Continuous population estimates forDendroctonus frontalis Zimm. (Coleoptera: Scolytidae) occurring in infestations

Infestations of Dendroctonus frontalisZimm. are often observed to enlarge continuously by the colonization of new hosts in a pattern similar to a forest fire. This pattern of infestation growth presents unique problems in quantitatively estimating populations of D. frontalis. Beetle populations on each infested tree in an infestation go through five processes: attack, oviposition, reemergence, survivorship, and emergence. These processes, which have been described mathematically in the literature, each take several days for completion. In order to follow the distribution and abundance of D. frontalis throughout the course of development of a spot, we need a daily estimate of the number of beetles involved in each process on every tree. Since it is not practical to sample each tree daily, we developed a procedure whereby quantitative estimation procedures for within-tree populations were used in combination with the mathematical models for the life processes to produce a daily record of the number of adults successfully attacking trees, the number of eggs oviposited, the number of beetles reemerging, number of beetles surviving within the trees, and the number of beetles emerging. These daily estimates were then summarized for all trees in the spot for the duration of the infestation. The daily record of populations of D. frontalis, used with information on infestation geometry, were suggested to be of value in describing and elucidating several important facets of population dynamics including dispersal patterns within infestations, between tree beetle loss (mortality), and time lags among the various population processes. The information reported can be used to develop simulation models of population dynamics or to validate existing models.     

Evaluation of sampling methods for determining infestation rates of the tracheal mite (Acarapis woodi R.) in colonies of the honey bee (Apis mellifera): Spatial,temporal, and spatio-temporal effects

Current sampling methods for estimating infestation rates of tracheal mites in colonies of the honey bee,Apis mellifera, assume that infested bees are randomly distributed and that temporal fluctuations in infestation rates occur homogeneously throughout the colony. We examined these assumptions. Samples of bees were collected from up to five locations in each of eight colonies, and colonies were sampled several times throughout the year. Estimates of infestation rates varied, depending on the location in the colony from which a sample was obtained. Temporal fluctuations in infestation rates did not always occur homogeneously with respect to sampling location. These results demonstrate that assumptions of current sampling protocols for estimating tracheal-mite infestation rates are often violated. Consequently, estimates derived using these methods may not be accurate, and conclusions based on such estimates may not be valid.     

Impact of foraging byMonochamus titillator [Col.: Cerambycidae] on within-tree populations ofDendroctonus frontalis [Col.: Scolytidae]

The impact of foraging byMonochamus titillator (Fab.) on within-tree populations ofDendroctonus frontalis Zimm. was described. Total population estimates forD. frontalis and inner bark area foraged byM. titillator were computed for 89 loblolly pine,Pinus taeda L., sampled over a 3 year period. Frequency histograms were prepared for the surface area of habitat infested, area foraged byM. titillator, and within-tree populations ofD. frontalis. These data were used to calculate the proportion of area foraged byM. titillator, the proportionalD. frontalis mortality for the entire tree, and the proportional mortality occurring in the foraged area. Histograms showing the variation of these components were prepared and described using nonlinear mathematical models. Ca. 20% of the infested surface area was foraged byM. titillator. Mortality toD. frontalis on a per tree basis was ca. 14%. Mortality in the area foraged byM. titillator was ca. 70%. These estimates were highly variable between individual trees and a procedure for predicting the probability of a given level of foraging and mortality was described. Sources of variation influencing foraging byM. titillator and mortality toD. frontalis were investigated. Variation between years and season followed similar trends with both foraging and mortality increasing from the base to the top of the infested bole. Variation between tree size-class was highly significant. Larger size-class trees had substantially greater foraging and mortality than did the smaller size-classes. Mortality within the foraged area was also found to be greater at the extremes of the infested bole.

