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.     

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.     

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.     

The activity patterns of Ommatoiulus moreletii (Diplopoda: Iulidae) in South Australia

Pitfall traps were set and visual observations made along regularly walked transects to assess the patterns of surface activity of Ommatoiulus moreletii in open grassland and dry sclerophyllous woodland in South Australia. Active O. moreletii were most commonly seen and trapped in autumn, to a lesser extent in spring, occasionally in summer and rarely in winter. Day to day variations in activity were related to changes in temperature and moisture. The activity of older stadia was greater than that of younger stadia.     

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.     

The water and temperature relationships of Ommatoiulus moreletii (Diplopoda: Iulidae)

During summer, Ommatoiulus moreletii aggregates in cool, moist sites. O. moreletii prefers high relative humidities and temperatures of 20 to 25°C throughout the year. These preferences explain the summer aggregations.
O. moreletii develops a strong ability to survive exposure to low humidity and high temperature during summer. This may explain why summer mortality of the millipede has only been demonstrated during an extremely hot and dry summer. The percentage water contents of O. moreletii are higher during spring and summer than during autumn and winter. They are also higher in small animals than in large. The ecological and physiological significances of these facts are discussed.     

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.     

The population dynamics of the millipede, Ommatoiulus moreletii(Diplopoda: Iulidae)

The population dynamics of Ommatoiulus moreletii were studied in an open grassland and a dry sclerophyllous woodland in South Australia. During summer, O. moreletii aggregated in cool, moist sites (e.g. beneath tussocks of Lomandra fibrata (Liliaceae)) or burrowed underground. In autumn, the animals surfaced and dispersed. In a particularly hot and dry summer, mortality was demonstrated in the grassland but not in the woodland which was relatively cooler and moister. The survival of females from an autumn breeding season to the subsequent spring was inversely correlated with their maturity in autumn. The advantage to O. moreletii of survival of females after a poor breeding season is discussed with reference to den Boer's (1968) concept of "spreading of risk".     

Changes in desiccation resistance during development in the millipede Polydesmus angustus

Water loss at increasing temperature in dry air (< 5% r.h.) was measured for the eight stadia of Polydesmus angustus Latzel (Diplopoda, Polydesmida: Polydesmidae). Instantaneous rates of water loss, expressed as percentages of original body water, were calculated at 20, 25 and 30°C. At each temperature, the rate of water loss varied greatly among stadia, decreasing between 15‐ and 18‐fold from stadium I to stadium VII, and then significantly increasing from stadium VII to stadium VIII (adult). In all stadia, the rate of water loss increased with rising temperature, but the temperature effect was much more pronounced in juveniles than in larger stadia. The threshold temperature of heat stupor, which was measured from stadium IV onwards, increased significantly during development, from 35.6°C in stadium IV to 39.5°C in adults. Survival at 20°C and 76% r.h. varied as could be expected from the water loss rates in dry air, survival time increasing from stadium I to stadium VII and then decreasing from stadium VII to the adult stage. The maximum tolerable water loss, estimated for stadia V, VII and adults, did not change significantly among stadia (mean: 47% of the original water content), suggesting that survival times under desiccating conditions depended primarily on the rates of water loss. The relevance of the results in terms of summer survival under natural conditions is discussed, and it is concluded that juvenile mortality could be substantial in the field. This is important for understanding the adaptive value of cohort‐splitting in P. angustus.     

Anamorphosis and life-history of the millipede Nopoiulus kochii (Gervais, 1847), new for Egypt

Abstract

Monthly samples were taken to determine the anamorphosis and the life-history of the millipede Nopoiulus kochii, new to Egypt. Stadial determination was possible from counting the podous and apodous rings, rows of ocelli, and serial additions of defence glands. Sexual dimorphism occurred at stadium IV. Maturity is achieved at stadium VII. Few males, however, attained maturity at stadium VI. Further moultings of adult stadia occur so that additional adults are found from stadia VII to XIV. N. kochii breeds at the age of six months.     

