Neighbourhood analyses of tree seed predation by introduced rodents in a New Zealand temperate rainforest

House mice Mus musculus and other introduced rodents represent a novel source of predation on tree seeds in New Zealand forests. In the northern temperate forests where these rodents are native, spatial and temporal variation in tree seed production can result in dramatic fluctuations in the distribution and abundance of seed predators, with subsequent feedbacks on the distribution and abundance of seedlings. We use neighbourhood models to examine variation in rodent predation on seeds of 4 tree species of the temperate rainforests of New Zealand as a function of 1) spatial variation in local canopy composition and 2) spatial and temporal variation in mouse activity. We placed seeds throughout mapped stands of mixed forests in alluvial valley bottoms and on elevated marine terraces in the Waitutu Forest, South Island. The risk of predation on seeds of 2 dominant canopy trees – rimu Dacrydium cupressinum and mountain beech Nothofagus solandri var . cliffortioides – peaked in neighbourhoods dominated by those species and by silver beech N. menziesii , particularly in a year of plentiful seed rain from these species. The risk of predation on rimu and beech seed was also related to measures of local mouse activity. These relationships suggest that the highest local abundance of mice was concentrated in rimu and beech neighbourhoods because of the food provided by seed rain from those trees. Predation on seed of miro Prumnopitys ferruginea , which is eaten by rats but not mice, was low in rimu neighbourhoods and where mouse activity was high. These patterns may reflect spatial segregation in the activity of rats versus mice within stands. Our results suggest that the spatial distribution of canopy trees translates into predictable patterns of variation in mouse activity and seed predation. Heterogeneity in rodent activity and seed predation within stands may have important implications for tree population dynamics.     

Variation in body size, sexual dimorphism and age-specific survival in stoats, Mustela erminea (Mammalia: Garnivora), with fluctuating food supplies

Most hypotheses attempting to explain the evolution of pronounced sexual dimorphism in body size in the three species of weasels (Mustela erminea, M. frenata, M. nivalis) assume that sexual dimorphism is a long-term adaptation, associated with the different reproductive strategies of the two sexes. We here examine an auxiliary hypothesis which predicts that the degree of sexual dimorphism may also vary over the short-term, because when food is temporarily abundant, sexual selection should favour a greater growth rate of males than of females. This hypothesis concerns a phenotypic response which could introduce temporarily increased variation into an existing genotypic trait. We document the present size and sexual dimorphism of stoats introduced last century to New Zealand from Britain in relation to between-year variation in food supply in a single habitat (forests of southern beech, Nothofagus sp.). Southern beech trees produce heavy crops of flowers and seed at 3–5 year intervals, which are associated with very variable supplies of important prey of stoats, including several species of seed-eating birds, litter-feeding insects, and feral house mice (Mus musculus). Alternative prey are scarce. Regressions of condylobasal length and head-body length on mouse population indices were significant in both sexes. Mean condylobasal length was larger in both male and female stoats born after a heavy seedfall compared with those born in non-seedfall years. However, the largest males born in years of heavy seedfall were removed by selective mortality before the age of 3 years, so the condylobasal lengths for old (≥ 3.0 yr) males converged on a common mean regardless of food supply in their birth year. Sexual dimorphism did not vary with food supplies (as reflected in seedfall records or mouse population indices) at any age. First-year survivorship, at least from the age of independence, was significantly negatively correlated with density of stoats in the summer of their birth year.     

The Effects of a Natural Increase in Food-Supply on a Wild Population of House Mice

Changes in density and breeding of the house mouse (Mus musculus) in a New Zealand forest dominated by hard beech (Nothofagus truncata) were monitored for 2.5 years. Mice bred during winter and increased dramatically in density only during a beech mast year. Mice readily ate the endosperm and embryo of hard beech seed in die laboratory and chemical analysis showed it to be a very nutritious food source, similar in quality to Fagus beech seed in the northern hemisphere. Thus the mouse, introduced to New Zealand, responds to a Nothofagus mast year in a similar way to other rodent species in the northern hemisphere during a Fagus mast year.     

