The post-embryonic development and life history of the millipede, Ommatoiulus moreletii (Diplopoda: Iulidae), introduced in south-eastern Australia

The life cycle of Ommatoiulus moreletii consists of an egg, pupoid and up to sixteen stadia. Sexes are differentiated at the sixth stadium. Males may mature at any moult from the eighth to the twelfth stadia, but most are mature by the tenth or eleventh. Maturation is more difficult to determine in females but seems similar to the males with respect to stadial age. O. moreletii is periodomorphic.
Field studies were made of O. moreletii in an open grassland and a dry sclerophyllous woodland in South Australia. Females matured their eggs in late summer-autumn. They mated and oviposited during autumn-winter. After one year, O. moreletii was in the seventh, eighth or ninth stadium. After two years, the tenth or eleventh stadia were reached and after three years the twelfth or thirteenth. The moulting of individuals older than about one year was confined to moults in (1) spring and (2) summer. Adult males were mature from summer to spring and intercalary from spring to summer.     

Notes on the life-history and ecology of Tachypodoiulus niger (Diplopoda, Iulidae) in Britain

In some iulids females can moult several times after first attaining maturity but males cannot. Males of Tachypodoiulus niger (Leach) are exceptional in that they can moult, but they lose their functional intromittent organs in the process, regaining them after a second moult. The process can be repeated, a series of functional males alternating with nonfunctional males. This extension of the life and the consequent increase in number of segments led Verhoeff (1928) to postulate that the ancestral millipede was short-bodied and that many-segmented forms were derived from it. We think that this extension of life is of ecological rather than phylogenetic significance–adapting the species to disperse widely to scattered habitats. In particular, the extension of life of males as well as of females may ensure a reasonable sex ratio in those areas where the species is least dense.
In Britain, Tachypodoiulus niger lays eggs in spring which reach the fourth and fifth stadia by their first winter and the seventh, eighth and ninth stadia by their second winter. Maturity is usually attained by males in the eighth stadium but occasional specimens mature at stadium seven and others defer maturity until the ninth. Both sexes can proceed to the fourteenth stadium but adults in Britain usually belong to two generations of two and three years old, divided mainly between the eighth, ninth and tenth stadia. Details are given of a large collection made by the late Dr Scott of animals taking refuge in his house at Henley-on-Thames. Animals in Britain appear to have a similar life history to those in Germany as described by Verhoeff, but effective comparison has depended on correcting and re-interpreting some of Verhoeff's data.     

Reproductive tactics of the ringed plover Charadrius hiaticula

Reproductive tactics of ringed plovers Charadrius hiaticula were studied at three localities in SW Sweden during five seasons. The usual clutch size is four, but removal experiments showed that females can produce five eggs in succession, with similar intervals between all eggs. High predation made mean breeding success per clutch low, 6.3% of eggs resulting in fledged young. Replacement clutches were common, and many pairs laid again after rearing their first brood to fledging. Egg laying spanned three months, much longer than for other waders in this region. Between years, reproductive success varied unpredictably with time of the season, but averaged over several years, the expected success was low and similar for the different parts of the breeding season. Chicks from late clutches had similar survival and recruitment as others. Because of the long breeding season and high rate of nest failure a female may produce up to five clutches of four eggs per season, containing in total about 3.7 times her own mass. Yearly local survival of adult females and males was estimated to 84.6 and 88.6%, respectively. Ability to produce many clutches with similar expected success throughout the season favours a long reproductive period, sometimes leading to double-brooding. Possible life-history trade-offs are discussed.     

The life history and post-embryonic development of "Spirobolus" bivirgatus (Diplopoda: Spirobolida) on Aldabra, Western Indian Ocean

Spirobolus bivirgatus, recently known as Mystalides bivirgatus, passes through 15 stadia. These were differentiated by the ocular field method. Development is anamorphic from stadia I to VIII and epimorphic from stadia IX to XV. Maturity is reached in a few males in the eighth stadium but more normally it occurs in the ninth to twelfth stadium. Females are mature in the ninth to fifteenth stadium. Adult males possess fully developed gonopods as well as soft pads on the ventral surface of the tarsi. The pads are illustrated with scanning electron micrographs. Males die after breeding but females probably breed in three or more successive years. Eggs are laid during the wet season, December to April, and these reach maturity within the second or, possibly, the first year of growth. The density, distribution and food of S. bivirgatus is briefly described.     

