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These second edition guidelines, updated from the 2007 version (Marchiondo et al., 2007), are intended to assist the planning and conduct of laboratory and clinical studies to assess the efficacy of ectoparasiticides applied to dogs or cats for the purpose of treating, preventing and controlling flea and tick infestations. Major revisions to this second edition include guidelines on the assessment of systemic flea and tick products, an update of the geographical distribution of the common fleas and ticks species on dogs and cats, determination of flea and tick efficacy based on geometric versus arithmetic means with respect to geographic regulatory agencies, modification of tick categorization in the assessment of efficacy, expanded guidelines on repellency and anti-feeding effects, enhanced practical field study guidance, and considerations on the ranges of flea and ticks for infestations in laboratory studies. The term ectoparasiticide includes insecticidal and acaricidal compounds, as well as insect growth regulators. The range of biological activities from animal treatment that are considered include: repellency and anti-feeding effects, knockdown, speed of kill, immediate and persistent lethal effects, and interference with egg fertility and subsequent development of off-host life cycle stages. Information is provided on the selection of animals, dose determination, dose confirmation and field studies, record keeping, interpretation of results and animal welfare. These guidelines are also intended to assist regulatory authorities involved in the approval and registration of new topical or systemic ectoparasiticides, and to facilitate the worldwide adoption of harmonized procedures.

The objective of this study was to determine the species spectrum of ixodid ticks infesting birds in Free State Province, South Africa. To this end a large number of birds belonging to several species were examined for ticks and a total of 180 birds belonging to 39 species at 17 localities were infested, and ticks belonging to eight species were recovered. The immature stages of only two, namely Amblyomma marmoreum and Hyalomma marginatum rufipes, were sufficiently prevalent and numerous to safely assume that they regularly use birds as hosts. Helmeted guineafowls, Numida meleagris, were the most heavily infested and one harboured a total of 319 larvae and four nymphs. Amongst the other species an eastern clapper lark, Mirafra fasciolata, was infested with 69 larvae and a nymph, but no other bird harboured more than 40 ticks. The larvae and nymphs of H. m. rufipes were most numerous on birds from April to August.

Two hundred and seventy-three elephant shrews, consisting of 193 Elephantulus myurus, 67 Elephantulus edwardii and 13 animals belonging to other species, were examined for ixodid ticks at 18 localities in South Africa and Namibia. The immature stages of Ixodes rubicundus, Rhipicentor nuttalli, Rhipicephalus warburtoni and a Rhipicephalus pravus-like tick were the most numerous of the 18 tick species recovered. Substantial numbers of immature Rhipicephalus arnoldi, Rhipicephalus distinctus and Rhipicephalus exophthalmos were also collected from elephant shrews at particular localities. Larvae of I. rubicundus were most numerous on E. myurus in Free State Province from April to July and nymphs from June to October. Larvae of R. nuttalli were most numerous on these animals during April, May, August and September, and nymphs in February and from April to August. The immature stages of R. warburtoni were collected from E. myurus only in Free State Province, and larvae were generally most numerous from December to August and nymphs from April to October.

Two hundred and twenty-five small mammals belonging to 16 species were examined for ticks in Free State, Mpumalanga and Limpopo Provinces, South Africa, and 18 ixodid tick species, of which two could only be identified to genus level, were recovered. Scrub hares, Lepus saxatilis, and Cape hares, Lepus capensis, harboured the largest number of tick species. In Free State Province Namaqua rock mice, Aethomys namaquensis, and four-striped grass mice, Rhabdomys pumilio, were good hosts of the immature stages of Haemaphysalis leachi and Rhipicephalus gertrudae, while in Mpumalanga and Limpopo Provinces red veld rats, Aethomys chrysophilus, Namaqua rock mice and Natal multimammate mice, Mastomys natalensis were good hosts of H. leachi and Rhipicephalus simus. Haemaphysalis leachi was the only tick recovered from animals in all three provinces.