Variation in mandible shape and body size of house mice Mus musculus in five separate New Zealand forest habitats

This study investigates variation in house mouse Mus musculus body size and mandible shape across New Zealand, using geometric morphometrics and biomechanical advantage analyses. The Mus phylogroups currently known in New Zealand include Mus musculus domesticus, M. m. musculus and M. m. castaneus. We examined samples of house mice inhabiting five different podocarp and beech forest environments across the North and South Islands (Pureora Forest, Zealandia Wildlife Sanctuary, Craigieburn Forest Park, Eglinton Valley and Hollyford Valley). Significant variation in mandible shape and body size was found between all five forest populations. South Island mice had larger bodies and greater mechanical advantage in the temporalis muscle compared with their North Island counterparts. Zealandia Sanctuary mouse mandibles were broader and shorter than South Island mouse mandibles, and had greater masseter muscle advantage. Centroid size and body weight, but not head-body length, varied significantly with two distinct genetic haplotypes. Finally, annual rainfall was the most significant covariate with mandible shape. Higher rainfall locations were generally associated with soft-food related mandible shapes, while lower rainfall correlated with hard-food mandible shapes. This preliminary investigation provides the framework for further research into mandible shape and body size variation in New Zealand house mice.     

Heavy rimu (Dacrydium cupressinum) mast seeding and rat (Rattus spp.) population eruptions on Stewart Island/Rakiura

Abstract

This study aimed to quantify changes in rat abundance and population structure before, during, and after a rimu (Dacrydium cupressinum) mast seed event in lowland forest on Stewart Island, New Zealand. Rats, primarily ship rats (Rattus rattus), were trapped in low numbers throughout the study period (March 2000‐March 2003), except when they erupted to very high abundance in spring 2002, shortly after heavy rimu seed fall. In the immediate post‐peak phase, scavenging of trapped rats increased substantially; rats were seen and trapped in daylight; and weights of adult female rats were low in relation to their size, which suggests that food shortage was the cause of the subsequent steep decline in abundance. Rat eruptions have been observed on Stewart Island after heavy rimu seed fall several times over the past 40 years. Eruptions of rats caused by heavy rimu seed‐fall may have triggered the invasion of nearby islands by rats, and caused the extinction of several native species on Stewart Island.     

The effects of mice on stoats in southern beech forests

Introduced stoats (Mustela erminea) are important invasive predators in southern beech (Nothofagus sp.) forests in New Zealand. In these forests, one of their primary prey species – introduced house mice (Mus musculus), fluctuate dramatically between years, driven by the irregular heavy seed‐fall (masting) of the beech trees. We examined the effects of mice on stoats in this system by comparing the weights, age structure and population densities of stoats caught on two large islands in Fiordland, New Zealand – one that has mice (Resolution Island) and one that does not (Secretary Island). On Resolution Island, the stoat population showed a history of recruitment spikes and troughs linked to beech masting, whereas the Secretary Island population had more constant recruitment, indicating that rodents are probably the primary cause for the ‘boom and bust’ population cycle of stoats in beech forests. Resolutions Island stoats were 10% heavier on average than Secretary Island stoats, supporting the hypothesis that the availability of larger prey (mice verses wētā) leads to larger stoats. Beech masting years on this island were also correlated with a higher weight for stoats born in the year of the masting event. The detailed demographic information on the stoat populations of these two islands supports previously suggested interactions among mice, stoats and beech masting. These interactions may have important consequences for the endemic species that interact with fluctuating populations of mice and stoats.     

Understanding contributions of cohort effects to growth rates of fluctuating populations

1. Understanding contributions of cohort effects to variation in population growth of fluctuating populations is of great interest in evolutionary biology and may be critical in contributing towards wildlife and conservation management. Cohort-specific contributions to population growth can be evaluated using age-specific matrix models and associated elasticity analyses. 2. We developed age-specific matrix models for naturally fluctuating populations of stoats Mustela erminea in New Zealand beech forests. Dynamics and productivity of stoat populations in this environment are related to the 3-5 year masting cycle of beech trees and consequent effects on the abundance of rodents. 3. The finite rate of increase (lambda) of stoat populations in New Zealand beech forests varied substantially, from 1.98 during seedfall years to 0.58 during post-seedfall years. Predicted mean growth rates for stoat populations in continuous 3-, 4- or 5-year cycles are 0.85, 1.00 and 1.13. The variation in population growth was a consequence of high reproductive success of females during seedfall years combined with low survival and fertility of females of the post-seedfall cohort. 4. Variation in population growth was consistently more sensitive to changes in survival rates both when each matrix was evaluated in isolation and when matrices were linked into cycles. Relative contributions to variation in population growth from survival and fertility, especially in 0-1-year-old stoats, also depend on the year of the cycle and the number of transitional years before a new cycle is initiated. 5. Consequently, management strategies aimed at reducing stoat populations that may be best during one phase of the beech seedfall cycle may not be the most efficient during other phases of the cycle. We suggest that management strategies based on elasticities of vital rates need to consider how population growth rates vary so as to meet appropriate economic and conservation targets.     

