The cervid ked ( Lipoptena cervi, L. , Diptera, Hippoboscidae ) is an ectoparasitic louse fly of the elk ( Alces Alcess ) and other deers. It spread to Finland in the early 1960 ‘s from the South-East ( Hackman et al. , 1983 ) and at present it is a common parasite in the southern parts of Finland with its northern distribution bound bit by bit spreading due norths. This means that it will be shortly in contact with another possible host, the semi-domesticated caribou ( Rangifer tarandus tarandus ) . In the former Soviet Union, the cervid ked parasitized practically every elk in Belarus with 1144-5082 keds per elk ( Ivanov, 1981 ) and in the Leningrad part between 200-300 keds per elk with a maximal figure of about 1000 flies ( Popov, 1965 ) . In Poland, this parasite could be found in 93 % of elk, 78 % of ruddy cervid ( Cervus elaphus ) , 64 % of roe cervid ( Capreolus Capreolus ) ( Kadulski, 1996 ) and 76 % of fallow cervid ( Dama Dama ) ( Szczurek & A ; Kadulski, 2004 ) . The general public considers the cervid ked a plague. It attaches thirstily to human vesture and hair, drops it wings and bites rather frequently but has non been observed to parasitize worlds for good. However, inauspicious tegument reactions largely interpreted as allergic are on a regular basis encountered ( Rantanen et al. , 1982 ; Reunala et al. , 2008 ) .
As a louse fly fly, the cervid ked is morphologically and physiologically adapted to the ectoparasitic life ( Metcalf & A ; Metcalf, 1993 ) . It is dorsoventrally flattened with a difficult exoskeleton. It drops its wings instantly upon fond regard on the host ( Hackman et al. , 1983 ) and has big claws to heighten the fond regard and to forestall withdrawal ( Haarlov, 1964 ) . The cervid ked is a pool feeder: it cuts the tegument of the host and consumes the blood accumulating in the lesion. The generative scheme of the cervid ked is live-bearing ( Meier et al. , 1999 ) . The egg hatches in the generative piece of land and the development larva is fed by secernments produced by the female. One female can bring forth 20-32 pupae ( Popov, 1965 ; Ivanov, 1981 ) , somewhat more than the related sheep ked ( Melophagus ovinus ) ( Metcalf & A ; Metcalf, 1993 ) . The pupae autumn from the pelt of the host to the forest floor or snow and during the following fall the grownup flies emerge ( Popov, 1965 ) . The cervid ked does non wing actively seeking for a host but stays near to its hatching site and delaies for a suited host to get. Between 14-24 A°C the flight distance is ad 50 m, at 7-11 A°C up to 15 m and below 7 A°C keds become inactive. The cervid ked has one coevals per twelvemonth, the outgrowth of grownups starts at the terminal of July and the flight season ends at the beginning of November ( Ivanov, 1981 ) . A cervid ked lives for 120-180 yearss after it has selected a host. The grownups die out bit by bit and from February-March elk remain unparasitized until the following fall ( Popov, 1965 ) .
The purpose of this survey was to find the basic forms and features of cervid ked parasitism on the elk in Eastern Finland. The specific purposes were to analyze 1 ) how the age and sex of the elk affect the parasitism and 2 ) how the denseness of cervid keds differs between anatomical parts of the moose tegument. It can be hypothesized that the cervid ked would be present in high Numberss on Finnish elk and that the choice of an anatomical part of the host would depend on, e.g. , fur features such as hair length.
