Emergence of flies from overwintering populations of cabbage root fly pupae

ABSTRACT.

• 1 Once pupal diapause had been terminated, over-wintering cabbage root fly (Delia radicum (L.)) pupae from Wellesbourne required a further 14 days at 20°C for most of the flies to emerge.
• 2 There were considerable variations in the rates of fly emergence from thirteen populations of cabbage root fly pupae collected between latitudes 50° 42′ and 54° 59′ in England and Wales. These thirteen populations could be grouped into early-, intermediate- and late-emerging types. In the early-emerging type, flies emerged within 14 days at 20°C whereas in the late-emerging type emergence was protracted and was completed only after 100 days at 20°C in one population from Halsall, Lancashire. In the intermediateemerging type, approximately two-thirds of the flies emerged within 14 days at 20° C, the remainder taking considerably longer.
• 3 The intermediate-emerging types could be just mixtures of the early- and late-emerging types.
• 4 Subjecting pupae to diapause-breaking temperatures (4°C) for up to 1 year failed to shorten the time to subsequent fly emergence in any of the populations.
• 5 Populations of early, intermediate- and late-emerging fies could be selected from a parental population, heterogeneous with respect to emergence, within one generation.
• 6 The type of emergence that occurred in a locality was not correlated with latitude.
• 7 Any models developed for forecasting the most appropriate time to apply insecticide in a locality will have to include information about the emergence pattern of the fly population in that locality.

     

An analysis of the deterrent effect of aphids on cabbage root fly (Delia radicum) egg-laying

Abstract. 1. The cabbage root fly, Delia radicum (L.), was deterred from laying eggs on brassica plants with >250 cabbage aphid, Brevicoryne brassicae (L.), or peach-potato aphid, Myzuspersicae (Sulz.).
2. Flies did not lay on plants infested with >250 aphids.
3. Preparations of (E)-β-farnesene, the aphid alarm pheromone, deterred the flies from laying only at the extremely high dose of 32 mg/plant.
4. Although M. persicae secreted large (1 ng/insect) amounts of alarm pheromone and B. brassicae extremely small (<0.01 ng/insect) amounts, both aphids equally deterred D. radicum from laying.
5. The deterrent effect appeared to result from the aphids physically disturbing the flies during host-plant selection.     

Improving the selectivity of water traps for monitoring populations of the cabbage root fly

Traps placed within brassica crops to monitor changes in cabbage root fly activity could be made more selective by painting black the inner wall of the standard fluorescent yellow water-traps. Traps could also be made more selective by covering them with cylinders of Netlon mesh, although this procedure reduces the numbers of cabbage root fly caught by about 70%. Of the single-coloured traps tested, those painted “marigold” yellow were the most selective in capturing female cabbage root flies. Although white and certain blue traps were as effective as the best yellow traps at capturing cabbage root flies, such traps should be avoided, as they catch 4–5 times as many of the closely-related bean seed fly. The presence of bean seed flies makes cabbage root fly identification more difficult and adds considerably to the time required to sort trap catches.     

The relationship between cabbage root fly (Delia radicum) larval feeding and the freeze-dried matter and sugar content of Brassica roots

Five genotypes of swede (Brassica napus var. napobrassica), two genotypes of kale (B. oleracea var. acephala), and two genotypes of rape (B. napus var. napus) were each inoculated at the 8–10 true leaf stage with five cabbage root fly (Delia radicum) eggs. The percentage pupation after larval feeding on individual plant genotypes ranged from 45 to 78%, and the mean pupal weight from 6.5 to 13.0 mg. After 5 weeks, larval feeding damage had reduced root weight by up to 47%, compared with uninoculated plants. The dry matter content of undamaged roots was higher in the kales and rapes than in the swedes. Whilst the dry matter content of the rapes and swedes were not changed by D. radicum damage, that of the kales was elevated. The ethanol-soluble sugar content of the root was reduced in all cases by D. radicum larval damage. However, the effect of D. radicum damage on the concentrations of individual sugars (glucose, fructose and sucrose) was crop- and genotype-dependent. In the roots of kales and rapes, the glucose and fructose concentrations were either very low or unaffected by D. radicum damage, whilst both glucose and fructose were generally reduced in swede roots by D. radicum damage. The root sucrose concentration was either reduced or not significantly affected by D. radicum damage in all of the crop types tested. The percentage pupation and the mean pupal weight of D. radicum were inversely correlated to root freeze-dried matter content. D. radicum pupal weight was positively correlated with root fructose, glucose and ethanol-soluble sugar contents.     