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Résumé L’impact de l’activité deMonochamus titillator (Fab.) sur les populations à l’intérieur de l’arbre deDendroctonus frontalis Zimm. est décrit. Le total des évaluations de population deD. frontalis dans des zones à l’intérieur du tronc attaquées parM. titillator a été soumis à l’ordinateur pour 89 pins,Pinus taeda L., échantillonnés pendant 3 ans. Les histogrammes de fréquence ont été établis pour la surface de l’habitat infesté, la surface explorée parM. titillator et pour les populations deD. frontalis présentes dans l’arbre. A partir de ces données on a calculé la part de surface utilisée parM. titillator, la mortalité deD. frontalis extrapolée pour l’arbre entier et la mortalité se produisant dans la surface exploitée. Les histogrammes traduisant les variations de ces composantes ont été dressés en ayant recours à des modèles mathématiques non linéaires. Environ 20% de la surface infestée a été exploitée parM. titillator; la mortalité du scolyte fut d’environ 14% dans l’arbre et de 70% dans la zone attaquée parM. titillator. Ces évaluations sont très variables d’un arbre à l’autre et on a mis au point un procédé pour prévoir la probabilité d’un taux donné d’activité alimentaire du cerambycide et de mortalité du scolyte. Les causes de cette variabilité ont été étudiées. La variation suivant les années et les saisons suit les mêmes tendances à savoir l’augmentation à la fois de l’activité alimentaire et de la mortalité de la base au sommet du pin attaqué. La variation selon les classes de taille des arbres est hautement significative: les arbres les plus gros présentent une exploitation et une mortalité beaucoup plus importantes que les arbres plus petits. La mortalité du scolyte dans la zone attaquée par le cerambycide est également plus élevée aux extrémités du pin infesté.

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Texas Agricultural Experiment Station No.TA15244.     

Incidence of honeydew in southern pine-hardwood forests: implications for adult parasitoids of the Southern Pine Beetle,Dendroctonus frontalis (Coleoptera: Scolytidae)

The incidence of homopteran honeydew was measured under pines and hardwoods located within 10 Dendroctonus frontalis Zimmermann infestations on the Talladega National Forest (Alabama) during summer, 2000. There were more honeydew droplets beneath hardwood trees compared to pines, but no difference beween D. frontalis infested and uninfested pines. The number of honeydew droplets per tree differed between sampling dates, likely due to the effect of mean weekly temperature and rainfall on homopteran populations. The presence of honeydew on hardwoods may explain why D. frontalis parasitoids forage in areas that lack hosts and suggests that the fecundity of D. frontalis parasitoids may not be limited by adult nutrition.     

Partial age-specific life tables for Monochamus titillator in Dendroctonus frontalis infested loblolly pines

Partial age-specific life tables were constructed for Monochamus titillator(Fabricius) (Coleoptera: Cerambycidae) in Dendroctonus frontalisZimmermann (Coleoptera: Scolytidae) infested loblolly pine (Pinus taedaL.) trees. Successive life stages including egg, early larvae, mid-stage larvae, late larvae, and adult emergence were sampled within six sample trees. Generation mortality ranged from 60.94% to 98.61% in sample trees. Highest mortality typically occurred to eggs and mid-stage larvae. Possible mortality factors included resinosis, predaceous beetles, parasitoids, and woodpeckers. Dendroctonus frontalisbrood stages were determined for consecutive M. titillatorsampling. Monochamus titillatorand D. frontaliscoexisted and likely interacted in the phloem of host trees for at least 20 days.     

Mathematical description of within-tree distributions of the various developmental stages ofDendroctonus frontalis Zimm. (Coleoptera: Scolytidae)

Probability density functions for the various within-tree developmental life stages of D. frontalis are derived. The differences between some of these distributions are discussed. The possibility of the existence of a negative feed-back mechanism which enables the beetle to make optimum utilization of the available resources is proposed. In addition, the concept of an optimum location on the infested bole where the environmental (tree) conditions are such that maximum survivorship occurs is advanced.     

Über morphologische folgen der parasitierung von Euscelis-männchen (Homoptera auchenorrhyncha) mit dryiniden-larven

Summary At the end of July 1959 in a field population (from Taucha near Leipzig) all males of the leafhoppers Euscelis plebejus Fall. and E. distin, guendus Kbm. parasitized by Dryinid larvae (Gonatopus sepsoides Westwood) had a significantly larger Aedoeagus than the not infested ones. With E. plebejus the infested males were larger on the whole too. This finding is in contrast with the more or less strong reduction of the outer genitalia normally produced by entomophagous parasites in Homoptera. A thorough discussion of this discrepancy by all available observations on the bionomical and morphological effects of Dryinid infestation leads in connection with ecomorphological statements on the relation between generative and vegetative development in leafhoppers to the conception, that the infestation by Dryinids only then causes a reduction of the host genitalia, if it happens very early (Aphelopinae). If the parasitization occurred in already older host larvae, the transformation of metabolism caused by check of the gonads only effects a delayed development and by this an increase of growth. By this the male host at least, reaching the adult stage at all, shows a general enlargement of size, only faint indeed—due to the relatively short parasitization timend surely to be proved only with eurymorph species as E. plebejus or with organs positively allometrical growing as the Aedoeagus of the Euscelis species. Along with this the size of the latter is demonstrated to be modificable in the second line by parasitic infestation.