Materials and methods
Whole elk teguments ( n = 23 ) were collected during the legal hunting season between October 7 and November 26, 2006 in Liperi commune, Eastern Finland ( 62A°31’N, 29A°08’E ) . The age, sex and general status of the elk were determined. The organic structure mass was estimated from the weight of the carcase ( Wallin et al. , 1996 ) . The thickness of hypodermic fat reflecting the nutritionary province was measured from an scratch in the gluteal part after clambering every bit instructed by the Finnish Game and Fisheries Research Institute ( 2009a ) . Immediately after clambering, the tegument was divided into six subdivisions ( caput, anterior back, buttocks back, front limbs, hind limbs and venters ; Fig. 1 ) based on the anatomical part and the length of the pelt ( Sokolov & A ; Chernova, 1987 ) . Each subdivision was sealed in a plastic bag and frozen at -20A°C. In the research lab, the hair was cut with scissors and all keds and other arthropod ectozoans were collected by manus. The country of the tegument was determined by puting a metal grid ( 20 A- 20 millimeter ) on a skin subdivision and ciphering the figure of squares covering the subdivision.
The cervid keds were divided into classs as follows: late attached winged, late attached wingless, blood-consumed ( expanded venters incorporating blood ) , mating braces ( ever blood-consumed 1s ) and pupae. Other arthropod parasites were besides determined. A random subsample ( n = 200 unless the entire number/section was less ) of all subdivisions was divided by sex. Both sexes were separately weighed and the average weights of the cervid keds of both sexes were calculated. Based on the sex ratio and the average weights of the male and female keds the entire Numberss of flies of both sexes on each tegument subdivision were estimated.
Comparisons of the entire Numberss of cervid keds and their densenesss per tegument country harmonizing to the age, sex and different anatomical parts of the elk were performed with the one-way analysis of discrepancy ( ANOVA ) and the Duncan ‘s station hoc trial with the SPSS-program ( v. 15.0, SPSS Inc. , Chicago, IL, USA ) . For nonparametric informations, the Kruskal-Wallis ANOVA on ranks followed by the Dunn ‘s station hoc trial was performed with the SigmaPlot-program ( v. 11.0, Systat Software Inc. , San Jose, CA, USA ) . Correlations were calculated with the Spearman Correlation Coefficient ( R ) . A p-value & lt ; 0.05 was considered statistically important. The consequences are presented as the mean A± S.E.
The burden of cervid keds on the elk was high and varied depending on the sex and adulthood of the host. The bulls had on the mean 10616 A± 1375 cervid keds, the cattles 3549 A± 587 keds and the calves 1730 A± 191 keds, all these values being significantly different from each other ( Table 1 ) . The maximal Numberss of cervid keds on the bulls, cattles and calves were 17491, 5130 and 2309, severally ( Table 2 ) . The bulls had more copulating cervid ked braces and pupae than the cattles and calves. Other ectozoans were scarce with merely few ticks ( Acari ) nowadays.
The bulls had more keds per tegument country than the cattles and calves ( Table 3 ) . The denseness of cervid keds was the highest on anterior back, where about half of all keds were found ( Table 4, Fig. 1 ) . This was followed by posterior back with about one 4th of all keds. The denseness on other parts decreased harmonizing to the sequence: front limbs & gt ; venters & gt ; caput & gt ; hind limbs. The Numberss of late attached winged and wingless cervid keds and their densenesss correlated negatively with the day of the month of trying ( rs = -0.712- -0.936, P & lt ; 0.01 ) , while the figure of mating braces on the bulls correlated positively with the trying day of the month ( rs = 0.795, P & lt ; 0.05 ) . The thickness of gluteal fat did non correlate with the Numberss or densenesss of cervid keds.
The sex ratio of the cervid keds was equal ( 1:1 ) in all anatomical parts. Weights of late attached winged and wingless keds did non differ significantly but blood-consumed keds were heavier ( Table 5 ) . After cervid keds had consumed blood, males were heavier than females. The average weight of pupae was 9.996 A± 0.391 milligram. The weights of the cervid keds did non differ harmonizing to the age, sex or anatomical part of the elk.