Turnover and dispersal in a Peregrine Falco peregrinus population

In south Scotland, most Peregrines returned to the same territories to breed in successive years, though a few females changed territory from one year to the next.
Annual mortality among breeding birds was at most 9% among females (or 11% in both sexes combined). There may have been considerable annual variation, however, and excluding one exceptional year out of five reduced the estimate for females to 7%. These estimates are maxima, but are still considerably lower than those obtained from ring recoveries of dead birds reported by members of the public.
Among trapped birds, four males first bred at age two years, one at three and another at four or five; two females first bred at one year, 13 at two years old and one at three. Five other females which were seen to be in first-year plumage but were not trapped, also laid eggs, and 12 other such paired females held territory but did not lay. Only one paired male held territory in first-year plumage.
In their movements between natal and breeding territories, some females moved further than males, with median distances of 83 and 58 km respectively. In addition, of birds trapped breeding in the study area, a greater proportion of the males than of the females had been born locally, despite an equal sex ratio among fledglings; this was also consistent with a greater dispersal of females. In general, Peregrines made much longer movements in their first year of life than subsequently. Movements were in any direction.     

The millipedes of a Cheshire wood

The density and surface activity of the millipedes in a quarter of an acre of a sycamore ash wood are described. Seven species were extracted by Tullgren funnels from samples of soil and litter over five years and were also caught in pitfall traps during a further two years; four other species occurred occasionally in the traps. Each square metre of the site supported 100 individuals over the winter, rising to 300 in the summer. Of these, 85 % belonged to three species, Iulus scandinavius, Polydesmus angustus and P. denticulatus .
Male I. scandinavius become adult in either the ninth, tenth or eleventh stadium; females in the tenth and eleventh. Eggs are laid in spring and these take three years to become adults which breed and then die. The majority of Polydesmus spp. in the samples are young belonging to the first six of the eight free-living stadia. The adults fell into the traps in the summer and newly emerged young appear in the samples at this time. They overwinter in their first year mainly as fifth stadia; some might reach maturity (eighth stadia) in the summer following, but it is not certain that they could breed at this time.
The pattern of dispersion of lulus is fairly even and is correlated with the distribution of leaf litter but the Polydesmus spp. are highly aggregated. All stadia of lulus fall into the traps but only the last two of Polydesmus . The aggregation of Polydesmus spp. appears to be correlated with the relative inactivity of the younger stadia. The estimates of density of Polydesmus spp. are unreliable because of their aggregation but those of lulus have determinable limits and it is possible to derive rough though meaningful standing crop, production and life-table data. The overwintering standing crop of lulus consists of the survivors of three generations of 5, 2.2 and 1.4% of the original eggs laid; it has a fresh weight of about 1.25 g and a production in the order of 1.5–2.5 g/m².     

Dispersal, recruitment and migratory behaviour in a hawksbill sea turtle aggregation

We investigated the dispersal, recruitment and migratory behaviour of the hawksbill sea turtle ( Eretmochelys imbricata ), among different life-history stages and demographic segments of the large hawksbill turtle aggregation at Mona Island, Puerto Rico. There were significant differences in both mitochondrial DNA (mtDNA) haplotype diversity and haplotype frequencies among the adult males, females and juveniles examined, but little evidence for temporal heterogeneity within these same groups sampled across years. Consistent with previous studies and the hypothesis of strong natal homing, there were striking mtDNA haplotype differences between nesting females on Mona Island and nesting females in other major Caribbean rookeries. Breeding males also showed strong, albeit weaker, genetic evidence of natal homing. Overall, Bayesian mixed-stock analysis suggests that Mona Island was the natal rookery for 79% (65–94%) of males in the aggregation. In contrast, the Mona Island rookery accounted for only a small subset of the new juvenile recruits to the foraging grounds or in the population of older juvenile hawksbills turtles on Mona. Instead, both new recruits and the older juvenile hawksbill turtles on Mona more likely recruited from other Caribbean rookeries, suggesting that a mechanism besides natal homing must be influencing recruitment to feeding habitats. The difference in the apparent degree of natal homing behaviour among the different life-history stages of hawksbill turtles at Mona Island underscores the complexity of the species' life-history dynamics and highlights the need for both local and regional conservation efforts.     

