Ventilation rate and copulation in Octopus vulgaris

Typically three events occur when Octopus vulgaris Cuvier mates. The male inserts its hectocotylus into the mantle cavity of the female; the hectocotylus and the bodies of the male and female become quiescent; and the female increases its ventilation rate about 2.5 times, on the average, above its precopulatory base rate. It is hypothesized that the increased ventilation rate of the female provides the stimulus to the male to transfer its spermatophore to the oviduct. Since the insertion of the hectocotylus and the female's increased ventilation rate may be dissociated, it is suggested that copulation be defined as the attachment of the hectocotylus to the oviduct. A conditioned ventilation-rate increase in the female was observed with repeated testing. Observation of arching and pumping movements of the male as well as measurements of its ventilation rate before and during copulation indicate that only a few spermatophores are transferred per copulation.     

Choice in Varroa jacobsoni Oud. between honey bee drone or workerbrood cells for reproduction

Summary The effect of variation in copulation duration on sperm precedence (P₂) of the last male was investigated in a non-territorial libellulid using irradiated-male techniques and observations in a large field enclosure. Copulation duration varied between 6 min and more than 1 h (X = 23.19 min ± 12.74). In the clutch deposited immediately after the copulation, most copulation durations resulted in high sperm precedence for the last male. Several experimentally interrupted short matings still resulted in 100% precedence. However, most interrupted copulations of 10 min or less resulted in a low P₂. After long, uninterrupted copulations, P₂ remained high in clutches that were collected up to 9 days after the mating. After interrupted short copulations, P₂ fluctuated or remained at a low level from the first clutch onwards. This indicates that sperm mixing probably starts within 24 h and that after a long copulation virtually no rival sperm is left in the female's sperm storage organs. As most natural copulations are long, complete removal of rival sperm seems to be common in S. danae. Only 3 f 17 variables investigated explained a significant amount of the variation in copulation duration. Copulations shortened with increasing temperature and for each additional mating of the male. The strongest effect however, was due to time of day: early morning copulations lasted longer than midday copulations. Adjustment of the copulation duration to the decreasing chance of obtaining additional matings was found to be the most likely of five explanatory hypotheses that were evaluated.     

Information content of female copulation calls in wild long-tailed macaques (<Emphasis Type="Italic">Macaca fascicularis</Emphasis>)

Primates are unusual in that many females display sexual signals, such as sex skin swellings/colorations and copulation calls, without any sex role reversal. The adaptive function of these signals remains largely unclear, although it has been suggested that they provide males with information on female reproductive status. For sex skin swellings, there is increasing evidence that they represent a graded signal indicating the probability of ovulation. Data on the functional significance of copulation calls are much scarcer. To clarify the information content of such calls, we recorded copulation calls in wild long-tailed macaques (Macaca fascicularis) and analysed the structure of these calls during the ovarian cycle. Specifically, we correlated selected call parameters with the female oestrogen to progestogen ratio (obtained from faecal samples), which are known to be elevated during the female's fertile phase. In addition, we ran a general linear mixed model for these call parameters, testing factors (cycle phase, occurrence/absence of ejaculation, male dominance status, occurrence/absence of mate guarding) which potentially influence female copulation calls in primates. Our results show that copulation calls of female long-tailed macaques signal mating outcome and rank of the mating partner, but not female reproductive status. They also show for the first time on primates that copulation calls can convey information on whether a female is mate guarded or not. We suspect that the function of these calls is manipulation of male mating and mate-guarding behaviour and that in this way the degree of sperm competition and ultimately male reproductive success is influenced.     

Direct and indirect estimates of sperm precedence and displacement in the dragonfly Sympetrum danae (Odonata: Libellulidae)

Summary The non-territorial dragonfly Sympetrum danae shows a high degree of post-copulatory mate-guarding, which suggests precedence of the last male's sperm. Irradiated male techniques revealed that the last male that mates with a female fertilizes an average of 95% (100% in 28% of the cases) of the clutch laid immediately after copulation, irrespective of any previous matings. Sperm volume estimates in both sexes and sperm opacity changes in females at 7 stages of copulation and oviposition indicated that males remove 41–87% of previously deposited sperm from the female's storage organs during the first 5 min of the copulation. Ejaculation takes place during the remaining 10–15 min. We maintain that indirect estimates of precedence in libellulids, based on sperm volume changes, always understimate reality because of the supplementing effect of a first-in, last out mechanism. Because males are skilled at achieving sperm precedence, they are forced to be good guards as well, since a risk of take-overs exists. Females are believed to benefit from contact guarding because it results in a lowered risk of male harassment and predation as well as a lower energy expenditure during oviposition.     