The susceptibility of the larval offspring of engorged female Boophilus decoloratus, and of the engorged females, collected from cattle on the dairy farms Brycedale, Sunny Grove and Welgevind in the Eastern Cape Province, South Africa, was tested against the acaricides amitraz, chlorfenvinphos and cypermethrin. Resistance was determined by means of the Shaw Larval Immersion Test (SLIT) for larvae and the Reproductive Estimate Test (RET) and Egg Laying Test (ELT) for adults. At Brycedale the tests all indicated resistance to chlorfenvinphos, and RET and ELT indicated resistance to amitraz and emerging resistance to cypermethrin. At Sunny Grove, B. decoloratus was resistant to cypermethrin using SLIT and exhibited emerging resistance to chlorfenvinphos with SLIT and to cypermethrin with both RET and ELT At Welgevind, resistance was recorded against chlorfenvinphos (SLIT) and against cypermethrin (ELT), and emerging resistance against permethrin (RET). The results obtained with RET and ELT were generally comparable, but often differed from those obtained with SLIT. Resistance could be detected within 7 days with ELT compared to 42 days with RET and 60 days with SLIT.

Several surveys of ticks infesting dogs belonging to owners in resource-limited and more affluent communities have been conducted in South Africa, but no such investigation has been carried out in the Free State Province of this country. The present study was initiated to meet this shortcoming. Ticks were collected from dogs at six localities in, and to the east of the city of Bloemfontein in the central region of the province. Three of these localities could be classed as resource-limited and two as affluent, while the sixth locality was an animal shelter serving all members of the public. Adult ticks belonging to nine ixodid tick species were collected, of which Rhipicephalus sanguineus was the most numerous. Significantly more R. sanguineus was collected from dogs at resource-limited than at more affluent localities. The greatest proportions of these ticks attached to the backs and necks of the dogs, with the proportions being larger in long-haired than in short-haired dogs. Most R. sanguineus were collected during the warmer months particularly from January to April. The greatest proportions of Haemaphysalis leachi, the next most numerous species, were also collected from the backs and the necks of the dogs. Most of these were present during the period September to November.

Ixodid ticks were collected from 104 wild carnivores belonging to 23 species in various nature reserves and on several farms in all nine provinces of South Africa. Seven feral cats in a nature reserve were also examined. Twenty-four tick species belonging to seven genera were recovered and identified. Amongst these ticks we consider the adults of Haemaphysalis leachi, Haemaphysalis spinulosa, Haemaphysalis zumpti, Ixodes rubicundus, Rhipicentor nuttalli, Rhipicephalus simus and Rhipicephalus turanicus to be true parasites of wild carnivores. Although numerous adult Rhipicephalus appendiculatus and Rhipicephalus zambeziensis were collected from some lions these were either sick or old animals. The immature stages of seven species regularly utilized wild carnivores as hosts on an opportunistic basis.

This questionnaire survey was conducted amongst 200 farmers in the resource-poor, urban and per-urban environments of Botshabelo and Thaba Nchu towns in the Free State Province of South Africa. The questionnaire was divided into seven sections, namely demography, livestock, cattle, parasites, parasite control, livestock diseases and problems experienced. A total of 87.5% of the livestock owners were not employed (unemployed or pensioner). Cattle constituted more than 50% of the live-stock units owned in the area and farmers owned an average of 9.33 +/- 0.812 head of cattle. A total of 193 (96.5%) of the farmers indicated that milk was the most important product from their cattle. Only 26% of them slaughtered their own cattle for meat consumption. Eighty-eight percent of them indicated that external parasites on their livestock presented a problem, but only 72.9% of farmers implemented any tick control measures. Less than half (45.5%) of the farmers who attempted to control ticks used commercial acaricides. The remainder used various other methods, including the application of used engine oil and household detergents. Amongst the clinical diseases observed in their cattle, dry gallsickness was mentioned most often (20%). This figure, however, is believed to be inaccurate because dry gallsickness may be a clinical manifestation of some of the other diseases mentioned by the farmers, such as anaplasmosis, foreign body obstruction of the gastro-intestinal tract by plastic bags, pneumonia and mastitis. Animal husbandry problems experienced by the farmers included pollution (i.e. ingestion of plastic bags and string by their livestock), availability of water and theft.