Stage-specific effects of temperature and dietary protein on growth and survival of Manduca sexta caterpillars

The effects of temperature and dietary protein concentration on growth and survival of Manduca sexta L. (Lepidoptera: Sphingidae) caterpillars during different larval stages were examined. Sets of caterpillars were raised from hatching at one of five constant temperatures (18, 22, 26, 30 or 34°C) and on one of two artificial diets (low or high protein concentration). Mass gain, duration (development time) and mean growth rate were measured for each caterpillar for the 1st to 3rd stadia, the 4th stadium, and the 5th stadium. Temperature significantly affected mass gain during each larval stage, resulting in smaller mass gains at higher temperatures at each stage. This effect was strongest at high temperatures during the 5th stadium. Temperature significantly affected durations of each larval stage, but the effect varied among stages: for example, the duration of stadia 1–3 decreased continuously with increasing temperature, whereas the duration of the 5th stadium was shortest at 26–30°C and increased at lower and higher temperatures. The effect of temperature on mean growth rate changed dramatically across larval stages: maximal growth rate occurred at 34°C during the 1st to 3rd stadia, at 30°C during the 4th stadium and at 26°C during the 5th stadium. Higher dietary protein concentration significantly decreased the duration of stadia 1–3 and of the 4th stadium, but had no significant effect on the duration of the 5th stadium. Temperature and dietary protein had little effect on mortality rates during any larval stadium, with one exception: mortality during the 5th stadium increased dramatically at temperatures of 30 and 34°C. These results demonstrate that the effects of temperature and dietary protein concentration on growth, development and survival in M. sexta vary markedly in different larval stadia during development; 5th instar caterpillars are particularly sensitive to higher temperatures.     

Survival and Cause-Specific Mortality of Male Mule Deer Under Different Hunting Regulations in the Bridger Mountains,Montana

Abstract: We selected 2 adjacent populations of mule deer (Odocoileus hemionus hemionus) in the Bridger Mountains, Montana, USA, to measure effects on survival rates and causes of mortality of 2 hunting regulations designed to enhance representation of mature males. We compared male survival between the West Slope and South 16 Mile populations considering both hunting and nonhunting sources of mortality with respect to age (fawn, yearling, and mature), month (Jun-May), and year (1990-1995). Harvest rates of mature males were greater than for yearlings, demonstrating hunter preferences. We found no differences in yearling monthly survival rates between October and November or between areas or years. In contrast, we found survival of mature males differed between October and November and across years and study areas. During these months, survival rates of mature males averaged 0.602 on the West Slope under the 2-point regulation and 0.762 on South 16 Mile under the outfitted hunt. Monthly survival during summer also differed by age class, but not area, with estimates of 0.963 for yearling males, and corresponding mature male survival estimates of 0.991, demonstrating greater yearling summer mortality. Winter survival rates of yearlings and matures were similar for both areas with a monthly estimate of 0.986. We found differences in spring monthly survival estimates for the 2 areas, mainly for matures. Yearling male monthly survival estimates were 0.959 and 0.958 for the 2 areas, whereas corresponding mature male estimates during spring were 0.991 and 0.936 on the West Slope and South 16 Mile, respectively. Fawn survival rates varied from 0.101 to 0.770 among years and between areas overwinter. Cumulative effects of nonhunting mortality among all age classes reduced the effectiveness of 2 hunting regulations designed to enhance survival of males to age classes ≥4 years associated with maximum antler development despite accomplishing reductions in harvest rates. Low and variable fawn survival and relatively high nonhunting-related losses of yearling and mature males might be typical of many populations in the Northern Rocky Mountains. Deer managers should avoid populations coexisting with a diversity of large predators in environments with strong year effects when considering opportunities for implementing harvest regulations to improve representation of mature males.     

Maturation in chinook salmon, Oncorhynchus tshawytscha (Walbaum): early identification based on the development of a bimodal weight-frequency distribution

Male chinook salmon, orhynchus tshawytscha (Walbaum), are known to mature sexually after one summer in sea water, one year earlier than any of the females. Two year classes of first generation domestic chinook salmon stock reared to sexual maturity on a commercial salmon farm in British Columbia, Canada were sampled for wet weight at various times during the spring after their first winter in the sea. Frequency distributions of wet weight were developed for both year classes. The weight-frequency distributions for both cohorts became statistically bimodal in May of their first year in the sea. The fish from the two modal groups for both cohorts were sorted and kept separate over the summer. The difference in the mean weight for the two groups increased throughout the summer in both years. The higher weight group in each cohort proved to be >95% early maturing males, and represented 64–46% of all the early maturing males evident in the September following the sort. The potential use of this information for commercial salmon growers and fisheries scientists interested in sexual maturation of the chinook salmon is discussed.     