Variation in copulation duration in Mnais pruinosa pruinosa Selys (Odonata: Calopterygidae)

Summary The Japanese calopterygid damselfly Mnais pruinosa pruinosa shows three distinct copulation durations (Siva-Jothy and Tsubaki 1989). A variety of factors which might influence copulation duration were investigated. Results indicate that the temperature in the vicinity of copulating pairs, the rate at which males encountered receptive females, territory quality, the rate of agonistic interactions between males, female gravidity and female willingness to oviposit have no effect on copulation duration. The most significant factor was the location of the site at which males captured their mates. We suggest that under natural conditions the location of the capture site provides males with reliable, indirect information about a female's intention to oviposit: it is important for males of this species to acquire such information since sperm precedence, and therefore reproductive success, is in part dependant on the interval experienced by females between copulation and subsequent oviposition. The results are discussed and it is suggested that the three observed mate-securing tactics of M.p. pruinosa are facultative and information-dependant.     

Movement and environmental preferences of Greenland sharks (<Emphasis Type="Italic">Somniosus microcephalus</Emphasis>) electronically tagged in the St. Lawrence Estuary,Canada

Three Greenland sharks (Somniosus microcephalus) were tagged with electronic tags, in Baie St. Pancrace, St. Lawrence Estuary, Quebec, Canada. One shark was tagged on 23 July 2004, with an acoustic telemetry tag. Two sharks were each tagged with a pop-up satellite archival tag (PSAT) on 27 August 2004. Two of the sharks remained in or close to the bay, one for 47 days and the other for at least 66 days. The third shark left the bay immediately after tagging on 27 August 2004. This shark entered the main channel of the St. Lawrence Estuary, and had moved 114.9 km upstream by 1 November 2004 when the tag reported to ARGOS satellites. The tags provided a total of 179 days of data on the movement and environmental preferences of Greenland sharks in the St. Lawrence Estuary. Sharks that reported depth and ambient water temperature data from the bay showed significant diel differences in depth preferences and corresponding ambient temperatures. The sharks remained near the bottom of the water column during the day and displayed increased vertical movements at night. The shark that resided in the main channel did not show this pattern, but generally remained at depths between 325 and 352 m. Sharks in the bay experienced water temperatures that ranged from −1.1 to 8.6°C at depths from 0 to 67 m. In the main channel the shark experienced temperatures that ranged from 1.0 to 5.4°C at depths from 132 to 352 m. This is the first report of numerous Greenland sharks inhabiting shallow near shore bays during summer and autumn.     

Fertilisation success in the fly Dryomyza anilis (Dryomyzidae): effects of male size and the mating situation

Male fertilisation success in relation to male size and the mating situation (ordinary pair formation with a single, nonvirgin female vs. take overs) was examined in the fly Dryomyza anilis. In ordinary matings, large males achieved higher fertilisation success than small ones when they were the second to mate with the female. Take overs differ from ordinary pair formation in that the second male experiences intensified sperm competition. This is because in take overs the female is not able to discharge any of the sperm inseminated by the first male as she usually does before a new mating. Compared with ordinary matings, take overs reduced the fertilisation success of the second male by 8–10%, whereas that of the first male was 7–14% higher in take overs. Even though the intruder was always larger than the paired male his superior fertilisation success did not compensate for the effect of the sperm already present in the female. In D. anilis, males can increase their fertilisation success by tapping the female's external genitalia with their claspers or having several copulation bouts per mating. Thus, in a take over, the intruder could respond to the intensified sperm competition by performing more tapping sequences per copulation bout or more copulation bouts per mating. In matings observed in the wild, males performed more tapping sequences after a take over than after pair formation with a single female, although the difference was not significant. The results show that there are differences in fertilisation success between males of different size. In addition, different mating situations can result in considerable variation in the fertilisation success of an individual male. Higher fertilisation success for the first male after a take over may be significant, in particular, for the reproductive success of small males, which frequently lose their females to large males.     