Relative resistance levels of cattle against tick infestations in the communal grazing area of Botshabelo in the south-eastern Free State were determined. The objective was to establish whether differences in resistance can be exploited to contribute to tick control methods used by small-scale farmers in resource-poor environments. Ten cows (Bos taurus crosses) between the ages of 18 months and four years were used and tick counts were conducted once a month over a period of 12 months to compare their total tick burdens. Tick burdens of the various animals were compared mutually as well as with the mean tick burden of the group as a whole. Tick numbers varied throughout the year on all individuals but some animals consistently tended to have either higher or lower numbers than the mean of the group. Tick burdens on cattle classified as having a relatively low resistance to tick infestations increased eleven-fold from January to June 1996 compared to a six-fold increase on cattle categorized as belonging to the high resistance group. Twenty-eight percent of the cattle in the total study group carried 50% of the ticks collected (60,079). It is recommended that farmers in the region visually assess B. decoloratus burdens, the most abundant tick species, and sell or cull the most susceptible animals first in their normal program of utilization of the animals. This should eventually result in the direct improvement of the overall tick resistance of their cattle herds.

The objectives of this study were to determine the diversity, seasonal dynamics and abundance of ticks infesting cattle in urban, small-scale farming communities in and around Botshabelo and Thaba Nchu in the eastern Free State Province, South Africa. A total of ten cattle, ear-tagged for individual identification, were investigated monthly at each of five localities. Adult ticks were removed from the right hand side of each animal and placed in containers filled with 70% ethanol. They were subsequently identified and their numbers quantified. Immature Otobius megnini were counted but not removed. A total of 244,538 adult ticks of ten different species were collected over the 12-month study period. The tick species, in decreasing order of relative abundance, were: Boophilus decoloratus (87.26%), Rhipicephalus evertsi evertsi (6.86%), Hyalomma marginatum rufipes (2.42%), Otobius megnini (1.85%) Rhipicephalus follis (0.76%), Rhipicephalus gertrudae (0.54%), Rhipicephalus sp. (0.21%), Ixodes rubicundus (0.08%), Hyalomma truncatum (0.01%) and Margaropus winthemi (0.004%). The three most abundant species, namely B. decoloratus, R. evertsi evertsi and H. marginatum rufipes, occurred at all localities but with significant differences in abundance. M. winthemi ticks occurred only in the Thaba Nchu area and were not found at any of the there localities in Botshabelo. Significant differences in tick burdens between the six warm months (September to February) and the six cooler months (March to August) were found for most of the species recorded. Boophilus decoloratus occurred in significantly higher numbers in autumn (March to May) and winter (June to August) compared to spring (September to November) and summer (December to February), with 76.8% of the total B. decoloratus burden occurring during the cooler months.

A seroepidemiological study was conducted on 151 cattle from the Botshabelo and Thaba Nchu areas in the central Free State Province of South Africa, two areas where small scale, peri-urban cattle farming is practised. An indirect fluorescent antibody test was used to test for Babesia bigemina and B. bovis antibodies. To test for Anaplasma marginale antibodies a competitive inhibition enzyme-linked immunosorbent assay method was used. There were no significant differences in serological test results between the cattle from Botshabelo and those from Thaba Nchu. The herd (two areas combined) had an average seroprevalence of 62.42% to B. bigemina, 19.47% to B. bovis and 98.60% to A. marginale. Based on the percentage of cattle that were seropositive to B. bigemina the immune status of cattle in the Botshabelo-Thaba Nchu area is approaching a situation of endemic stability. With reference to A. marginale, the high seroprevalence is indicative of a situation of endemic stability. The occurrence of B. bovis antibodies in the cattle is difficult to explain as Boophilus microplus ticks do not occur in the area in which the study was conducted.