Maturational changes in patterns of association in male and female humpback whales, Megaptera novaeangliae

The patterns of association of juvenile male and female humpback whales, Megaptera novaeangliae , in the southern Gulf of Maine were studied for evidence of maturational changes. Both males and females became less solitary with age. In males, time spent alone changed from a mean of 55.8% of observations at age one to 26.8% at age six. Females were alone in a mean of 49.9% of observations at age one, but in only 20.5% by age six. However, females that produced calves at five, six or seven were associated with no whales but the calf in 73.8% of observations. Males exhibited a clear age-related trend of increasing associations with adults, notably with adult females which constituted approximately 80% of the associates of males aged six years or more. Females showed a similar trend of increasing associations with adults of both sexes. Tests of association data for whales of known age with similar data for adults of the same sex showed that the association patterns of young males and females became statistically indistinguishable from those of adults by the ages of five and four, respectively. The data suggest that the observed changes in social behaviour are closely linked to the attainment of sexual maturity and preparation for adult roles. The different patterns of males and females after maturity may reflect differing reproductive and life-history strategies.     

Millipedes faced with drought: the life cycle of a Mediterranean population of Ommatoiulus sabulosus (Linnaeus) (Diplopoda,Julida, Julidae)

Growth, development and life-cycle duration of the millipede Ommatoiulus sabulosus (f. aimatopodus) were studied in a Mediterranean shrubland of southern France and compared with previous data from northwest Europe. Changes in the proportions of stadia during the course of the year were analysed in several generations. The results show that stadia VII and VIII are consistently reached after the first year of growth, and stadia IX and X after the second year. First reproduction may occur at the age of two years in males reaching maturity at stadium X, but not until the age of three in those reaching maturity at stadia XI and XII. Reproduction cannot occur until at least the age of three in females, which carry mature eggs from stadium XI onwards. In comparison with more northern populations, life-cycle duration is not shorter in the Mediterranean population but there are marked differences in its phenology: the breeding period is in autumn, so that juveniles of stadia II to VI are never faced with the summer drought, and larger individuals are mostly inactive in summer; moreover, all individuals moult once every winter. The results illustrate how julid millipedes of humid temperate regions could respond to higher temperatures and drier summer conditions in the context of climate change.     

Effect of duration of larval development on sexual competence in young adult male Diploptera punctata

ABSTRACT. The duration of larval development was 27% longer, adult weight was 10% greater, and adult head capsule was 2% wider in Diploptera punctata (Eschscholtz) males with four larval stadia compared with those with three larval stadia. Four-instar males transferred significantly more sperm than three-instar males 8, 14 and 28 days after adult eclosion, but there was no difference in the length of sperma-tophore transferred by three- and four-instar males at these ages. Eight-day-old four-instar males were more successful than 8-day-old three-instar males in fertilizing complete batches of eggs. For both three- and four-instar males, males that mated when they were older (measured in days after adult eclosion) transferred more sperm and larger spermatophores than younger males. Body size did not have a significant effect on the number of sperm or size of spermatophore transferred by males.     

The Cooperative Breeding System of the Red-cockaded Woodpecker

The breeding system of the red-cockaded woodpecker is described based on data collected over six years from a population of 500 marked individuals in the Sandhills of North Carolina. Male-female pairs were the most common social unit (59%), but 30% of social units contained one or more adult helpers, and 11% consisted of solitary males. Helpers were almost exclusively male: 27% of males remained in their natal group as helpers for at least one year, whereas only four (1%) females did. Most breeding females remained as breeders in the same group from one year to the next (56%), but a surprising number (12%) moved to another group. Many movements were related to incest avoidance or mate death, but 39% involved deserting a mate, usually following successful reproduction. We suggest that females sometimes are forced from groups by immigrants or other group members. The median distance of movements by adult females was only 1.3 km. In contrast to females, no breeding males switched groups. Survival of both breeding (76%) and helper (80%) males was higher than that of breeding females (69%). Males exhibited two distinct life-history strategies. Some remained as helpers on their natal territory for one or more years, and became breeders by inheriting breeding status in the natal group (17% per year) or by replacing a deceased breeder in a nearby group (13% per year, median distance moved 1.0 km). Other males dispersed from their natal group permanently during their first year. Some of these males were floaters at age one year, others were solitary, and a few became helpers in a non-natal group, but many were breeders. In contrast to males that first functioned as helpers, those that dispersed after fledging moved long distances (median dispersal distance 4.5 km), longer even than dispersing female fledglings moved (median distance 3.2 km). The habitat saturation model of the evolution of cooperative breeding is based on selection between the two life-history strategies exhibited by male red-cockaded woodpeckers. The model therefore may be tested directly with this species. Another indication that this model is appropriate for this species is the existence of a resource (cavity trees) that might provide an ecological basis for habitat saturation.     