Sperm transfer,displacement and precedence in Ischnura graellsii (Odonata : Coenagrionidae)

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     

Observations on the short-term movements and behaviour of whale sharks (Rhincodon typus) at Ningaloo Reef, Western Australia

The short-term movements and behaviour of whale sharks (Rhincodon typus Smith, 1828) during March 1994 and April 1997 are reported from data collected by acoustic tracking and archival tags at Ningaloo Reef on the north west coast of Western Australia. Sharks were tracked for up to 26 h and generally swam slowly at ≃0.7 m s⁻¹ parallel to the reef edge; occasionally they swam in a wide arc adjacent to passes in the reef. All tracked sharks made regular dives through the water column, mostly from the surface to near the bottom. These dives did not appear to be related to hydrographic features, and the sharks were probably searching the water column for food. Most sharks were accompanied by other fishes, usually the golden trevally Gnathanodon speciosus. Received: 19 January 1999 / Accepted: 22 June 1999     

Ultrasonic tracking of Greenland sharks,<Emphasis Type="Italic"> Somniosus microcephalus</Emphasis>, under Arctic ice

Six Greenland sharks, Somniosus microcephalus (Bloch and Schneider, 1801), 190–355 cm fork length, were tracked under land-fast sea ice off northern Baffin Island (73.2°N; 85.3°W) between 16 and 28 May 1999, using ultrasonic telemetry. The sharks were tracked continuously for periods of 5.5–13.0 h, with the tracks of two individuals lasting 31.4 and 42.8 h, respectively, each with an interval when the track was lost. Several sharks dove after release and moved along the ocean bottom for the duration of the tracking period, while others varied their movements regarding course and depth. Two sharks made repeated visits to within 11 m of the ice–water interface from deeper water. The tracked sharks exhibited no apparent depth or temperature preferences, and pooled data indicated that sharks remained deep during the morning and gradually moved into shallower depths through the afternoon and night. Rates of descent (average=0.099 m s^–1) were significantly greater than rates of ascent (average=0.058 m s^–1) for all sharks, and the average rate of horizontal movement over ground was estimated as 0.215 m s^–1. Based on the movements of tracked sharks and information contained in the literature, S. microcephalus may prey on seals in areas covered by land-fast sea ice.Communicated by J.P. Grassle, New Brunswick     

Quantitative analysis of aggregation behavior in juvenile blacktip sharks

Aggregation patterns of a population of juvenile blacktip sharks (Carcharhinus limbatus) within a summer nursery area were examined over a period of 3 years using an array of acoustic monitors. Individually coded acoustic tags were surgically implanted into 33–40 neonate sharks each year from 2000 to 2002 and their presence monitored continuously by the acoustic array. Data from the acoustic monitors was processed to estimate the center of activity location of each tagged shark every 30 min. Nearest neighbor analysis of shark locations revealed that sharks aggregated within the nursery in all years of the study. Sharks were aggregated most commonly during the early and late summer months (June, October and November) and became less common through the middle of the study period each year (July–September). Temporal periodicity within the data revealed a strong diel pattern with sharks aggregating during the day and dispersing at night, except in June when aggregations often persisted through the night. Aggregations were generally restricted to the mid and northern sections of the study site. Reasons for aggregations may include predator avoidance or improved feeding efficiency.     

Female choice of multiple male criteria in guppies: interacting effects of dominance,coloration and courtship

Summary I experimentally examined the relative importance of social dominance, color patterns, and courtship behavior in male mating and reproductive success in the guppy Poecilia reticulata. Female choice of males is based on a complex set of behavioral and morphological traits. The results of 59 paired-male one-female visual choice and mating trials showed that male mating success was positively correlated with dominance, courtship intensity, and male coloration. Only dominant males engaged in full copoulations, and they sired two-thirds of the broods. An analysis of the paternity of broods and results of mating trials showed that a female's visual response when the sexes are separated by a glass partition is a good predictor of a male's reproductive success when the partition is removed and they are allowed to mate. A canonical correlation analysis of male behavioral and morphological traits indicated that female visual response and male mating success were positively correlated with male courtship and with agonistic behavior. However, the relative importance of color varied. Carotenoid and iridescent spots were important both in attracting the female's attention and in enhancing male mating success. Melanins were not correlated with either mating success or female response. There was a relatively low correlation (48%) between male behavioral and morphological variables and female response variables (full copulation and female visual response). These results suggest that female choice is subtle, and is based on a complex suite of male behavioral and morphological traits as well as on competitive interactions among males.     

