The results presented here show the duration of reductions in cattle TB incidence associated with widespread badger culling. Beneficial effects inside culled areas were greatest shortly after culling ended, but then declined over time and were no longer detectable four years after the last annual cull (i.e. three years into the post-trial period). On adjoining lands, the effects of culling were estimated to be beneficial only for the first 18 months of the post-trial period but never significantly so.
Although there have been a number of assessments of the effects of badger culling on cattle TB, our study provides the only experimentally-derived estimate of the duration of effects following the cessation of culling. There has been one other large-scale replicated trial of the effects of badger culling on cattle TB incidence, albeit without the randomised allocation of treatments, or the no-culling control 
. This study, conducted in the Republic of Ireland and known as the Four Areas Trial, found reductions in cattle TB incidence ranging from 51% to 68% over a five-year culling period 
. One explanation for the larger beneficial effect of ongoing culling observed in the Four Areas Trial is that greater reductions in badger density may have been achieved, because (i) land occupier compliance was higher; (ii) the use of snares, rather than cage traps, probably allowed a higher proportion of badgers to be captured; and (iii) the culling areas were selected to have geographical barriers such as coastline and rivers which would impede badger recolonisation. However, since culling is still ongoing in the Four Areas, that study provides no data on the duration of impacts post-culling which can be compared with the results presented here. Similarly, Kelly et al
studied the long-term effects of badger culling on cattle TB using 16 years of observational data, but badger culling was ongoing throughout (with some periods having more intensive culling than others).
In the absence of data on badger populations during the post-trial period, we cannot be certain of the ecological and epidemiological mechanisms underlying the changes in cattle TB risks that we documented in and around former RBCT culling areas. However, we suspect that these changes reflect recovery of badger numbers and spatial organization following the cessation of culling. Proactive culling markedly reduced local densities of badgers 
, which would be expected to reduce the overall risk of cattle coming into contact with badgers. However, culling also prompted expansions of badger ranging behaviour 
, increasing the number of herds that each badger could potentially contact. Moreover, culling increased the prevalence of M. bovis
infection among badgers 
; this, combined with badgers' expanded ranging, would increase the probability of badger-to-cattle transmission, undermining the beneficial effects of reduced badger density. In another study, cessation of culling prompted a contraction of badger ranging within about two years, but recovery of badger numbers took around 10 years 
. We previously suggested 
that the marked reductions in cattle TB incidence observed immediately after the cessation of culling might reflect contraction of badger home ranges (and consequently reduced contact with cattle) prior to substantial recovery of badger numbers. We further speculate that the subsequent decline and disappearance of these beneficial effects may reflect increasing badger numbers, and consequently increased badger-cattle contact. While it is impossible to determine whether the system has now returned to equilibrium, in other studies badger numbers have taken five 
to ten 
years to recover from culls, suggesting that growth of the badger populations in RBCT proactive areas may continue for several more years. As the prevalence of M. bovis
infection in badgers was found to rise on successive culls 
, it is possible that the prevalence in badgers might still be elevated in RBCT areas (although no data are available to test this hypothesis). Were this the case, however, continued growth of the badger populations might be associated with future increases in the risk of TB transmission to cattle herds in areas proactively culled during the RBCT. Continued surveillance of cattle herds will allow characterisation of any further changes in cattle TB incidence, while studies of badger population density, spatial organization, and M. bovis
infection prevalence could provide ecological and epidemiological insights into the long-term impacts of culling, and its cessation, on bovine TB dynamics.
It is important to note that the effects described here relate only to culling as conducted in the RBCT, i.e.
deployment of cage traps by highly trained staff in coordinated, large-scale, simultaneous operations, repeated annually for five years and then halted. As described elsewhere, culling-induced changes in badger numbers and movement patterns mean that culling which is small-scale, patchy, short-term or asynchronous is very unlikely to provide comparable reductions in the incidence of cattle TB and could well prompt increases 
. Other culling methods, such as snares or gassing, might be expected to remove a higher proportion of local badger populations than did cage traps (albeit with a likely cost in terms of badger welfare). However, since there is both ecological 
and genetic 
evidence that badger culling prompts substantial immigration from surrounding lands, improvements in culling efficiency might not result in proportional reductions in badger density, and would not therefore be expected to greatly improve the beneficial effects of culling. In principle, such immigration could be limited by culling within geographical features which present barriers to badger movement (as in the areas selected for culling in Ireland's Four Areas Trial 
). However, such geographical barriers are sparse in TB-affected areas of Britain [e.g., 28]
. Detailed consideration of other potential forms of badger culling 
suggests that no practicable methods would be likely to yield benefits markedly greater than those achieved in the RBCT.