Social influences on nymphal development in the cockroach, Diploptera punctata

Solitary male nymphs of the cockroach Diploptera punctata (Eschscholtz) (Blattaria: Blaberidae) took significantly longer to reach adulthood than males paired with either a male or female nymph or grouped with four other male nymphs since birth. When isolated throughout nymphal development, 15.8% of males passed through 3 stadia before adult eclosion, and the remainder went through 4 stadia. In contrast, 61.3% of paired males became adults in 3 stadia. Males need not, however, be isolated or paired for the entire nymphal period to express isolated or paired patterns of development. About 60% of males paired in just the first stadium or its initial 9 days became adults in 3 stadia, and only 20.4% of males isolated in the first stadium and the first 3 days of the second reached adulthood within 3 stadia. Although the first stadium was a critical period in which social condition determined the course of future development, analyses of covariance showed that isolated males gained less weight than paired ones, not only in the first stadium, but in the second as well. Moreover, the degree of growth of a male in the second stadium, measured as either weight gain or relative growth rate, did not depend on the male's social condition in the first stadium, because isolated second-instar males grew less than paired ones, even when both sets of insects had been paired in the first stadium. Female nymphal development, unlike that of males, was not greatly affected by social factors.     

Life history and distribution of the small south-western Atlantic octopus, Octopus tehuelchus

Although this species has been reported from shallow waters down to 90 m depth, knowledge is almost entirely based on intertidal samples. In this study both intertidal and subtidal samples were taken during 1982–1987, in northern San Matias Gulf (41° S, 63° 30' W). This is a large-egged (eggs: 9–12 mm long ×–5 mm wide, stack of 4–6 mm long) and small-sized (up to 150 g) octopus. Egg laying occurs between autumn and winter. Embryonic development takes about four months (water temperature:–19°C). Large hatchlings (DML: 5·64 mm, TL: 14·23 mm, TW: 0·139 g) emerge over spring and early summer, and development is direct. Maximum size is reached after 17–18 months; mating takes place in summer. Females reduce their feeding activity when they reach maturity, and cease eating while brooding. Mean life-span is two years, but some individuals (mostly females) may live up to three years. Females approaching the beginning of the brooding period move to the subtidal zone, where males outnumber females until the end of summer and females (mostly brooders) then outnumber males. In the intertidal zone sex ratio was 1:1 from December to late March, but in April males outnumber females.
These life-history traits are compared with those of other large-egged octopuses and are discussed in relation to environmental conditions prevailing in the San Matias Gulf.     

Spatial genetic analysis and long-term mark-recapture data demonstrate male-biased dispersal in a snake

Dispersal is an important life-history trait, but it is notoriously difficult to study. The most powerful approach is to attack the problem with multiple independent sources of data. We integrated information from a 14-year demographic study with molecular data from five polymorphic microsatellite loci to test the prediction of male-biased dispersal in a common elapid species from eastern Australia, the small-eyed snake Rhinoplocephalus nigrescens. These snakes have a polygynous mating system in which males fight for access to females. Our demographic data demonstrate that males move farther than females (about twice as far on average, and about three times for maximum distances). This sex bias in adult dispersal was evident also in the genetic data, which showed a strong and significant genetic signature of male-biased dispersal. Together, the genetic and demographic data suggest that gene flow is largely mediated by males in this species.     

Age and growth analysis of the shortfin mako, <Emphasis Type="Italic">Isurus oxyrinchus</Emphasis>, in the western and central North Pacific Ocean

We determined the age and growth rates of male and female shortfin makos, (Isurus oxyrinchus), from the western and central North Pacific Ocean. Growth band pairs were counted on half-cut vertebral centra using a shadowing method. In this method, we focused on the ridges on the surface of the centra, consisting of a convex and concave structure. After comparing four enhancing methods, we decided on the use of shadowing method for aging. Vertebrae from 128 males and 147 females were examined. The centrum edge analysis suggested annual band pair formation. Von Bertalanffy growth curves were fitted separately to the length-at-age data for males and females with birth length fixed. Until approximately 7 years of age, both sexes showed similar growth rates; thereafter, males showed a significantly slower growth rate compared to females. It was suggested males and females mature at approximately 6 years and 16 years, respectively. These life-history characteristics suggest relatively low productivity for this species, which agrees with reports on populations in other geographic regions.     