Possible impact of global warming on cabbage root fly (Delia radicum) activity in the UK

A program for simulating the patterns of egg-laying by populations of the cabbage root fly was used to model the effects of global warming on future cabbage root fly attacks. An increase of 3°C in mean daily temperature would cause the cabbage root fly to become active about a month earlier in the year than at present. Under such conditions, the emergence of flies from the overwintering population would be less synchronised, as the completion of diapause and post-diapause development would occur at the same time in different individuals within the population. However, there would continue to be only three generations of fly each year, even in the south of England. With temperature increases of 5°C or 10°C, the fly would complete four generations each year and aestivation would seriously disrupt egg-laying. These rises in temperature would have a major impact on cabbage root fly activity and would require new strategies for controlling this pest.     

Effect of trap background on cabbage root fly landing and capture

Pieces (600 mm × 800 mm) of coloured board, plastic sheeting and woven materials, place beneath water traps prevented the traps from becoming soiled during rainy weather. Such backgrounds are not recommended for use with traps for monitoring populations of the cabbage root fly (Delia radicum L. — Diptera: Anthomyiidae), however, as, instead of increasing trap catch they reduced the numbers of female flies caught by 70%–90%. The main effect was that the visually attractive stimuli from the introduced backgrounds competed with those from the trap. A white background competed with a white trap on a direct fly/unit area basis. Green backgrounds stimulated males to land and the vertical stems of both real and artificial grass induced trivial flights that resulted in greater numbers of males entering traps resting on short grass. Most females were caught over bare soil. To minimize the variation in catch between traps used for monitoring cabbage root fly populations, the background beneath each trap should be similar. For maximum capture, the background should be of grass for male flies and of bare soil for female flies. Care is required if data from traps within mulched crops are used to make pest control decisions, as they will underestimate considerably the numbers of flies in such crops.     

Role of visual and olfactory stimuli in host finding of adult cabbage root flies, Delia radicum

Visual host finding and form recognition by the cabbage root fly, Delia radicum, were studied using four shapes of yellow sticky traps as plant models. Cross, disc, and Y traps caught similar numbers of flies. Cross traps caught more females than vertical or horizontal rectangular traps. Black borders did not increase rectangle trap catches. Discs (8 cm diam.) at ground level caught more females than discs at 40 cm above ground. Larger discs (11.2 and 13.8 cm diam.) caught more females per trap than 8 cm diam. discs, but not as many per unit area.Volatile mustard oils were added to sticky cross traps as single and multiple-component baits. Traps with multiple-component baits did not consistently catch more females than single baited traps. Yellow traps baited with isothiocyanates caught 4–7 times as many females as clear plastic allylisothiocyanate (ANCS)-baited traps.Results indicated yellow discs or crosses at ground level beited with ANCS effectively monitored female D. radicum. Main factors affecting landing of female D. radicum are suggested to be color of substrate, height above ground, presence of host volatiles in vicinity, visual prominence, and area of attractive color. The view that host finding in some insects is mediated by complex responses to multichannel stimuli was supported.