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Herrn Professor Dr. Arno Wetzel zum 70. Geburtstage gewidmet.     

Probability functions for components of theDendroctonus frontalis—Host tree population system and their potential use with population models

Detailed investigation of the within-tree population system of Dendroctonus frontalisZimmermann has resulted in a large data base consisting of abundance estimates for various life stages. This data base was used to construct histograms for transformed estimates and several life stage indices. Histograms were also constructed for transformed values of adult residence time, brood development time, and several host-tree characteristics. Probability and cumulative density functions of the Weibull distribution were fitted, in tandem, to the scaled frequencies and interval means for each histogram. The inverse cumulative function is known, and with a uniform random number generator, allows the selection of random deviates from each distribution. This technique can be used for generating initial (starting) values in D. frontalis population models.     

Detectability of the emerald ash borer (Coleoptera: Buprestidae) in asymptomatic urban trees by using branch samples

The emerald ash borer, Agrilus planipennis Fairmaire, is an exotic invasive insect causing extensive mortality to ash trees, Fraxinus spp., in Canada and the United States. Detection of incipient populations of this pest is difficult because of its cryptic life stages and a multiyear time lag between initial attack and the appearance of signs or symptoms of infestation. We sampled branches from open-grown urban ash trees to develop a sample unit suitable for detecting low density A. planipennis infestation before any signs or symptoms are evident. The sample unit that maximized detection rates consisted of one 50-cm-long piece from the base of a branch ≥6 cm diameter in the midcrown. The optimal sample size was two such branches per tree. This sampling method detected ≈75% of asymptomatic trees known to be infested by using more intensive sampling and ≈3 times more trees than sampling one-fourth of the circumference of the trunk at breast height. The method is less conspicuous and esthetically damaging to a tree than the removal of bark from the main stem or the use of trap trees, and could be incorporated into routine sanitation or maintenance of city-owned trees to identify and delineate infested areas. This research indicates that branch sampling greatly reduces false negatives associated with visual surveys and window sampling at breast height. Detection of A. planipennis-infested asymptomatic trees through branch sampling in urban centers would provide landowners and urban foresters with more time to develop and implement management tactics.     

An invasion revisited: the African big-headed ant (<Emphasis Type="Italic">Pheidole megacephala</Emphasis>) in northern Australia

Long-term studies provide the best information for invasion ecology as multi-temporal sampling can illuminate invasion dynamics, such as population changes and rate of spread. In 1996, Hoffmann et al. surveyed an infestation of African big-headed ant, Pheidole megacephala, in semi-natural rainforest of northern Australia. Here, we re-survey this infestation 9 years later to assess the dynamics of this invasion against the key finding of the initial study. Importantly, we re-sample a site sampled prior to invasion to demonstrate the causal link between an infestation and changes in a native invertebrate community. We found the area infested had almost doubled, and P. megacephala biomass in infested sites was up to 18 times greater than that of native ants in uninfested sites. In the two sites with the youngest infestations in 1996, P. megacephala abundance had increased more than 20-fold than that measured in 1996. Native ant abundance and species richness in infested sites were notably lower in 2005, with only one native ant specimen found in the most recently infested site. The abundance of other macro-invertebrates was the lowest in the three oldest infested sites. Coleoptera and Orthoptera were less abundant in infested sites. This study supports the findings of the first ‘snapshot’ study, and has shown that the area infested has almost doubled, populations are increasing, and ecological impacts remain severe. It reinforces that P. megacephala represents a serious ecological threat, and we argue that there is a greater need for management of this ant globally.     