Tiger shark (Galeocerdo cuvier) abundance and growth in a subtropical embayment: evidence from 7 years of standardized fishing effort

The tiger shark (Galeocerdo cuvier Peron and Lesueur 1822) is a widely distributed predator with a broad diet and the potential to affect marine community structure, yet information on local patterns of abundance for this species is lacking. Tiger shark catch data were gathered over 7 years of tag and release research fishing (1991–2000, 2002–2004) in Shark Bay, Western Australia (25°45′S, 113°44′E). Sharks were caught using drumlines deployed in six permanent zones (~3 km² in area). Fishing effort was standardized across days and months, and catch rates on hooks were expressed as the number of sharks caught h⁻¹. A total of 449 individual tiger sharks was captured; 29 were recaptured. Tiger shark catch rate showed seasonal periodicity, being higher during the warm season (Sep–May) than during the cold season (Jun–Aug), and was marked by inter-annual variability. The most striking feature of the catch data was a consistent pattern of slow, continuous variation within each year from a peak during the height of the warm season (February) to a trough in the cold season (July). Annual growth rates of recaptured individuals were generally consistent with estimates from other regions, but exceeded those for populations elsewhere for sharks >275 cm fork length (FL), perhaps because mature sharks in the study area rely heavily on large prey. The data suggest that (1) the threat of predation faced by animals consumed by tiger sharks fluctuates dramatically within and between years, and (2) efforts to monitor large shark abundance should be extensive enough to detect inter-annual variation and sufficiently intensive to account for intra-annual trends.     

The scent of dominance: female field crickets use odour to predict the outcome of male competition

Pheromones may convey information about mate quality and social status. In the field cricket Gryllus integer, females mount the males for copulation, such that males cannot coerce females to mate. We examined whether virgin G. integer females preferred the scent of potentially dominant males to that of subordinate males. First, we collected pheromones by confining males on filter paper. Next, we offered filter paper from each of two size-matched males and control paper to females that had never been exposed to males, and measured the time spent by the female on each kind of paper. Finally, dominance status of the males in each size-matched pair was determined by pitting the two males against one another in agonistic contests. When offered filter paper from subsequently dominant versus subsequently subordinate males, females spent more time on the paper from the dominant male than the subordinate male, and much less time on control paper. Thus, pheromones may inform female G. integer about a male's potential to achieve dominant social status. Male pheromones were also associated with the female's tendency to mount a male. In contrast to cockroaches, where females prefer the scent of subordinate males (presumably to avoid risk of injury), female crickets prefer the scent of potentially dominant males and are more likely than males to wound their mating partners.     

Movement patterns of young Caribbean reef sharks, Carcharhinus perezi, at Fernando de Noronha Archipelago, Brazil: the potential of marine protected areas for conservation of a nursery ground

The movement patterns and long-term site-fidelity of primarily juvenile Caribbean reef sharks, Carcharhinus perezi, were investigated using tag-recapture and automated telemetry at an insular nursery area, the Fernando de Noronha Archipelago, Brazil. Of the 143 externally tagged juvenile sharks (<110 cm), 22 (15.3%) were recaptured between 0 and 5 km from the site of tagging after 5–800 days at liberty, suggesting some site-fidelity in young individuals of this species. Site-fidelity and movement patterns of ten juvenile sharks ranging from 78 to 110 cm total length (TL) and one opportunistically captured adult female (224 cm TL) were also investigated for periods of up to 2 years with an array of automated telemetry receivers. Tagging and telemetry data from both inside and outside a marine protected area (MPA) show that shark abundance and activity is greatest along the part of the archipelago’s coastline least disturbed by human activity. Telemetry tracking also showed that juvenile reef sharks demonstrated a high degree of site-fidelity and occupied specific locations along the coast throughout the year, with some evidence of an increase in activity space with ontogeny. Sharks appeared to range more widely at night and there were no seasonal variations in habitat use. Our results suggest that MPAs may be a useful conservation tool to protect young C. perezi and potentially other reef-dwelling carcharhinid sharks during their early life history.     

Mating behavior and sperm competition in the fly,Dryomyza anilis

Summary Mating behavior in D. andis was studied both in nature and in the laboratory. Each mating in nature consisted of several repeated copulation and oviposition bouts during which the male stayed mounted. One copulation bout included an intromission in the beginning followed by several tapping sequences. During one tapping sequence the male touched the external genitalia of the female with his claspers several times. The experimental matings with irradiated sterile males contained one copulation bout after which the female laid her eggs. In these matings the percentage fertilization for the last male to mate with the female increased with increasing number of tapping sequences. The percentage fertilization was affected positively also by the time between the end of the copulation bout and the start of oviposition, but the time spent in the copulation bout was not important. Furthermore, there was a negative correlation between the length of intromission and the percentage fertilization. After the copulation bout, the percentage fertilization was higher among the first eggs laid. This difference between the first and later eggs laid increased with the increasing number of tapping sequences, suggesting that copulating and ovipositing in several bouts further increase the male percentage fertilization. In matings recorded in nature, both the number of tapping sequences and the number of copulation and oviposition bouts varied greatly. Of the variables recorded during these matings, individual size and the number of eggs the female contained in the beginning of a mating affected the number of copulation bouts and the number of tapping sequences. Males also mated differently with females containing immature or mature eggs. The male behavior resulting in sperm precedence and the variability of matings in nature in relation to male sperm competition is discussed.     