Successful prophylaxis of paralysis, induced in small stock by feeding female Ixodes rubicundus, is dependent on the accurate determination of the commencement of seasonal activity by the tick. The commencement of this activity was recorded for 11 consecutive years on a farm in the south-western Free State, South Africa, and for shorter periods on other farms, some of these in regions with markedly colder climates. The colder the mean minimum atmospheric temperatures during the 2 months preceding the start to tick activity, the earlier it commenced. This could differ by 4 weeks from year to year on the same farm. In a region with a low effective temperature activity commenced between 3-8 weeks earlier than in a region with a higher effective temperature.

A study was conducted to compare the species of adult ixodid ticks on Friesian and Bonsmara cattle and their seasonal dynamics. Between five and ten animals of each breed were kept on natural False Upper Karoo vegetation, typical of the south-western Free State. Between May 1990 and October 1991, the animals were examined once or twice a month for the presence of adult ticks which were removed, counted and identified. Six species of ixodid ticks were recorded. The species composition on the two cattle breeds was remarkably similar. Ixodes rubicundus was dominant (55-57.7% of all ticks collected), followed by Hyalomma marginatum rufipes (28.9-31.5%) and Rhipicephalus punctatus *6.2-6.7%). Only small numbers of Hyalomma truncatum, Rhipicephalus evertsi evertsi and Rhipicephalus gertrudae were recorded. Almost twice as many ticks were collected from Friesian as from Bonsmara cattle. Ixodes rubicundus showed a typical winter-activity period. The other species were either active mostly during the warmer months, or their numbers were to low to determine any seasonal pattern. Apart from information on the diversity of ticks which infest cattle in the region, the study has also shown a greater resistance by Bonsmara cattle (a mixed Bos taurus/Bos indicus breed) to tick infestation. Bos indicus cattle can play an increasingly important role in integrated tick-management practices in southern Africa.

The responsiveness of Ixodes rubicundus ticks on questing substrates and the success of their attachment to non-living substrates were investigated. The purpose of this study was to relate responsiveness to conditions of temperature and humidity, to compare the efficacy of flagging and dragging methods to estimate the size of populations of adult I. rubicundus in the field, and to determine the possible influence of host odour on the efficacy of these methods. Responsiveness was tested under varying conditions of temperature and humidity under field conditions, and the same ticks were used to determine the duration of attachment to a flannel cloth, either impregnated with host odour from sheep wool or without it. Flagging and dragging methods were compared under laboratory conditions with cloths either treated with host odour or not. Within the range of ambient temperatures recorded during this study (7-25 degrees C) most of the ticks (86%) were responsive. No obvious relationship between temperature, relative humidity and responsiveness of ticks was evident. Ticks remained on average 32.9 seconds (n = 64) on cloth treated with host odour compared to 9.1 seconds (n = 54) on untreated cloth. Flagging was 1.5-1.7 times as effective as dragging and treatment of the cloths with host odour increased the efficacy 2.4 (dragging) to 2.8 (flagging) times. The reactions of ticks to external stimuli should, therefore, be taken into account to maximize field sampling success.

Engorgement, mass at repletion and moulting success of immature Ixodes rubicundus after sequential infestations of their natural hosts, rock elephant shrews (Elephantulus myurus), were investigated under laboratory conditions. The reaction of E. myurus is characterized by inefficient or non-existent anti-tick immunity which enables immature I. rubicundus to attach and engorge successfully and ensures a high moulting success rate.