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Résumé L'étude de la perception visuelle et de la reconnaissance des formes par les D. radicum adultes a été réalisée en utilisant des pièges jaunes gluants de quatre formes différentes comme leurres de plantes. Des pièges en croix, en disque et en Y ont permis la capture des mêmes nombres de mouches. Plus de femelles ont été capturées avec des pièges en croix qu'avec des pièges rectangulaires verticaux ou horizontaux. Les captures n'ont pas augmenté avec des pièges bordés de noir. Des disques de 8 cm de diamètre disposés au niveau du sol ont permis la capture de plus de femelles que des disques à 40 cm au-dessus du sol. La capture par piège a été supérieure avec des pièges de 11,2 et 13,8 cm de diamètre qu'avec ceux de 8 cm, mais moindre par unité de surface.Des essences volatiles de moutarde ont été ajoutées aux pièges en croix comme appâts simples ou composés. Les pièges avec appâtes composés n'ont pas été significativement plus efficaces que les pièges à appâts simples. Les pièges jaunes appâts avec de l'isothiocyanata ont entraîné la capture de 4 à 7 fois plus de mouches que des pièges en plastique transparent appâtés avec de l'allylisothiocyanate (ANCS). Les résultats ont montré que des disques ou des croix jaunes appâtés avec de l'ANCS au niveau du sol permettent d'organiser un avertissement agricole contre D. radicum. Les principaux facteurs intervenant dans l'atterrissage des femelles de D. radicum semblent être la couleur du substrat, l'élévation audessus du sol, le contraste visuel, la présence de substance de l'hôte dans le voisinage, et l'aire d'attractivité de la couleur. En conclusion, il est exact de considérer que la perception de l'hôte est provoquée chez quelques insectes par une réponse complexe à des stimuli multicanaux.

     

The effect of colour of the substrate on the landing and oviposition behaviour of the cabbage root fly

Field and laboratory experiments were made in order to understand the relation between the spectral characteristic of a substrate and its attractiveness for Delia radicum (L.) (Diptera: Anthomyiidae) flies to land or oviposit. Landing females preferred substrates with high reflectance of green and particularly of yellow wavelengths, but substrates with a high proportion of the blue and green reflectance simultaneously (light blue and white) were also very attractive. Unattractive substrates had either low reflectance across the whole insect-visible spectrum (dark blue) or increased reflectance only in orange or red region of the spectrum (red) or a large proportion of UV reflectance (aluminium). Landing males were most attracted to the substrates with the highest total reflection. Oviposition attractiveness of a substrate grew with an increase in the proportion of reflectance in the blue and a decrease in the green regions of the spectrum. In addition, the oviposition attractiveness increased with increasing total reflection and contrast with the background, and decreased with a high proportion of UV reflectance.     

Foliar form, colour and surface characteristics influence oviposition behaviour in the cabbage root fly Delia radicum

The effects of foliar form, colour and surface characteristics on the oviposition behaviour of the cabbage fly were investigated by presenting females with various leaf models made of paper. The models differed in shape, size, colour, surface coating, the presence of 3 dimensional folds and a stem. It was shown that in the presence of host leaf surface extracts physical factors can strongly influence oviposition behaviour. Females laid the most eggs around the base of bright green or yellow models having a stem, vertical folds and covered with a thin layer of paraffin. The vertical folds in the surface increased the chance of a behavioural transition from leaf run to stem run.

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Zusammenfassung Mit Hilfe von Papiermodellen von Kohlblättern wurde der Einfluss von visuellen und mechanischen Eigenschaften des Blattes auf das Eiablageverhalten der kleinen Kohlfliege, Delia radicum, untersucht. Die verwendeten Modelle unterschieden sich in Form, Grösse, Farbe, Oberflächen struktur und Stiel des Blattes. Es wurde gezeigt, dass physikalische Reize zusammen mit den chemischen Stoffe auf der Blattoberfläche stimulierend wirken. Ein natürliches Eiablageverhalten und die stärkste Eiablage wurde durch ein Modell ausgelöst, welches sich durch leuchtend grüne Farbe, Stiel und vertikale Falten und einen Paraffinnüberzug auszeichnete.