Multiple mating reduces male survivorship but not ejaculate size in the polygamous insect Stenomacra marginella (Heteroptera: Largidae)

In polygamous species, successful males should be able to inseminate multiple females, to defeat sperm from previous males, to avoid sperm displacement by other males, and to induce females to use his sperm during fertilization. Since resources are limited, adaptations to perform any of these functions may conflict with each other (and with other life-history traits) and trade-offs are expected to evolve. We studied if males of the polygamous true bug Stenomacra marginella face a trade-off between multiple mating and survivorship, by comparing the survivorship of virgin and multiply mated males. We also looked for physiological costs of ejaculate production by examining ejaculate production in consecutive matings in multiple mated males. Multiply mated males were able to produce ejaculates of similar size in up to six consecutive copulations but they had decreased survivorship in comparison with virgin males. There was no difference in survivorship between males mated three and six consecutive times, suggesting that the negative relation between survivorship and number of copulations is not linear. The decrease in survivorship seems to be a cost of mating and ejaculate production. This cost could favor the evolution of prudence in the allocation of resources to ejaculate production (e.g., cryptic male choice).     

Sex- and age-specific survival of the highly dimorphic Alpine ibex: evidence for a conservative life-history tactic

1. Age-specific survival of 215 males and 117 females of the highly sexually dimorphic Alpine ibex Capra ibex (L.) was assessed from a 21-year capture-mark-recapture (CMR) programme (1983-2004). The study covered two contrasted periods of population performance (high performance from 1983 to 1997 vs. low performance from 1998 onwards). 2. Based on current life-history theories for sexually dimorphic species, we expected that survival should decrease with age in both sexes, female survival should be buffered against environmental variations, male survival should decrease during the low performance period, and adult survival should be lower in males than females during the low performance period. 3. Survival of both sexes was strongly affected by age, with the four age classes (yearling, prime-aged adults of 2-8 years of age, old adults of 8-13 years of age, and senescent adults from 13 years of age onwards) generally reported for large herbivores. 4. Survival of females at all ages, and of yearling and prime-aged males, was buffered against environmental variations and was the same during periods of high and low population performance. The survival of old males decreased in years of low population performance. 5. All marked yearlings (32 females, 56 males) survived to age 2. Survival of prime-aged females (0.996 +/- 0.011) was higher than for other large herbivores, but similarly to other large herbivore species, it declined slowly and regularly with increasing age afterwards. Male survival was 5-15% higher each year than that of males of other large herbivores. Males enjoyed very high survival when prime-aged (0.981 +/- 0.009) and as old adults (high-performance period: 0.965 +/- 0.028, low-performance period: 0.847 +/- 0.032). 6. The very high survival of males, coupled with their prolonged mass gain, suggests a highly conservative reproductive tactic. Male ibex differ from similar-sized herbivores by showing a nearly indeterminate growth in horn size and body mass. By surviving to an advanced age, males may enjoy high reproductive success because of their large size.     

Chromosome banding homologies in three species of Aedes (Stegomyia)

The chromosome complements of the mosquitoes Aedes aegypti, Aedes mascarensis, and Aedes albopictus, belonging to the subgenus Stegomyia, gave a uniform response to the Q-, H-, and R-banding techniques. Of the three homomorphic chromosome pairs, only the shortest or sex pair (I) showed a consistent banding pattern. In the three species, a bright yellow intercalary band was present on one arm of both chromosomes of the sex pair after heat treatment and staining with acridine orange. The rest of the chromosome and the other two pairs fluoresced orange-red. The same intercalary region appeared completely dark with the fluorochromes quinacrine and Hoechst 33258, while the rest of the chromosomes fluoresced dull. The same banding pattern was present in males and females. Size variations of the Q- and H-negative and R-positive intercalary bands were observed within each species. The results are interpreted in terms of structural homology of the sex-determining chromosomes, which is retained within the subgenus.     

The reproductive cycle of yellowfin bream, Acanthopagms australis (Günther), with particular reference to protandrous sex inversion

Yellowfin bream, Acanthopagrus australis , of all age classes were collected from Moreton Bay, Australia. The species possessed typical sparid ovotestes in which the testis and ovary occur in separate zones. During the spawning period (June-August) juveniles, functional males and functional females could be distinguished by the macroscopic appearance of the gonad. The sex ratio of males to females decreases with age, indicating protandrous sex inversion.
Histological and structural study of the ovotestis showed all fish have previtellogenic cells in the ovarian zone but only juvenile and male fish have developing spermatogenic cells in the testis. Most juveniles become functional males by the age of two years but a small proportion of juveniles develop directly into functional females (primary females). Protandrous sex inversion commences after the spawning period when male fish appear with spermatozoa and no other spermatogenic cells in the testis. During the period November-January male fish with no spermatogenic cells are common and a reduction in size of the testis occurs so that by March-April the ovotestis becomes structurally and histologically similar to the female ovotestis. Some fish remain functional males during their whole life-history (primary males). In functional females vitellogenic cells are present in the ovary only during the spawning period and the testis remains very small in size.