Mixed reproductive strategy and mate guarding in a semi-colonial passerine,the swallow Hirundo rustica

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.     

Why do male Callosobruchus maculatus beetles inseminate so many sperm?

Male Callosobruchus maculatus F. (Coleoptera: Bruchidae) inseminate more sperm than females can effectively store in their spermathecae. This study examines the adaptive significance of excess sperm transfer by measuring components of male and female reproductive success in response to manipulating the number of sperm inseminated. The number of sperm transferred during copulation was reduced from 56,000 ±4,462 to 8,700±1,194 by sequentially mating males to virgin females. Reducing the number of sperm inseminated by the first male to mate had no effect on the extent of sperm precedence, but reducing the number of sperm inseminated by the second male resulted in a significant reduction in the extent of sperm precedence. When large numbers of sperm are inseminated the remating refractory period of females is increased. These results indicate that males transferring large numbers of sperm during copulation have a two-fold advantage at fertilization; they are more effective at preempting previously stored sperm and they are likely to father more offspring by delaying the time of female remating. The transfer of excess sperm does not appear to serve as nonpromiscuous male mating effort; the number of eggs laid, their fertility and the subsequent survival of zygotes were unaffected by manipulating the number of sperm inseminated. The underlying mechanisms of sperm precedence were also examined. Simple models of sperm displacement failed to accurately predict the patterns of sperm precedence observed in this species. However, the results do not provide conclusive evidence against the models but rather serve to highlight our limited understanding of the movement of sperm within the female's reproductive tract.     

Predicting occurrence of juvenile shark habitat to improve conservation planning

Fishing and habitat degradation have increased the extinction risk of sharks, and conservation strategies recognize that survival of juveniles is critical for the effective management of shark populations. Despite the rapid expansion of marine protected areas (MPAs) globally, the paucity of shark‐monitoring data on large scales (100s–1000s km) means that the effectiveness of MPAs in halting shark declines remains unclear. Using data collected by baited remote underwater video systems (BRUVS) in northwestern Australia, we developed generalized linear models to elucidate the ecological drivers of habitat suitability for juvenile sharks. We assessed occurrence patterns at the order and species levels. We included all juvenile sharks sampled and the 3 most abundant species sampled separately (grey reef [Carcharhinus amblyrhynchos], sandbar [Carcharhinus plumbeus], and whitetip reef sharks [Triaenodon obesus]). We predicted the occurrence of juvenile sharks across 490,515 km² of coastal waters and quantified the representation of highly suitable habitats within MPAs. Our species‐level models had higher accuracy (? ≥ 0.69) and deviance explained (≥48%) than our order‐level model (? = 0.36 and deviance explained of 10%). Maps of predicted occurrence revealed different species‐specific patterns of highly suitable habitat. These differences likely reflect different physiological or resource requirements between individual species and validate concerns over the utility of conservation targets based on aggregate species groups as opposed to a species‐focused approach. Highly suitable habitats were poorly represented in MPAs with the most restrictions on extractive activities. This spatial mismatch possibly indicates a lack of explicit conservation targets and information on species distribution during the planning process. Non‐extractive BRUVS provided a useful platform for building the suitability models across large scales to assist conservation planning across multiple maritime jurisdictions, and our approach provides a simple for method for testing the effectiveness of MPAs.     

Pre-mating sperm removal in the bushcricket Metaplastes ornatus Ramme 1931 (Orthoptera, Tettigonoidea, Phaneropteridae)

Summary Mating in the bushcricket Metaplastes ornatus Ramme 1931 entails a number of peculiar genital couplings that precede the transfer of the large spermatophore. During these phase-I couplings, the male introduces his specially structured subgenital plate into the female's genital chamber, performs back-and-forth movements, and turns her genital chamber inside out when he withdraws, whereupon the female carefully cleans her everted genital chamber with her mouthparts. During the last coupling (phase II) the male's subgenital plate is not introduced but the large spermatophore, which averages 22% of a male's body weight, is transferred. Counts of sperm in the spermathecae of females suggested that the phase-I couplings, which occur prior to spermatophore transfer, function to remove, or at least to reduce, the sperm of a female's previous mates. The form of the keel of the male's subgenital plate, its position within the female's genital tract during phase-I couplings, and the back-and-forth movements suggest that the male may stimulate release of sperm from the female's spermatheca by a mechanism similar to fertilization as eggs pass through the genital chamber during oviposition.