     

Tarsal chemoreceptors and oviposition behaviour of the cabbage root fly (Delia radicum) sensitive to fractions and new compounds of host-leaf surface extracts

Abstract. Contact chemoreception plays a decisive role in host selection and oviposition behaviour of the cabbage root fly, Delia radicum L. (Diptera, Anthomyiidae). Glucosinolates (mustard oil glucosides) are known to be perceived by the flies, and when sprayed on paper leaf-models induce oviposition. Recently it has become clear that other non-volatile types of compounds must also be involved in host selection.
A pair of ventro-medial C sensilla on die fifth tarsomere respond strongly to a novel compound called tentatively 'cabbage identification factor' (CIF), but not to sucrose, glucose, fructose and proline. CDF is a new non-glucosinolate oviposition stimulant. A single neurone in each sensillum is activated by this compound and the same is true for glucosinolates. In some flies a mixture of bom types of stimuli evoked an apparent mononeural spike train, whereas in odiers spikes of two separate cells were activated. The significance of this variability is not yet clear.
The new stimulant, CIF, does not evoke responses in glucosinolate receptors in the D sensilla. The involvement of the C₃ sensilla in the detection of host-specific compounds constitutes the first known function for C sensilla in D. radicum. CIF appears to be present in leaf surface extracts from the host-plant Brassica oleracea in quantities as low as 1 ng per gram leaf. In spite of this low level, it stimulates oviposition significantly better than glucobrassicin at higher concentrations, which up till now was known as the most powerful stimulant for D.radicum.     

Preference for plants damaged by conspecific larvae in ovipositing cabbage root flies: influence of stimuli from leaf surface and roots

In laboratory dual-choice assays females of the cabbage root fly, Delia radicum, prefer for oviposition plants with roots damaged by conspecific larvae to undamaged controls. Cauliflower and kale plants were inoculated with root fly eggs (25 per plant) and the hatching larvae were allowed to feed on the roots for various periods of time (1–17 days). After 4 (cauliflower) or 5 (kale) days of larval feeding the oviposition preference was most pronounced and flies laid between 64% and 68% of their eggs near plants with damaged roots. Later, with increasing damage but fewer surviving, and thus actively feeding, larvae, the magnitude of the preference declined. The preference for plants already damaged by conspecific larvae may contribute to the previously observed aggregated distribution of D. radicum eggs in Brassica crop fields.Further experiments revealed that the sensory cues inducing this oviposition preference originate from the complex consisting of the damaged roots, the surrounding substrate (soil) and associated microbes, rather than from the aerial plant parts. In choice assays using the root-substrate complex of damaged and control plants (aerial parts removed), the observed preference for damaged roots was similar to that found for the entire plant but was more pronounced. The damaged roots alone, compared to control roots, received up to 72% (cauliflower) and 75% (kale) of the eggs. By contrast, surrogate leaves sprayed with methanolic leaf surface extracts from the most preferred plants which had been damaged were not discriminated from surrogate leaved sprayed with extracts of the respective control plants. Analysis of glucosinolate levels in methanolic leaf surface extracts revealed that root damage resulted in enhanced concentrations of indole-glucosinolates on the leaf surface in kale but not in cauliflower. Although indole-glucosinolates are oviposition stimulants for the cabbage root fly, the induced changes were apparently too small to influence oviposition behaviour.     

Evaluating the enemies hypothesis in a clover‐cabbage intercrop: effects of generalist and specialist natural enemies on the turnip root fly (Delia floralis)

• 1 The relative importance of the resource concentration hypothesis and the enemies hypothesis was investigated for the turnip root fly Delia floralis in a cabbage–red clover intercropping system compared with a cabbage monoculture.
• 2 Delia floralis egg densities were measured as well as the activity‐densities of generalist predators in a field experiment during two growing seasons. In the second year, a study of egg predation with artificially placed eggs was conducted, in addition to a predator exclusion experiment, to estimate total predation during the season. Parasitization rates were estimated from samples of pupae.
• 3 Delia floralis oviposition was greater in the monoculture during both years. The predator activity‐densities differed between treatments and study years. The known natural enemies of Delia spp., Bembidion spp. and Aleochara bipustulata showed a strong response to a cultivation system with higher activity‐densities in the monoculture. The response, however, appeared to be caused primarily by habitat preferences and not by D. floralis egg densities.
• 4 The reduction in the number of D. floralis pupae in the intercropping may be explained by a disruption in oviposition behaviour caused by the presence of clover because neither predation, nor parasitization rates differed between cultivation systems.