Age-Specific Prevalence and a Possible Transmission Route for

The prevalence of infestation of the skulls of stoats with the parasitic nematode Skrjabingylus nasicola was previously described in a national survey by King and Moody (1982). Since then, more samples from Craigieburn Forest Park and from the Eglinton Valley, Fiordland, have been collected, and a method of determining the actual ages of adult stoats has been developed. The extended samples are here examined for a relationship between infestation and age, which could not previously be tested. Prevalence generally increases with age, significantly so at Craigieburn. Stoats which had lived through one or more beech (Nothofagus solandri:) mast years at Craigieburn were significantly more likely to be infested, when the effects of age were allowed for. The hypothesis is advanced that the paratenic host for S. nasicola in New Zealand is the feral house mouse, Mus musculus, which is more numerous after a heavy beech seed fall.     

Stoat density, diet and survival compared between alpine grassland and beech forest habitats

In New Zealand, alpine grasslands occur above the treeline of beech forest. Historically stoat control paradigms in New Zealand?s montane natural areas have assumed alpine grassland is a marginal habitat that limits dispersal between beech forest stoat populations. We compared the summer-to-autumn (January?April) density, weight, diet and winter survival of stoats between these two habitatsduring years of low beech seedfall. Stoats were live-trapped, marked and released in alpine grassland and low-altitude beech forest in the Borland Valley, Fiordland National Park, during 2003 and 2004, and were caught and euthanased for necropsy in 2005. Stoat density was estimated using spatially explicit capture?recapture (SECR). The proportion of stoats marked in one year but recaptured in the next was used as a measure of ?observed survival?. Prey remains were identified from scats collected during 2003 and 2004 and stomachs from stoats killed in 2005. Stoat density was similar in both habitats over the two years, about one stoat per square kilometre. Observed survival from 2003?2004 was also similar, but survival from 2004?2005 was higher in alpine grassland than in beech forest. In 2003, male stoats were on average heavier in alpine grassland than in beech forest, although average weights were similar in the other years. Diet differed significantly between the two habitats, with stoats in alpine grasslands eating mainly ground weta (a large invertebrate) (72%) and hares (23%), while stoats in beech forest ate mainly birds (31%) and mice (19%). Collectively these results suggest that alpine grasslands are not a poor quality habitat for stoats. Traditionally it has been thought that stoats cannot survive on invertebrate prey alone. This research demonstrates that stoats relying largely on invertebrate prey can occur at similar densities and with equivalent survival to stoats relying on vertebrate prey.     

Plant-Species Preferences of Birds in Lowland Rimu (

Plant species preferences of birds were determined by comparing the proportional bird use of plant species during direct observations with the proportions of plant species present on point-height intercepts in lowland rimu (Dacrydium cupressinum) forest in North Okarito, Westland. Plant species and bird use of plant species were divided into 5 m height classes, and rimu trees were divided into four age classes (sapling, pole, mature, and old). The frugivorous New Zealand pigeon (Hemiphaga novaeseelandiae) used mature and old rimu more than expected from the proportion of these age classes present, and it preferred the upper tiers of the forest. The omnivorous tui (Prosthemadera novaeseelandiae) had similar preferences to the pigeon. Other omnivorous species, viz., the bellbird (Anthornis melanura) and silvereye (Zosterops lateralis), shared preferences with both the pigeon and insectivorous species. Most insectivorous species, viz., the brown creeper (Mohoua novaeseelandiae), grey warbler (Gerygone igata), New Zealand fantail (Rhipidura fuliginosa), New Zealand tomtit (Petroica macrocephala), and rifleman (Acanthisitta chloris), used pole rimu more than expected and preferred the middle and lowest tiers. The insectivorous New Zealand robin (Petroica australis) had no preference for any plant species but had a strong preference for deadwood and the lowest tier of the forest. Of the 10 species sufficiently abundant to be monitored, the pigeon, bellbird, and tui are most likely to be detrimentally affected by selective-logging of mature and old rimu.     