Detachment rhythms of immature Karoo paralysis ticks (Ixodes rubicundus) from their natural host, the rock elephant shrew (Elephantulus myurus), were investigated under laboratory and natural conditions. Larvae and nymphs detach mainly during the day. Peak detachment always occurred after the commencement of artificial as well as natural light cycles. The detachment of larvae appears to be more synchronous than that of nymphs, with a larger percentage of larvae detaching over a shorter period of time. The detachment rhythms of I. rubicundus are strongly correlated with the activity patterns of their hosts. This enhances not only the survival, but also the dispersion and host contact of subsequent stages.

Mortality due to Karoo paralysis amongst livestock is of considerable economic importance in South Africa. Indications are that the extent of vertical migration of adult ticks, to reach questing positions on grasses, is influenced by sudden climatic changes. Such influences as well as selection of questing substrates were investigated under controlled conditions. Vertical migration of adult Ixodes rubicundus on copper rods was determined in a square metal arena within a closed glass container in a room in which temperature and light could be controlled. Significantly more female and male ticks migrated vertically on 1.5 mm diameter rods than on 4.0 mm rods and on 450 mm long rods than on 100 mm rods. More ticks of both sexes migrated vertically at RH > 80% than at RH < 50% and at 12 degrees C than at 21 degrees C. More female than male and more older than younger ticks migrated vertically in all experiments. The results of this study contribute towards an understanding of the epidemiology of Karoo paralysis.

The appetence responses of the ixodid ticks Ixodes rubicundus and Rhipicephalus punctatus to various stimuli were quantified under laboratory conditions. A greater percentage of I. rubicundus generally responded to the different stimuli at lower than at higher temperatures while the converse was usually true for R. punctatus. Radiation heat and shadowing evoked a response in the greatest percentage of I. rubicundus, whereas CO2 and host odour were responsible for a similar reaction in R. punctatus. A single perturbation of the substrate caused more than 50% of I. rubicundus to respond while constant perturbation resulted in a progressive decrease in the proportion of ticks reacting. Significantly more R. punctatus than I. rubicundus were attracted to a CO2 trap and the R. punctatus were also attracted to the trap over a greater distance. Significantly more older than younger ticks of both species responded to the various stimuli.

The sites of attachment of female Karoo paralysis ticks (Ixodes rubicundus) were determined on Friesian, Bonsmara and Nguni cattle in the south-western Orange Free State. The largest proportion of the ticks (> 30%) attached to the forelegs and shoulders of the cattle. Contrary to the findings on small stock, no significant differences in the preferred sites of attachment were evident between the breeds of cattle.

The seasonal occurrence of Karoo paralysis in relation to the infestation density of female Ixodes rubicundus was investigated in an enzootic area of the southwestern Orange Free State, South Africa. Young and adult Angora goats were exposed to infestation during two paralysis tick seasons. Only the young goats became paralysed and their infestation densities (female ticks kg-1 host body weight) were significantly higher than those of healthy animals. The infestation densities on paralysed goats varied between 0.64 and 3.56 ticks kg-1. During the 1988 season, 30% of the naive young goats became paralysed compared with 67% the following season. Twenty-nine percent of previously paralysed goats were paralysed again during the same season. Paralysis occurred during periods of peak tick activity (April-June).

The aim of this study was to quantify the infestation densities of Karoo and brown paralysis ticks on sheep and goats and explain it in terms of the etho-ecology of these ticks and their domestic hosts. The Karoo paralysis tick usually quests from a vantage point on the vegetation whereas the brown paralysis tick displays an appetence response from the ground and mainly engages hosts that are prostrate. Both tick species are confined chiefly to hilly areas. Temporal differences in the infestation densities of the hosts within and between tick species were evident. These differences were related to disparities in the spatial distribution of the hosts, their activity patterns and the specific appetence responses of the two tick species. Differential climatological conditions affected the quality of forage in areas with a varied topography and the feeding preferences of hosts influenced tick-host sympatry and hence infestation densities.