Our results suggest that culling would need to be targeted at circular areas larger than 141 km2
for long-term benefits to be realised. Because the relative benefits improve only slowly with increasing area culled (), even larger areas would need to be targeted to be confident of benefits substantially greater than break-even. For example, to be confident of achieving at least a 10% reduction in the overall incidence of cattle TB would require targeting culling at circular areas ≥568 km2
. These extrapolated figures are somewhat larger than those published most recently, because earlier extrapolations assumed that the benefits of culling increased at greater distances inside the culling area boundary 
. Since no such trend is detectable in this updated dataset, it was excluded from the calculations presented here. All such extrapolations are illustrative: in reality, deviations from perfectly circular culling areas would increase edge effects and reduce overall benefits, while positioning of culling areas close to cattle-free areas or geographic barriers to badger movement might potentially reduce edge effects and increase net benefits 
. Nevertheless, such extrapolations give a rough indication of the minimum areas within which culling would need to be conducted for benefits to be realised.
These updated findings also allow an assessment of the financial costs and benefits of badger culling as a tool to control cattle TB. The overall number of breakdowns estimated to be preventable by proactive culling is fairly modest in comparison with background TB incidence (e.g. 22.6 breakdowns prevented over 7.5 years in an area that would otherwise experience roughly 187 breakdowns), and the consequent financial savings much too low to offset the costs of culling using cage traps, snares, or gassing. Defra estimated that the costs of culling would be substantially lower if implemented by licencing of farmers (roughly £1,000/km2
, hence £562,500 for the idealised five-year 150 km2
area described above; note that the Welsh Assembly Government recently published updated cost estimates of £4,200/km2
/year for government-delivered cage trapping and £1,500/km2
/year for farmer-delivered culling 
.). However, this assumed that farmers would conduct the culling themselves (and so included only minimal capital costs) and excluded the costs of training farmers or coordinating their efforts 
. In the absence of such training and coordination, licenced culling would almost certainly be patchy, asynchronous, unsustained and uncoordinated, circumstances highly likely to prompt increases, rather than reductions, in the incidence of cattle TB 
. Hence, although the total cost of licenced culling is slightly lower than the potential benefits projected from RBCT results (using 2005 cost estimates 
), it is extremely unlikely that such benefits could in fact be realised by this culling method. The costs of conducting badger culls thus substantially exceed the long-term financial benefits likely to be achieved.
Our findings are broadly consistent with those of a recent analysis 
which assessed the potential financial outcomes of badger culling by combining a transmission model (incorporating aspects of badger ecology such as post-cull disruption of badger social organization, as well as farm management such as cattle movement) with data on costs and benefits. In this model, cage-trapping of badgers (assumed to remove 70% of badgers), produced a net economic loss in all simulations, with these losses being greater than those associated with the other culling options considered (shooting free-ranging badgers, snaring and gassing). The authors concluded “Model results strongly indicate that although, if perturbation
[of badger social groups] were restricted, extensive badger culling could reduce rates in cattle, overall an economic loss would be more likely than a benefit
Predicting the financial implications of continuing (rather than halting) annual proactive culls is speculative. However, we can estimate the financial costs and benefits to be incurred annually in and around the idealised circular area of 150 km2
(with a herd density of 1.25/km2
and a background incidence of 0.08 breakdowns/herd/year) based on the impacts of culling estimated between the fourth proactive cull and the end of the during-trial period (the latest estimates available while the proactive culling treatment was ongoing, and ). On this basis, each year of annual proactive culling in the circular area would be expected to prevent 31.5% of 15 breakdowns inside the culled area (4.7 breakdowns prevented), while increasing the number of breakdowns on adjoining land by 14.7% (prompting 1.4 additional breakdowns), giving an overall total of 3.3 breakdowns prevented on average. This constitutes an annual saving of £89,100 at £27,000/breakdown 
. For comparison, the cost of conducting an annual culls over a 150 km2
area, 75% of which was accessible for culling, is estimated as £427,500 for cage trapping (as undertaken in the RBCT) at £3,800/km2
/year, or approximately £270,000 for snaring or gassing at roughly £2,400/km2
. Clearly, continuing to cull would be relatively costly were the benefits of ongoing annual culling to continue at the levels observed following the fourth and subsequent proactive culls in the RBCT.
Our findings have important implications for the development of cattle TB control policies throughout the British Isles. They show that, although widespread badger culling can achieve overall reductions in the incidence of cattle TB, these benefits are not sustained in the long term once culling is halted. Moreover, the financial costs of conducting the culling substantially exceed the overall benefits accrued. In the absence of other practicable culling methods likely to yield greater benefits, our findings indicate that, on the basis of cost-effectiveness, badger culling is unlikely to contribute to the control of cattle TB in Britain.