Ecology of the stoat in

We studied the ecology of a high-density population of stoats in Fiordland, New Zealand, in the summer and autumn of 1990-91 following a Nothofagus seeding in 1990. Results are compared with findings from the same area in 1991-92, a period of lower stoat density. In the high-density year, minimum home ranges (revealed by radio-tracking) of four females averaged 69 ha and those of three males 93 ha; range lengths averaged 1.3 km and 2.5 km respectively. Neither difference was statistically significant. For combined sexes, average range area in the high-density year was significantly less, and range length was significantly shorter, than in the following year. When we compared stoat diet in the high-density year with that in the following two years, there were no significant differences in the frequencies of occurrence of birds or invertebrates in stoat guts. Overall, bird remains were found in 56% of guts, and invertebrates in 28%. Possum remains occurred in 6% of male stoats but were never found in females. Mice were only detected in stoats in the high-density year, when they occurred in 54% of guts. Lagomorphs occurred significantly more often in the guts of stoats during lower-density years (26%) than the high- density year (7%). Seedfall in Nothofagus forest is synchronous and periodic. Following seedfall, mouse density rises dramatically, followed by a sharp rise in stoat numbers. It has been suggested that mice feed on the abundant seed and that stoats in turn increase because of the large numbers of mice available to them. We suggest that the situation is more complex and that increases in not only mouse, but also bird (and possibly invertebrate), densities may contribute to the high productivity of stoats in the year following a Nothofagus seedfall.     

Breeding variation in female kakapo (

We investigated why some mature females of New?Zealand?s critically endangered parrot, the kakapo (Strigops habroptilus), did not attempt to breed during the 2005 breeding season on Codfish Island. At a population level, the initiation of kakapo breeding appears to correspond with years of mast fruiting of rimu (Dacrydium cupressinum) trees, with the proportion of females that breed each season dependent on the quantity of rimu fruit available. This research investigates possible links between habitat quality within individual home ranges and the breeding status of adult females during 2005, when the abundance of available rimu fruit was low. We assessed the importance of both home range size and habitat characteristics in determining breeding attempts. Foraging home ranges were characterised using radio-tracking and triangulation techniques. The relative importance of habitat variables in optimal breeding habitat was assessed using ecological niche factor analysis. Our results show that female kakapo breeding in 2005 had, on average, home ranges twice the size of those females that did not breed that season and the ranges contained a significantly greater quantity of mature rimu forest. Multivariate analysis illustrates female kakapo were effectively partitioning available habitat, as breeders? foraging locations were positively correlated with high-abundance rimu forest with a tall canopy, described as optimal breeding habitat. In contrast non-breeders? locations were weakly correlated with short forest containing little or no mature rimu forest. To maximise reproductive output each breeding season, conservation managers need to ensure that all breeding-aged females occupy optimal breeding habitat on Codfish Island. Removal to other islands of kakapo not required in the breeding population may enable females to increase their home range size and occupy better breeding habitat.     

Predicting the incidence of mohua predation from the seedfall,mouse, and predator fluctuations in beech forests

Abstract

Predator control will be required to save many mohua (Mohoua ochrocephala) populations from extinction. However, control may be required only in years when stoat (Mustela erminea) densities are high. To manage local stoat populations effectively, a reliable predictor of high risk years is required. We examined whether different levels of beech seedfall and mouse capture rates were related to the levels of mohua predation recorded in the Hawdon Valley, Arthur's Pass National Park, and the Eglinton Valley, Fiordland National Park, between 1989 and 1994. During this period there was only one full beech mast year in each study area during autumn. The full mast seedfall in Hawdon Valley was predominantly of mountain beech (Nothofagus solandri var. cliffortioides) and red beech (N. fused), and in Eglinton Valley it was predominantly silver beech (TV. menziesii). During the following summer, mouse and stoat densities, and the predation rate of adult mohua, all increased considerably. There was very little predation on adult mohua in the summers following poor seedfalls when mouse and predator densities remained low. In 1993, a partial mast did not trigger a mouse or stoat irruption.