One hundred and seventeen scrub hares, Lepus saxatilis, were examined for ixodid ticks in various regions of the Cape Province. They were infested with 18 tick species and the seasonal abundances of the immature stages of Amblyomma hebraeum, Amblyomma marmoreum, Hyalomma marginatum rufipes, Rhipicephalus appendiculatus and Rhipicephalus glabroscutatum and all stages of Rhipicephalus oculatus and the adults of a Rhipicephalus sp. (near R. oculatus) were determined. Seventy-two scrub hares on 3 farms in the Orange Free State harboured 10 ixodid tick species and the seasonal abundances of the immature stages of H. marginatum rufipes and Hyalomma truncatum and all stages of Rhipicephalus punctatus were determined. Thirty Cape hares, Lepus capensis, examined in the northern Cape Province harboured 5 tick species and the seasonal abundances of the immature stages of H. marginatum rufipes and H. truncatum were determined. Thirty-four Cape hares examined in the south-western Orange Free State carried 6 tick species and the seasonal abundance of immature H. marginatum rufipes was determined on these animals. Twenty-eight Smith's red rock rabbits examined on 2 farms in the south-western Orange Free State were infested with 7 tick species.

Significant differences in the distribution of brown paralysis ticks on various age classes of Angora goats were recorded. In kids, most (greater than 98%) of the ticks attached to the head and ears, whereas in older groups, in addition to the ears, a high proportion (greater than 20%) of ticks also attached to the ventral side of the neck. There were significant differences in the mean infestation densities of both male and female R. punctatus in kids and older animals. These differences were, however, only significant for the first two sample dates involving kids, and are probably related to behavioural attributes of the kids which enhance tick/host contact. Newly born Angora goat kids are considered a high-risk group with regard to paralysis caused by the brown paralysis tick. Methods of avoiding mortality amongst kids are suggested.

Sixteen species of ixodid ticks were collected over varying periods of time from 6 species of ground-frequenting birds and 15 species of small and large mammals in the Mountain Zebra National Park, Karoo, Cape Province. Margaropus winthemi followed by Rhipicephalus glabroscutatum and Rhipicephalus evertsi evertsi were the most abundant species. The host preferences of 14 tick species and the seasonal abundances of 13 species were determined. Small numbers of immature stages of only a few tick species were generally recovered from ground-frequenting birds, mice and rats. Amongst the slightly larger animals the smallest number of ticks and species were recovered from springhares and the largest numbers from scrub hares. Amongst the larger mammals, very few ticks were harboured by springbuck and black wildebeest, while heavy infestations of several species were encountered on Cape mountain zebras and eland.

Adult ixodid ticks were collected at 2-weekly intervals for a period of 23 consecutive months from 15 to 20 Angora goats on a farm in the south western Orange Free State. A total of 6 ixodid tick species were recovered. Rhipicephalus punctatus was the most abundant and prevalent tick. It was present from spring to late summer. Ixodes rubicundus was the next most abundant tick and was present mainly from March or April to July with peak numbers present in April or May. The onset of this tick's activity appeared to be stimulated by low atmospheric temperatures.

Spatial distribution of the Karoo paralysis tick (Ixodes rubicundus) has been studied in a paralysis enzootic area of the southwestern Orange Free State. Significantly more females than males were collected from the vegetation. Most ticks (99%) used grass as questing sites at a height of approximately 45 cm (range 10-97 cm), which correlates with the size of host animals. No significant differences between the questing height of male and female ticks were evident. The ticks were found on most of the grass species present and, except for the genus Eragrostis, no specific preference was evident. When questing, the capitulum of I. rubicundus is usually (90%) directed towards the ground. Seventy-eight percent of the ticks occurred singly on a specific grassblade or flowerhead. In those cases where two ticks occurred together, no sexual aggregation was evident. Most ticks occurred under or close (mean = 62 cm) to the crown cover of specific shrub or tree species. These plant species were characterized by a dense crown cover with a mat of decaying leaf litter underneath. The preference of the tick for wild olive trees (Olea europaea africana) suggests that, except for the physical characteristics of this tree species, host factors may also be important. Significantly more ticks were distributed on the cooler, more protected southern sides of vegetation in comparison to the northern sides.