The entire Numberss and densenesss of cervid keds on the elk were highly high. The examined elk had 1.5-2.1 times more cervid keds than reported for the elk in Soviet Belarus ( Ivanov, 1981 ) , 9-35 times more than on the elk in the Leningrad part of the former Soviet Union ( Popov, 1965 ) and 115-708 times more than on the ruddy cervid in Denmark ( Haarlov, 1964 ) . The cervid keds were much more legion than reported antecedently for other louse fly species on homeothermal animate beings. The maximal figure of sheep keds was 300-400 keds per sheep ( Ovis Ariess ) ( Legg et al. , 1991 ) , while the figure of louse flies on birds seems to be smaller as, for illustration, the maximal figure of Crataerina pallida on the European Swift ( Apus Apus ) was 31 flies ( Hutson, 1981 ) . Some homeothermal animate beings seem to be able to defy hippoboscid parasitism. After experimental infections of 50-1000 parasites followed by a period of parasite reproduction, sheep seemed to be able to develop partial opposition and the figure of sheep keds declined to & lt ; 50 keds per animate being ( Nelson, 1962 ) . In a similar mode, after experimental infections of cervid keds on caribou, merely 1-18 of the original 300 keds per caribou survived ( Kynkaanniemi et al. , unpublished information ) . As the elk of the present survey were so to a great extent parasitized, any similar opposition would be improbable.
There are no studies on other louse fly species on the elk that would be every bit legion as the cervid keds in the present survey, but in North America winter ticks ( Dermacentor albipictus ) , a one-host parasite overwintering on wild deers ( Mooring & A ; Samuel, 1999 ) , were reported to be present at the figure of 30000 ticks or more per elk ( Samuel & A ; Welch, 1991 ) . This parasite caused weight loss, anemia and perchance secondary bacterial infections for confined elk ( Glines & A ; Samuel, 1989 ) . Parasitism of winter ticks caused nuisance and moose tried to take ticks by preparing, which increased hair loss ( Mooring & A ; Samuel, 1999 ) . Winter ticks preferred shoulders, cervix and withers similar to the cervid keds in our survey and hair loss began and was the most terrible on these parts. The preparing rate and hair loss were connected to the developmental phase of ticks ; grownup ticks irritated moose more than nymphal phases ( McLaughlin & A ; Addison, 1986 ) . Hair loss remained at a low degree until February, in March it increased aggressively and peaked in April and May ( Mooring & A ; Samuel, 1999 ) . In our survey, merely one elk had minor hair loss ( about 5 dm2 ) , which suggests that cervid ked parasitism at the ascertained degree would non do alopecia for elk, but more surveies in winter and spring will be needed to corroborate this. Furthermore, the burden of cervid keds was non reciprocally associated with the estimation of organic structure adiposeness unlike observed antecedently in winter tick-infested moose enduring from hair loss ( McLaughlin & A ; Addison, 1986 ) .
The average denseness of cowss seize with teething lice ( Bovicola bovis ) on tips ( Bos Sanchez ) was 26.5 lice per tegument dm2 ( informations calculated from Watson et al. , 1997 ) , which was less than the average cervid ked denseness on the bulls in the present survey. The highest B. bovis densenesss on tips were on canvass and shoulder parts ( 88.6 and 79.5 per tegument dm2 ) , which were higher than the average cervid ked densenesss on anterior dorsum of the elk in the present survey. Arthropod ectozoans could be found in 44 % of domestic Canis familiariss ( Canis familiaris ) ( Aldemir, 2007 ) and 82 % of domestic cats ( Felis catus ) were parasitized by Ctenocephalides Felis with a mean of 16 parasites per cat when utilizing the random trying method of minor countries ( Hsu et al. , 2002 ) . Over 45 % of all fleas were located on caput and cervix parts, the following preferable part was dorsal with over 27 % of the fleas. The sheep ked preferable rib and shoulder parts ( Legg et al. , 1991 ) and the sheep with an unfastened, long and reasonably oily fleece were considered ideal hosts for the sheep ked ( Evans, 1950 ) . The cervid ked shows negative geotaxis and phototaxis ( Ivanov, 1981 ) , which may explicate why most of the keds were attached to the withers of the elk. The pelt is the longest in the withers, up to 200 millimeter ( Sokolov & A ; Chernova, 1987 ) , which could supply the best shelter for keds.