We conclude that counts of beech seedfall and indices of mouse density are potential predictors of an impending irruption of key predators. Winter mouse density appeared to be the most reliable indicator, because neither stoats nor mice respond to seedfall alone. A combination of these indicators could be used as a basis for management decisions on whether to undertake stoat control to protect mohua populations in the future. However, more information is required on the seedfall thresholds that may trigger sufficient increases in mouse and stoat numbers and, consequently, bird predation.     

Diet of ship rats following a mast event in beech (

Diet of 98 ship rats (Rattus rattus) was investigated by examination of stomach contents. Rats were trapped (as bycatch in stoat traps) from June to December 2006, in the Dart Valley near Lake Wakatipu, New Zealand, after a Nothofagus beech mast. Plant material was found in 43% of all stomachs, invertebrate fragments in 67%, feathers in 8%, hairs in 46% and skin with hairs attached in 12%, and animal (vertebrate and invertebrate) tissue in 76%. Hair samples examined by scanning electron microscopy were from mice, and rat and/or mouse DNA was found in all (10) samples subjected to DNA extraction. Mice may be an important food resource for rats following beech masts, when mouse population densities are high. Presence of feathers indicates predation of roosting or nesting birds, or scavenging. Consumption of beech seed beyond the time of germination may be due to caching of seed.     

Responses of house mice to the removal of mammalian predators and competitors

Efforts to eradicate multiple mammal pests from offshore islands and fenced mainland ‘habitat islands’ often fail to remove mice, and such failures can result in a dramatic change in the food‐web whereby the removal of larger mammal pests facilitates a population explosion of mice through predator and competitor release. We investigated the ecological responses of house mice to the removal of mammalian predators from a 500‐ha fenced sanctuary at Tawharanui, northern New Zealand. Data on population structure and body condition of mice trapped in 2007, in four habitat types within the sanctuary, were compared with baseline data collected in 2001, before mammal control operations commenced. We hypothesized that: (i) in the absence of mammalian predators mouse densities would increase in all habitat types that provide vegetation cover, and (ii) in the absence of mammalian competitors mice would become heavier due to greater access to food resources. Mouse densities were significantly higher in 2007 than in 2001 in three habitat types. The high density of mice in forest – where none were trapped prior to control – suggests a competitive release, in which mice profited from the removal of ship rats. No mice were caught in the presence of ship rats on a forest trap‐line at a control site outside the sanctuary. Mice trapped in 2007 were significantly heavier than those trapped in 2001, and significantly heavier than mice trapped at the control site. Greater access to food in the absence of competing and predatory mammals probably explains the heavier body weight of Tawharanui mice. There has been a significant change in the mammalian food‐web at Tawharanui, such that the house mouse is now the primary pest. A rapid and dramatic increase in mouse numbers is likely to adversely impact invertebrates and seedling recruitment, which in turn could affect ecosystem functions.     

Preference and performance of a gall-inducing sawfly: plant vigor, sex, gall traits and phenology

Four hypotheses regarding the role of predation in the population dynamics of eruptive small mammal communities were tested using the small mammal assemblage found in mixed forests in New Zealand. Large-scale (750 ha) predator removal was conducted, targeting stoats ( Mustela erminea ). House mouse ( Mus musculus ) and ship rat ( Rattus rattus ) population dynamics during an eruption were compared in areas with and without predator reduction. The success of predator reduction was measured by comparing live-capture rates of predators on treatment and non-treatment areas, and by recruitment rates of the threatened northern brown kiwi ( Apteryx australis mantelli ). Overall, predator reduction was successful, although there was a continual low rate of reinvasion. The predictions and results were that 1) Predators can slow but not prevent a population eruption. Supported: Populations of mice and rats erupted to high densities in areas with and without predator reduction, following synchronous southern beech ( Nothofagus spp.) seeding. 2) Predators cannot truncate peak prey population size. Supported: Peak densities of mice and rats were not significantly different between treatment and non-treatment areas. 3) Predators can hasten the rate of decline in prey populations during the crash phase. Not supported: There was evidence of populations of mice and rats declining slower in areas with predators removed, but none of the trends were significant. 4) Predators can limit low-phase prey populations. Equivocal: Populations of rats in beech forest, and population of mice and rats in coastline habitats were significantly higher in areas with predators removed, but were not significantly different in tawa-podocarp forest. Therefore, the role of food in driving the early stages of the mouse and rat eruption was demonstrated, but the role of predation in the decline and low phases is unclear.     

