature. In the tropics, where N. triops originated, males express only the fast summer-call phenotype. We tested two alternative hypotheses: (1) call plasticity in the population from North America evolved in response to selection by female preference after N. triops colonized North America, or (2) call plasticity evolved before N. triops expanded into North America and its expression in the novel environment led to adaptive change of female preferences. First, we tested whether call plasticity was present in tropical populations of N. triops. Tropical males expressed the winter-call phenotype when reared under winter conditions, indicating that call plasticity did not evolve in response to temperate climates. Second, we compared female preferences among temperate and tropical populations. We found that the temperature dependence of preferred AM rate was significantly steeper in temperate N. triops than in a tropical population of N. triops. Third, we compared temperature dependence of female preference of the N. triops populations to three Neoconocephalus species without call plasticity. Only temperate N. triops had significantly steeper temperature dependence than the other species. This steeper temperature dependence matched female preference to the fast summer call at high temperatures and to the slow winter call at low temperatures in temperate populations. These results support the hypothesis that female preference changed in N. triops in North America to compensate for the plasticity of male calls.     

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     

Overwinter Survival of Neotropical Migratory Birds in Early-Successional and Mature Tropical Forests

Many Neotropical migratory species inhabit both mature and earty-successional forests on their wintering grounds, yet comparisons of survival rates between habitats are lacking. Consequently, the factors affecting habitat suitability for Neotropical migrants and the potential effects of tropical deforestation on migrants are not well understood. We estimated overwinter survival and capture probabilities of Wood Thrush ( Hylochichla mustelina ), Ovenbird ( Seiurus aurocapillus ), Hooded Warbler ( Wilsonia citrina ), and Kentucky Warbler ( Oporornis formosus ) inhabiting two common tropical habitat types, mature and early-successional forest. Our results suggest that large differences (for example, ratio of survival rates (γ) ≤ 0.85) in overwinter survival between these habitats do not exist for any of these species. Age ratios did not differ between habitats, but males were more common in forest habitats and females more common in successional habitats for Hooded Warblers and Kentucky Warblers. Future research on overwinter survival should address the need for age- and sex-specific survival estimates before we can draw strong conclusions regarding winter habitat suitability. Our estimates of overwinter survival extrapolated to annual survival rates that were generally lower than previous estimates of annual survival of migratory birds. Capture probability differed between habitats for Kentucky Warblers, but our results provide strong evidence against large differences in capture probability between habitats for Wood Thrush, Hooded Warblers, and Ovenbirds. We found no temporal or among-site differences in survival or capture probability for any of the four species. Additional research is needed to examine the effects of winter habitat use on survival during migration and between-winter survival.     

Pre-reproductive survival in a tropical bird and its implications for avian life histories

The factors that affect survival until reproduction are essential to understanding the organization of life histories within and among species. Theory predicts, for example, that survival until reproduction influences the optimum level of reproductive investment by parents, which might partly explain prolonged parental care in species with high first-year survival. Tests and refinements of life-history theory have been hampered, however, by a lack of field-based estimates of pre-reproductive survival, especially for tropical species, which have been the subject of many comparative analyses. Tropical species are predicted to have higher first-year survival and delayed reproduction compared to Northern Hemisphere species. We estimated survival until reproduction, age at first reproduction, and sources of variation in juvenile survival in a Neotropical passerine, the Western Slaty-Antshrike (Thamnophilus atrinucha), in central Panama. We observed that fledged antshrikes had 76% survival through the dependent period and 48% survival to the age of 1 year; survival rate was lowest during the first week after leaving the nest. Timing of fledging within the breeding season, fledgling mass, and age at dispersal influenced survival, while sex of offspring and year did not. Individuals did not breed until two years of age, and post-fledging pre-reproductive survival was 41% of annual adult survival. High survival until reproduction in antshrikes balanced their low annual productivity, resulting in a stable population. Survival during the post-fledging period of dependence and the first year of independence in the Western Slaty-Antshrike exceeded estimates for Northern Hemisphere species. This difference appears to be associated with the extended post-fledging parental care, delayed dispersal, low costs of dispersal, and the less seasonal environment of antshrikes.     

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