     

A cabbage root fly oviposition deterrent in the frass of garden pebble moth caterpillars

The cabbage root fly, Delia radicum (L.), was deterred from laying eggs on cauliflower plants sprayed with a water-soluble extract of the frass of caterpillars of the garden pebble moth, Evergestis forficalis (L.), which had fed on a range of cruciferous species. Chemical analysis of the extract revealed the presence of a phenolic acid which was not present in the frass of larvae of two other Lepidoptera (Mamestra brassicae (L.) and Plutella xylostella (L.)) that had fed on the same range of cruciferous plants. The deterrent chemical was isolated and identified as sinapic (3,5-dimethoxy-4-hydroxycinnamic) acid. Spraying cauliflower plants with a buffered solution of the acid in water (0.1, 1 and 10 mM) reduced the numbers of cabbage root fly eggs laid by 60–70%. In field experiments, the deterrent effect persisted 5 days after leaves were sprayed with a buffered, aqueous solution of the acid (10 mM).

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Résumé D. radicum L. a pondu beaucoup moins sur des choux-fleurs pulvérisés avec un extrait (polaire) hydrosoluble d'excréments de chenilles de E. forficalis L. qui avaient consommé différentes crucifères. La pulvérisation des choux-fleurs avec une suspension aqueuse d'éluate de méthanol d'une colonne polyamide et d'éluate aqueux d'une colonne florosile a réduit respectivement de 50 et 66% le nombre d'oeufs pondus. L'analyse chimique des fractions actives révèle la présence d'un acide phénolique isolé et identifié comme acide sinapique (3,5-diméthoxy-4-hydroxycinnamique). Cet acide est absent des excréments des chenilles de Mamestra brassicae L. et Plutella xylostella L. qui avaient consommé la même gamme de crucifères. La pulvérisation des plants de choux-fleurs par une solution aqueuse tamponnée d'acide sinapique a réduit la ponte de D. radicum de 60 à 70%. Dans la nature, l'effet dissuasif persiste sur choux-fleurs 5 jours après la pulvérisation avec une solution aqueuse tamponnée à 10 mM d'acide sinapique.

     

The effect of a chemical deterrent, released from the frass of caterpillars of the garden pebble moth, on cabbage root fly oviposition

The cabbage root fly, Delia radicum (L.), was deterred from laying eggs on cauliflower plants that had been sprayed with a suspension of the frass of caterpillars of the garden pebble moth, Evergestis forficalis (L.). Polar extracts of the frass deterred oviposition irrespective of the cruciferous plant species on which the caterpillars had been feeding. Non-polar extracts of the frass had no effect. Spraying plants with macerates from Brassica leaves stimulated fly oviposition whereas spraying plants with macerates from garlic mustard leaves deterred fly oviposition. Macerates from the leaves of all other plants tested had no effect. In field experiments the deterrent effect persisted 2–3 days after leaves were sprayed with frass extracts. Plants infested with feeding caterpillars and contaminated with only a few discrete frass pellets were as deterrent to the fly as those sprayed with frass suspensions.

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Résumé D. radicum a évité de pondre sur des pieds de chou-fleur, qui avaient été pulvérisés avec une suspension d'excréments d'E. forficalis. Les extraits polaires de ces excréments empêchent la ponte quelle que soit l'espère de crucifère sur laquelle les chenilles se sont alimentées. Les extraits non polares sont sans effet. Dans la nature, l'action dissuadante persiste 2 à 3 jours après la pulvérisation des feuilles avec les extraits d'excréments. La pulvérisation tous les deux jours a maintenu l'action dissuadante. Des plantes contaminées par des chenilles en train de s'alimenter et souillées par quelques crottes sont aussi dissuadantes pour la mouche que celles pulvérisées avec une suspension d'excréments.