Bird Abundance in Different-Aged Stands of Rimu (

The abundance of birds in three different-aged stands (young, mature, and old) was examined at North Okarito, a lowland rimu (Dacrydium cupressinum) forest in Westland, using 5-minute counts, transect counts, and mist-netting. Most of New Zealand's common forest bird species were present in the study area, with relatively high numbers of brown creeper (Mohoua novaeseelandiae) and New Zealand robin (Petroica australis), and low numbers of kaka (Nestor meridionalis) and yellow- crowned parakeet (Cyanoramphus auriceps). Most insectivorous species were more abundant than expected (from sampling effort) in young and mature stands, the frugivorous New Zealand pigeon (Hemiphaga novaeseelandiae) was more abundant than expected in mature and old stands, and most omnivorous species, viz., bellbird (Anthornis melanura), silvereye (Zosterops lateralis), and tui (Prosthemadera novaeseelandiae), were more abundant than expected in young and old stands. North Okarito Forest provided an important source of seasonal foods (nectar, fruit, and seeds) for frugivorous, omnivorous, and introduced granivorous species, which tended to have greater changes in their seasonal abundance than did insectivorous species. Coupe- logging of old stands will affect all bird species because it will reduce the overall area of standing forest, but it will have a greater impact on the pigeon, bellbird, silvereye, robin, and tui because of their preference for old stands.     

Trappability and densities of stoats (

Stoat (Mustela erminea) density was estimated by live-trapping in a South Island Nothofagus forest, New Zealand, at 8-9 (Jan/Feb 1996) and 15-16 (Aug/Sep 1996) month intervals after significant beech seedfall in autumn 1995. Absolute densities were 4.2 stoats per km² (2.9-7.7 stoats per km², 95% confidence intervals) in Jan;Feb 1996 and 2.5 stoats per km² (2.1-3.5 stoats per km²) in Aug/Sep 1996. Trappability of stoats increased in the latter sampling period, probably because mice (Mus musculus) had become extremely scarce. accordingly, trapping rates of stoats may vary temporally and spatially with food supply rather than only with absolute abundance. Ship rats (Rattus rattus) capture rates doubled between Jan/Feb 1996 and Aug/Sep 1996, but rapidly declined shortly afterwards. Trappability of ship rats also increased in the latter sampling period. These factors must be considered when planning methods of indexing relative densities of stoats and rats.     

Selection of alpine grasslands over beech forest by stoats (

Predation by introduced stoats is now considered a major threat to the population viability of several New Zealand endemic bird species. Historically stoat research and management has focused on beech forests and little is known about the ecology of stoats in the alpine grasslands occurring above the natural altitudinal limit of beech forest. Several stoat control operations in beech forest valley floors in southern New Zealand assume that adjacent montane areas act as a barrier to stoat immigration. Stoats were live-trapped and radio-tracked in alpine grasslands above the Borland Burn, Fiordland National Park, during the summer and autumn of 2003 and 2004. Seventeen stoats were radio-collared and home ranges were estimated for 11 of them. These home ranges were used in a compositional analysis which showed that these stoats spent significantly more time in alpine grassland than in adjacent beech forest. Range cores calculated for six of these stoats were located high up in alpine grassland and contained very little beech forest. This means that montane areas that contain alpine grasslands are unlikely to be barriers to stoat immigration; rather they may be a source of dispersing stoats that reinvade control areas. Also, endemic animal species that inhabit alpine grasslands could be at risk from stoat predation.     

Large‐scale pest control in New Zealand beech forests

In 2014, baits laced with the poison sodium fluoroacetate (1080) were sown over 694 000 ha of mostly native beech forests in New Zealand to control rats, stoats and possums – a landscape‐scale pest control programme called ‘Battle for our Birds’. This large pest control operation was necessitated by the mast seeding of beech trees which led to irruptions of rodent and stoats which were predicted to lead to decreases in vulnerable native wildlife. In this article, we describe why and how this extensive pest control programme was developed and implemented. We describe the seedfall monitoring that was used to determine the need for large‐scale rodent and stoat control and the response of these predators to this control. We also provide a summary of the bird monitoring that was undertaken to demonstrate the effectiveness or otherwise of the programme.