Harmonizing to Kadulski ( 1974 ) , the grade of cervid ked parasitism correlated positively to the organic structure size of the host. In the present survey, the grownup elk had more keds than the calves, which fits the hypothesis, but the bulls had more keds than the cattles, despite of no important differences in the skin country and of the somewhat lower organic structure mass. Based on the natural history of the cervid ked, the elk with big place scopes and high degrees of activity could hold the highest hazard to go parasitized. In Finland, 64 % of all bulls move more than 15 kilometer between seasonal place scopes, while merely 30 % of cattles travel as far ( Heikkinen, 2000 ) . The moose travel several times between summer and winter place ranges before settling in their winter home grounds. Therefore the bulls could hold de facto larger place scopes in fall, which may partially explicate why they had higher Numberss of cervid keds on their teguments. In the present survey, the cattles had higher Numberss of keds than the calves, but the densenesss of cervid keds between these two groups were equal. Ivanov ( 1975 ) noticed that if two possible hosts of different size were in the locality, the cervid keds chose the bigger 1. Based on this, calves that still follow their dikes may pull less keds.
The negative correlativity between the entire Numberss and densenesss of late attached cervid keds and the sampling day of the month were logical when reckoned with the flight season of the cervid ked. The Numberss of late attached winged and wingless keds were minor on the elk teguments, as about all keds had consumed blood by the clip of the sampling. Recently attached wingless cervid keds were somewhat heavier than late attached winged keds bespeaking that wingless keds had already consumed blood, although they did non yet show the distended venters characteristic to the bulk of keds. In the Leningrad part of the former Soviet Union the average weight of the freshly hatched cervid keds was 8-9 milligram, and a weight loss of 3.0-3.5 milligram was considered fatal ( Popov, 1965 ) . In the present survey, late attached cervid keds were over 50 % lighter than cervid keds in the Leningrad part. This indicates that there could be important fluctuation in the weight of cervid keds between different geographical parts. This has to be confirmed in the hereafter as it is non impossible that some vaporisation could hold occurred during the storage of the keds.
Based on our consequences, we can preliminarily gauge the population size of cervid keds in Finland. The average figure of cervid keds was 5400 keds per elk. Based on the equal sex ratio ( present survey ) and the generative result of the cervid ked ( Popov, 1965 ; Ivanov, 1981 ) , these 5400 cervid keds could hypothetically bring forth 54000-86400 pupae yearly. Harmonizing to the Finnish Game and Fisheries Research Institute ( 2009b ) , the size of the moose population in Finland was 79000-93000 persons after the hunting season in 2006. These elks could host 400-500 million cervid keds that could bring forth 4-8 billion pupae. Ivanov ( 1981 ) estimated that in 1972 the size of the cervid ked population in Belarus was 36 million keds, much smaller than the estimated population size in Finland. The figure of cervid keds correlated positively with the population size of the elk, while the Numberss of roe cervid and ruddy cervid did non impact the size of the cervid ked population ( Ivanov, 1981 ) . In Eastern Finland the sizes of the populations of other deers are notably smaller than that of the moose population ( Juha Kuittinen, personal communicating ) . This together with consequences of Ivanov ( 1981 ) suggest that the elk could be the chief host of the cervid ked. Based on this, cut downing the figure of elk may be the most practical manner to command the size of the cervid ked population.
1 ) The elk were to a great extent parasitized by the cervid keds. The calves had less keds than the grownups perchance due to the inclination of cervid keds to take a host of a larger organic structure size, and the bulls had more keds than the cattles, which could be caused by their more active travel between seasonal place scopes. 2 ) The denseness of cervid keds varied between the different anatomical parts being the highest on anterior back, where half of all keds were located. This could be explained e.g. by the long hair on the part of the withers or by the negative geotaxis and phototaxis displayed by cervid keds. The degree of parasitism was so high that the effects of the cervid ked on the wellness of deers should be determined.