Evaluation of Beauveria bassiana (Ascomycota: Hypocreales) as a control of the coffee berry borer Hypothenemus hampei (Coleoptera: Curculionidae: Scolytinae) emerging from fallen,infested coffee berries on the ground

The effects of Beauveria bassiana strains on coffee berry borers (CBB), which emerge from infested berries left on soil, and its impact on the infestation of coffee berries on tree branches were evaluated at two Experimental Stations (Naranjal-Caldas and Paraguaicito-Quindio) in the Colombian coffee zone. Using a completely randomized design with 10 repetitions, 50 coffee berries artificially infested with CBB were placed on the base of a coffee tree. Four treatments including B. bassiana strain Bb9205, a mixture of Cenicafé strains (Bb9001, Bb9024 and Bb9119), a commercial formulation of B. bassiana and a control (water) were sprayed with 1×10⁹ conidia per tree. After 30 days, all fungal strains lowered the infestation levels of the coffee berries on the trees at both locations. The mixture of Cenicafé strains decreased the tree infestation between 50 and 30% at both locations. In the berries dissected from each treated tree, insect mortality was about 40% at both locations compared to 15% in the control. B. bassiana strains also decreased the insect population inside the newly infested berries on the trees by 55–75%. The mixture of Cenicafé strains was the most effective for decreasing insect populations. B. bassiana significantly decreased CBB populations that emerged from fallen, infested, coffee berries and reduced future insect generations.     

Host infestation patterns of the massive liana Hydrangea serratifolia (Hydrangeaceae) in a Chilean temperate rainforest

As competition from lianas reduces fitness of host trees, lianas could influence community composition and structure if potential host species differ in susceptibility to infestation. We quantified infestation frequencies of Chilean temperate rainforest tree species by the massive liana Hydrangea serratifolia (H. et A.) F. Phil (Hydrangeaceae), which climbs using adhesive adventitious roots, and examined relationships with host light requirements and stem diameter. We recorded presence or absence of H. serratifolia in a random sample of 515 trees ≥10 cm diameter. Fifty‐four per cent of trees were infested by at least one individual of H. serratifolia. Although there was significant interspecific variation in infestation frequency, this variation was not systematically related to light requirements of host tree species. Probability of infestation increased with diameter for most host tree species, and old trees were found to be infested by a wide range of liana size classes, including some stems <2 cm diameter. This evidence supports the proposal that lianas which attach by adhesive roots can colonize host stems of any size.     

Local and systemic responses induced by aphids in Solanum tuberosum plants

The aphids Macrosiphum euphorbiae (Thomas) and Myzus persicae (Sulzer) (Homoptera: Aphididae) are serious pests of potato (Solanum tuberosum L.) (Solanaceae), notably in transmitting several plant viruses. Heterospecific interactions may occur between these two species as they are often seen at the same time on the same potato plant in the field. As aphid infestation is known to induce both local and systemic changes, we conducted experiments to determine the effect of previous infestation on probing behaviour and feeding‐related parameters. We used the DC electrical penetration graph technique to characterize the influence of previous infestation by conspecific M. persicae or by heterospecific Ma. euphorbiae on M. persicae feeding behaviour at both local and systemic levels, i.e., on previously infested leaves and on non‐previously infested leaves of infested plants, respectively. Conspecific and heterospecific infestation led to similar modification of M. persicae feeding activities. However, the effects of previous infestation occurring at the local level were opposite to those observed at the systemic level. Myzus persicae food acceptance was slightly enhanced on previously infested leaves, whereas it was inhibited on non‐infested leaves of infested plants, which indicated an induced resistance mechanism. Our results advance the understanding of the mechanisms involved in aphid–host plant acceptance and colonization processes on potato plants in conspecific and heterospecific situations.     

Spatial spread and infestation risk assessment in the Asian longhorned beetle,Anoplophora glabripennis

The Asian longhorned beetle (ALB), Anoplophora glabripennis (Motschulsky) (Coleoptera: Cerambycidae), is recognised as potentially one of the most damaging invasive insects in Europe and North America. International trade has increased the risk of accidental introduction of ALB. An eradication programme was initiated in Northeast Italy in June 2009, when an ALB infestation was discovered. The infestation was monitored by annual surveys of all host‐tree species growing in the eradication area. Infested trees were cut down and chipped. This study analyses the spatiotemporal distribution of infested trees for a 5‐year period from 2008 to 2012 using a generalised linear model approach. The results show that spread and infestation risk were significantly affected by (1) distance of suitable hosts from the nearest infested trees, (2) number of infested trees in the surroundings, and (3) annual variation. The significant differences in beetle dispersal between years reflect to some extent the onset of the eradication programme. The model allowed the estimation of arbitrary probability‐based management boundaries surrounding ALB‐infested trees. For example, the model estimated a 0.1% probability of attack on a suitable host tree 1 910 m from an existing attack.     