Sensitive periods for wing development and precocious metamorphosis after precocene treatment of the brown planthopper,Nilaparvata lugens

The critical periods for juvenile hormone suppression of wing development and metamorphosis were examined in a pure brachypterous line of Nilaparvata lugens following topical application of Precocene II (PII) to various stages of the third and fourth nymphal stadia. When PII, in doses ranging from 10 pg to 100 ng, was applied to 12-h-old third or 6-h-old fourth stadium nymphs, long-wing formation (macroptery) was induced. Macropter induction ranged from 5 to 50% in females and from 30 to 50% in males, the effect being more prominent in males. The sensitive period for macropter induction lasted from the early second stadium through to day one of the fourth (penultimate) stadium. Beyond that period nymphs required sensitivity to PII and the numbers of macropters increased towards the final nymphal molt, again males were more sensitive. On the other hand, periods sensitive to PII for precocious metamorphosis induction appeared from early second stadium to 18 h after molting to the third nymphal stadium. Overlapping application of JH-III with PII at early third stadium could totally inhibit the occurrence of precocious metamorphosis and significantly rescue the macropter induction. From the data described, a scheme is presented for the control of wing development and metamorphosis by juvenile hormone, assuming two types of threshold value determining wing form and metamorphosis.     

Developmental changes in the patterns of feeding in fourth- and fifth-instar Helicoverpa armigera caterpillars

Data are presented for developmental changes in feeding behaviour within and across the fourth and fifth stadium of Helicoverpa armigera (Lepidoptera, Noctuidae) caterpillars fed nutritionally homogeneous semi‐synthetic foods. We recorded the microstructure of feeding over continuous 12‐h periods on consecutive days throughout the two stadia, and in one experiment recorded continuously for 21 h. Larvae in the two stadia showed the same general pattern of macro‐events in feeding, including a similar duration of post‐ecdysis fast, which was usually broken by consumption of the exuviae, and then a sustained period in which discrete meals on the experimental food were taken regularly. There were, however, some distinct differences in the patterns of meal‐taking both between stadia and across different one‐third time segments within stadia. Considering between‐stadium differences, the proportion of time spent feeding differed significantly only in the last segment of the feeding period of the two stadia, with the value for the fourth‐instar larvae being substantially greater than for fifth‐instar larvae. As regards within stadium changes, the proportion of time feeding increased from the first to the second segment of both stadia. However, whereas the proportion of time feeding increased from the second to the final segment of the fourth stadium, it decreased across the same period in the fifth stadium. These patterns of changes in the proportion of time feeding within and between stadia, and their behavioural mechanisms (combination of meal durations and meal frequencies), can be explained only partially with reference to increasing food requirements with development. Three areas are identified where further study might help elucidate the reasons for the observed developmental changes in the microstructure of feeding: allometric constraint, the dynamic links between ingestion and post‐ingestive processing, and ecological factors such as predation.     

Ecology of an Indian grassland millipede Streptogonopus phipsoni (Diplopoda: Polydesmoidea)

Each of stadia II-VII and adults (stadia VIII) of Streptogonopus phipsoni (Pocock) can be distinguished by segment numbers, length, breadth and paired legs. Sexes are separate from stadium IV and sex ratios in immature stages were more or less 50% throughout the year, but high proportions of adult females are noted from June to September. High ratios of immature to adult were related to the breeding periods, from May to October, and in other months adults increased in proportion as a result of heavy mortality of immatures.
Adults were aggregated in distribution from April to October while solitary individuals were common in other months of the year.
Population density of S. phipsoni varied from 0.25 to 98 m^-2 on the soil surface and 1 to 11.75 m^-2 in the lower layer (0–7 cm deep). Biomass ranged from 12.25 to 2253 mg^-2 on the surface and 50.75 to 424.75 mg^-2 in the lower layer (0–7 cm deep).
A positive and significant correlation was found between mean monthly population density and mean monthly rainfall.
Streptogonopus phipsoni appears to complete its life cycle in one or two years     

Life stages of the non-native Ommatoiulus moreleti (Lucas, 1860) (Julida,Julidae) in Australian small grain systems