     

Preference of the cabbage root fly, Delia radicum (L.), for coloured traps: influence of sex and physiological status of the flies, trap background and experimental design

Abstract. Catches of Delia rudicum (L.) (Diptera: Anthomyiidae) were compared in water traps that reflected predominantly wavelengths shorter (violet and blue traps) and longer (green and yellow traps) than 500 nm.Traps were positioned in choice and no-choice situations against backgrounds of bare soil and weeds in the field and against backgrounds of brown and green paper in the laboratory.The physiological status of the flies was modified in the laboratory by denying them access to food sources and oviposition sites.
Males discriminated significantly more clearly than females between yellow and blue traps.The discrimination between yellow and blue traps was significantly more pronounced when the traps were presented in the choice than in the no-choice situation in both sexes.Green background (weeds and green paper) was highly preferred for landing and thus competed with the traps to such an extent that few flies were caught when non-preferred violet and blue traps were sited on green backgrounds.Flies seldom landed on the brown background (soil and brown paper) which resulted in the relative increase of catches in the non-preferred violet and blue traps.The preference for yellow traps was innate even in young flies with immature egg-follicles.Females that were ready to lay eggs, even those deprived of an oviposition site till the age of 8 days, also preferred yellow traps.In the no-choice situation, flies deprived of food landed with the same frequency in yellow and blue traps.Food deprivation, however, did not affect preference for yellow traps over the blue traps presented in a choice situation.     

Ovarian development status and population characteristics of Sogatella furcifera (Horváth) and Nilaparvata lugens (Stål): implications for pest forecasting

Migratory behaviour in insects correlates with reproductive development in females, and migration often occurs during the pre‐reproductive stage of adults. The relationship between ovarian development and population status of the white‐backed planthopper Sogatella furcifera (Horváth) and the brown planthopper Nilaparvata lugens (Stål) was evaluated. Females of both species were captured in rice fields and light traps and then dissected in double‐season rice‐farming regions of southern China. The ovarian development of S. furcifera and N. lugens was divided into five levels, following previous studies. The population statuses of both species were examined based on the ovarian development of female adults caught in rice paddies. The ovarian development in N. lugens females caught in light traps mostly ranged from level I to level II, whereas that in S. furcifera females caught in light traps mostly ranged from level I to level III. During peak immigration, ovarian development in N. lugens females was mainly at level II, whereas that in S. furcifera females was mainly at level II and sporadically at level III. During peak emigration, both S. furcifera and N. lugens showed level I ovarian development. The temporal dynamics of ovarian development in light trap catches revealed that (i) significant emigration and partial immigration periods occur in S. furcifera, with ovarian development mainly at level I and sporadically from level II to level III and (ii) numerous immigrants of N. lugens were detected during sedentary and local breeding periods. The temporal dynamics of ovarian development provides more information than does the paddy population. Thus, this study proposes another method for pest forecasting, which is more precise and efficient than conventional forecasting methods such as light trap catching and monitoring population dynamics in rice fields.     

Patterns in root traits of woody species hosting arbuscular and ectomycorrhizas: implications for the evolution of belowground strategies