EFFECT OF ECTOPARASITIC MITES ON SEXUAL SELECTION IN A SONORAN DESERT FRUIT FLY

We conducted a field study and a laboratory experiment to test whether ectoparasitic mites, Macrocheles subbadius, generate parasite-mediated sexual selection in the Sonoran desert endemic fruit fly, Drosophila nigrospiracula. Male flies gather on the outer surfaces of necrotic saguaro cacti where they engage in male–male competitive interactions and vigorous female-directed courtship. At these sites, operational sex ratios were significantly skewed toward males. The degree to which mites were aggregated among flies varied across the 25 fly populations sampled. The degree of mite aggregation across fly populations was strongly positively related to the mean number of mites per fly (intensity of infestation). Both the intensity and prevalence of infestation (fraction of flies infested) increased with the age of the cactus necrosis. Infested flies of both sexes were significantly less likely to be found in copula than uninfested flies, and mean intensity of infestation was significantly more pronounced in noncopulating than in copulating flies. The effect of attached mites on copulatory success exhibited dose-dependency, and this effect was more stringent in males: males or females with more than two and four mites, respectively, were never found in copula. The magnitude of parasite-mediated sexual selection was estimated for 12 fly populations by calculating selection differentials for each sex separately. The relation between intensity of infestation and magnitude of parasite-mediated sexual selection was stronger in males but significant for both sexes. We also assayed copulatory success of field-caught males in the laboratory, both during infestation and after experimental removal of mites. Males infested with two mites copulated less frequently than uninfested individuals, and in mating trials after mites had been removed, previously infested males copulated as many times as flies with no history of infestation. These findings, and the lack of difference in the number of mite-induced scars on copulating and single individuals in nature, strongly suggest that the reduced copulatory success of infested flies is attributable to an effect of mites per se, rather than to a character correlated with parasitism or previous parasite infestation.     

Physiological and ecological hosts of Popenaias popeii (Bivalvia: Unionidae): laboratory studies identify more hosts than field studies

1. Freshwater mussels are critically endangered in North America, making it important to understand their environmental requirements at all life stages. As glochidia (larvae), they attach to fish hosts where they undergo substantial mortality, making this transition important in their life cycle. Larval host fish requirements have typically been described using data from laboratory infestations to determine suitable hosts. 2. Laboratory infestations circumvent many natural barriers that prevent infestation of physiologically compatible fishes by mussel larvae. While such methods are invaluable for identifying ‘physiological hosts,’ they cannot fully describe realised ‘ecological hosts’ in the field. 3. We studied Popenaias popeii in the Black River in New Mexico, because it is of conservation concern and it is the only mussel species present, facilitating identification of glochidial infestation. To explore the difference between physiological hosts and ecological hosts, we conducted a 3‐year field study of fishes infested by P. popeii glochidia. 4. Substantially fewer fish species were infested by P. popeii in the wild (10 of 20 observed) than had been identified as physiological hosts in laboratory trials (24 of 31). We combined data on fish abundance, proportion of fish hosts infested (prevalence) and the number of glochidia per fish (intensity) and identified three fish species that probably contributed substantially more to mussel recruitment by carrying more glochidia than other host species. 5. Similarities in behaviour among these fishes allowed us to hypothesise routes of infestation, such as benthos‐feeding by catostomids, that allow glochidia to infest these hosts at higher rates than other suitable hosts. Overall, this approach provides a method of quantifying the relative importance of different species of host fish in the mussel lifecycle.     