1. The Portuguese millipede, Ommatoiulus moreleti (Lucas, 1860), is increasingly a pest of grains crops in Australian agricultural ecosystems. With the rapid uptake of minimum-tillage practices, habitat quality has been enhanced for several species of soil dwelling invertebrates, in particular O. moreleti.
2. To understand the population dynamics of O. moreleti in Australian grains crops, populations were sampled at multiple sites for more than 2 years using pitfall traps and cardboard roll traps. Specimens were dissected to investigate reproductive status and developmental stage.
3. Millipede trapping rates varied between sites and across the year, and tended to be lower when soil water content (m³/m³) was high. Both sexes were active year round, however females were relatively more abundant when ground temperatures were higher.
4. Males in a copulatory state and females with mature eggs were collected year round although females lacking mature eggs were more common when ground temperatures were higher and the soil was drier, and female egg load was higher under cooler conditions. Females at a stadium 9 or later carried mature eggs and copulatory males tended to be from stadium 8 or later. Different developmental stages could be recovered at all times of the year.
5. These findings indicate persistent populations of O. moreleti across fields throughout the year, with a key breeding period in autumn but likely opportunistic breeding all year. Control of millipedes will need to focus on increasing the impact of biological agents, decreasing suitable habitat for millipede breeding and intensive control at susceptible crop periods.

     

The distribution and dispersal of the introduced millipede, Ommatoiulus moreletii (Diplopoda: Iulidae), in Australia

The millipede, Ommatoiulus moreletii, is indigenous to South-Western Europe. The species has been introduced to Australia where it is in large numbers and constitutes a severe nuisance.
The distribution of O. moreletii in Australia is increasing. Man appears to be a major factor in the establishment of new outbreaks of the species. Individual outbreaks spread slowly but steadily (an expansion of up to 200 metres in radius per year).
The abundance of O. moreletii in Australia relative to its apparent rarity in Southwestern Europe cannot be explained by differences in general climate.
The variety of habitats colonized by O. moreletii in Australia is already wide and there is no reason to doubt that the species will continue to spread much further.     

1-Phenyl-2-pentanone and methyl salicylate: new defense allomone components and their content shift during ontogenetic development of the millipede <Emphasis Type="Italic">Nedyopus tambanus mangaesinus</Emphasis> (Polydesmida: Paradoxosomatidae)

Twelve components were identified in hexane extracts of the polydesmid millipede Nedyopus tambanus mangaesinus (Attems, 1909) and their contents were examined at all stadia of ontogenetic development including two adult conditions (before and after hibernation). Two compounds, 1-phenyl-2-pentanone and methyl salicylate, were newly identified as components of the millipede together with six well-known polydesmoid compounds (benzaldehyde, benzyl alcohol, benzoic acid, benzoyl cyanide, mandelonitrile, and mandelonitrile benzoate) and four phenols (phenol, p-cresol, 2-methoxyphenol, and 2-methoxy-4-methylphenol). Benzaldehyde and benzoyl cyanide were distributed from nymphs at stadium I, and other components started to become detectable at more advanced stadia. The largest content (35.1%) of methyl salicylate was detected in nymphs at stadium I, together with benzaldehyde and benzoyl cyanide, implying the reinforcement of defensive functions during or after egg emergence. The content (18.8%) of 1-phenyl-2-pentanone reached a maximum in hibernated females among all stadia and conditions. Its female-biased distribution at the mating season (ca. 5.7-fold more than in males) implies its sexually related functions.     

The life-history of the julid millipede Cylindroiulus nitidus in a Derbyshire wood

A population of C. nitidus in a deciduous wood at Milldale, Derbyshire included adult males with 36 to 54 podous segments but the highest number of segments possessed by immature males was 38. The occcurrence of males in successive stadia is contrasted with the alternation of copulatory and intercalary males in some other juliform millipedes.
Samples were taken to determine the course of post-embryonic development and life-history in the field. Stadial determination by counting rows of ocelli was not possible but a separation was made by analysis of dimensions. These dimensions and segmental formulae are compared with those of German examples. No intercalary or other modified post-imaginal male stages were present.
From a comparison of the distribution of stadia in five samples it appears that males mature in three and females in four years. Males survive a further four to five, females a further three years after first achieving maturity. The significance of the special and general life-history characters of the species are discussed.