Root traits vary enormously among plant species but we have little understanding of how this variation affects their functioning. Of central interest is how root traits are related to plant resource acquisition strategies from soil. We examined root traits of 33 woody species from northeastern US forests that form two of the most common types of mutualisms with fungi, arbuscular mycorrhizas (AM) and ectomycorrhizas (EM). We examined root trait distribution with respect to plant phylogeny, quantifying the phylogenetic signal (K statistic) in fine root morphology and architecture, and used phylogenetically independent contrasts (PICs) to test whether taxa forming different mycorrhizal associations had different root traits. We found a pattern of species forming roots with thinner diameters as species diversified across time. Given moderate phylogenetic signals (K = 0.44–0.68), we used PICs to examine traits variation among taxa forming AM or EM, revealing that hosts of AM were associated with lower branching intensity (r_PIC = −0.77) and thicker root diameter (r_PIC = −0.41). Because EM evolved relatively more recently and intermittently across plant phylogenies, significant differences in root traits and colonization between plants forming AM and EM imply linkages between the evolution of these biotic interactions and root traits and suggest a history of selection pressures, with trade-offs for supporting different types of associations. Finally, across plant hosts of both EM and AM, species with thinner root diameters and longer specific root length (SRL) had less colonization (r_PIC = 0.85, −0.87), suggesting constraints on colonization linked to the evolution of root morphology.     

Characterizing differences in precipitation regimes of extreme wet and dry years: implications for climate change experiments

Climate change is intensifying the hydrologic cycle and is expected to increase the frequency of extreme wet and dry years. Beyond precipitation amount, extreme wet and dry years may differ in other ways, such as the number of precipitation events, event size, and the time between events. We assessed 1614 long‐term (100 year) precipitation records from around the world to identify key attributes of precipitation regimes, besides amount, that distinguish statistically extreme wet from extreme dry years. In general, in regions where mean annual precipitation (MAP) exceeded 1000 mm, precipitation amounts in extreme wet and dry years differed from average years by ~40% and 30%, respectively. The magnitude of these deviations increased to >60% for dry years and to >150% for wet years in arid regions (MAP<500 mm). Extreme wet years were primarily distinguished from average and extreme dry years by the presence of multiple extreme (large) daily precipitation events (events >99th percentile of all events); these occurred twice as often in extreme wet years compared to average years. In contrast, these large precipitation events were rare in extreme dry years. Less important for distinguishing extreme wet from dry years were mean event size and frequency, or the number of dry days between events. However, extreme dry years were distinguished from average years by an increase in the number of dry days between events. These precipitation regime attributes consistently differed between extreme wet and dry years across 12 major terrestrial ecoregions from around the world, from deserts to the tropics. Thus, we recommend that climate change experiments and model simulations incorporate these differences in key precipitation regime attributes, as well as amount into treatments. This will allow experiments to more realistically simulate extreme precipitation years and more accurately assess the ecological consequences.     

Phylogeography, genetic structure and population divergence time of cheetahs in Africa and Asia: evidence for long-term geographic isolates

The cheetah (Acinonyx jubatus) has been described as a species with low levels of genetic variation. This has been suggested to be the consequence of a demographic bottleneck 10 000–12 000 years ago (ya) and also led to the assumption that only small genetic differences exist between the described subspecies. However, analysing mitochondrial DNA and microsatellites in cheetah samples from most of the historic range of the species we found relatively deep phylogeographic breaks between some of the investigated populations, and most of the methods assessed divergence time estimates predating the postulated bottleneck. Mitochondrial DNA monophyly and overall levels of genetic differentiation support the distinctiveness of Northern‐East African cheetahs (Acinonyx jubatus soemmeringii). Moreover, combining archaeozoological and contemporary samples, we show that Asiatic cheetahs (Acinonyx jubatus venaticus) are unambiguously separated from African subspecies. Divergence time estimates from mitochondrial and nuclear data place the split between Asiatic and Southern African cheetahs (Acinonyx jubatus jubatus) at 32 000–67 000 ya using an average mammalian microsatellite mutation rate and at 4700–44 000 ya employing human microsatellite mutation rates. Cheetahs are vulnerable to extinction globally and critically endangered in their Asiatic range, where the last 70–110 individuals survive only in Iran. We demonstrate that these extant Iranian cheetahs are an autochthonous monophyletic population and the last representatives of the Asiatic subspecies A. j. venaticus. We advocate that conservation strategies should consider the uncovered independent evolutionary histories of Asiatic and African cheetahs, as well as among some African subspecies. This would facilitate the dual conservation priorities of maintaining locally adapted ecotypes and genetic diversity.