Evaluation of tricin,a stylet probing stimulant of brown planthopper,in infested and non‐infested rice plants

The flavone, tricin (5,7,4′‐trihydroxy‐3′,5′‐dimethoxyflavone), is a valuable secondary metabolite that is common in gramineous plants, including cultivated rice (Oryza sativa). It can defend the rice plant against infestation by the brown planthopper (BPH), Nilaparvata lugens Stål, one of the most important pests of rice. This study evaluated the tricin concentration in infested and non‐infested rice plants. The results of the liquid chromatography coupled to tandem mass spectrometry (LC–MS/MS) quantitative analysis showed that the tricin concentration in rice leaves was significantly higher than in the stems and roots. The mass concentration of tricin in the leaves at the leaf stage was significantly higher than at the tiller and booting stages. The relationship between rice variety, BPH resistance and tricin concentration was investigated. There was a significant negative correlation between tricin concentration and the injury severity scores for rice varieties. Moreover, BPH infestation caused variations in tricin concentration among rice plants. High BPH infestation levels can significantly reduce the tricin concentration in rice plants. However, there is no significant effect of the length of infestation times on tricin concentrations in rice leaves. These results suggest that there may be an elicitor in BPH saliva, which is injected into rice plants during BPH infestation and triggers the tricin metabolic system. Future studies need to identify the elicitor and clarify the mechanism underlying tricin reduction in infested rice plants.     

The use of egg counts and suction trap samples to forecast the infestation of spring-sown field beans, Vicia faba, by the black bean aphid, Aphis fabae

Daily suction trap samples at a height of 12·2 m collected throughout the year, winter egg and ‘spring’ population counts on the spindle tree, Euonymus europaeus, and initial infestations of the black bean aphid, Aphis fabae, on field bean, Vicia faba, crops are available from Southern England since 1970. In different areas, estimates of the sizes of the autumn migrations, the over-wintering egg populations, the spring fundatrigeniae and the spring migrations, have been used to forecast field bean crop infestation levels which, in turn, project subsequent trap catches of alatae. The forecasts of crop infestation become progressively more accurate from the autumn migration to the following spring migration, accounting for 28%, 54%, 54% and 64% of the variance respectively. In areas where traps are sited, the spring migration trap samples give the most accurate estimate of the size and timing of crop infestation. Autumn trap catches are particularly useful as very early forecasts of likely very large or very small populations on field beans about 8 months later, but otherwise they lack precision. Egg sampling in winter provides a considerably more accurate forecast approximately 5 months before crop infestations. In spring (May), fundatrigeniae sampled on spindle are most useful for predicting time of migration, and provide approximately 2 weeks' warning for insecticide application, if needed. Finally, trap sampling of the spring migration provides the latest estimate of both the sizes of crop infestations and the timing of insecticide treatment. The E. europaeus and aerial sampling systems are complementary, the traps providing systematic, continuous information and the E. europaeus samples greater detail. Combined, they can provide excellent long-term fore-warning of the need for chemical control and short-term warning of control timing. Forecasts have been 90% correct in eight years out of nine. The error in the ninth year may be due to immigration from the European mainland.     

Dispersion,contagion, and population stability of red scale,Aonidiella aurantii,in citrus orchards with low or zero insecticide use

Red scale, Aonidiella aurantii Maskell (Hemiptera: Diaspididae), has a contagious distribution among fruit and among trees largely due to the fact that most young settle on the same fruit as their mothers. The pattern of distribution and degree of dispersion of red scale within various sets of contiguous citrus trees in several orchards were recorded each autumn for 4 years. Changes in density and site occupancy within trees and within contiguous groups of trees were made to assess the strength of any link between populations and their dynamics on different trees. Aonidiella aurantii was found to have a very clumped distribution whereby trees with a level of scale infestation well above average for their block could be adjacent to trees with few or no detectable scale. However, patches of high density on a given tree usually did not recur for more than 2 years but were replaced with above average infestations on other trees. The stability of the red scale populations varied from block to block. Some blocks had few or no infested trees over the study period whereas others had widely fluctuating levels of infestation. Dispersion coefficients at ‘within tree’ and ‘within block’ levels were similar. Theoretical and statistical models of the relation of average scale levels to the percentage of fruit or trees infested were less precise than they are for other pests because of high variation of clumping intensity of scale within any block.     

The estimation of population size,migration rates and survival in a stratified population

Summary Estimates of survival, migration rates, and population size are developed for a triple catch marking experiment onn (n>-2) areas with migration among all areas and death in all areas occurring, but no recruitment (birth). This repressents the extension to three sampling times of the method ofChapman andJunge (1956) for estimates in a stratified population. The method is further extented to allow for ‘losses on capture’.