The ratio of variance to mean (s2/means) of numbers of Ixodes rubicundus (males and females) per sheep was used as a measure of the dispersion pattern of this tick within a naturally infested population of sheep. Sheep were divided into two groups, one with previous contact with I. rubicundus and the other with no previous contact. Samples of 10 sheep from each group were searched for ticks on a weekly basis for 19 weeks during the winter of 1986. In each group, both male and female ticks had an s2/means ratio greater than 1 for the majority of observation periods indicating an aggregated dispersion pattern. The degree of aggregation was greater with increasing density of the ticks. The implications of these results are discussed both in relation to sampling procedures and potential control strategies.

The weights of feeding female Ixodes rubicundus attaching to sheep was studied in a natural focus of Karoo paralysis. The studies showed fewer females in the middle weight ranges and more in the heavier and lighter categories than would be expected if the distribution was normal. Weights were also strongly skewed towards the lighter females between 0 and 400 mg. No differences in the pattern of attachment weights occurred over the four experimental groups used, suggesting that previous contact of sheep with I. rubicundus had no effect on the ability of the tick to engorge.

Male Ixodes rubicundus were found in copula with attached females in 59% of individuals. The number of males attaching to the female integument or to the the host negligible. In laboratory experiments, males enclosed with females in a small vial had significantly longer life spans than solitary males. Paired males were found to be in copula on 20-34% of observations. Solitary males lost weight while paired males gained weight over a 2-day period. However, the specific energy content of the 2 groups remained constant.

Abstract. The Karoo Paralysis tick, Ixodes rubicundus Neumann (Acari: Ixodidae), is a semi-voltine ixodid that survives stressful environmental conditions using morphogenetic diapause (eggs and engorged nymphs) and desiccation resistance. Both photoperiod and temperature influence diapause induction in the engorged nymph. Ixodes rubicundus nymphs are typical long-day photoperiodic organisms. The critical photoperiod is ∼ 13.5 h light, 10.5 h dark, and they display a thermolabile response. The period between detachment and apolysis in engorged nymphs is modified by photoperiod; however, apolysis to ecdysis is not affected by photoperiod. Thus, initiation of development, but not the actual process is controlled by photoperiod. Most engorged nymphs delayed metamorphosis when exposed to short-day regimen (LD 12 : 12 h) after feeding. Nymphs exposed to pre- and post-feeding long-day regimen (LD 14 : 10 h) developed. Times for 50% of nymphs to apolyse when exposed to photoperiods of LD 14 : 10 h, 13.5 : 10.5 h, 13 : 11 h and 12 : 12 h were 28, 36, 40 and 58 days, respectively. Times for 50% of engorged nymphs to ecdyse ranged from 38 to 40 days after apolysis. Nymphs were sensitive to photoperiodic exposures before, during and after feeding. Nymphs exposed to long day (LD 14 :10 h) before and during feeding, moulted at 20°C; however, most exposed to 10°C followed by 20°C (post-feeding) went into diapause. Both short- (10 : 14 h) and long- (14 : 10 h) day exposed engorged nymphs survived 45 days at 0% r.h. (n = 73), but diapause-destined ticks kept at 13°C lost the least mass (29.5 ± 9.5%, SD), while nondiapause ticks at 23°C lost the most (48.7 ± 8.2%, SD). Termination of diapause and transition to development probably coincides with a definite increase of water vapour uptake by engorged nymphs. Comparatively, I. rubicundus engorged nymphs are more desiccation tolerant than a North-American counterpart, Amblyomma... [ABSTRACT FROM AUTHOR]