Prevalence of ectoparasites in goats (Capra aegagrus hircus) brought for slaughter in the Gwagwalada area, Abuja, Nigeria

Ectoparasites from the bodies of 600 slaughtered goats between the ages of 1 and 50 months, collected within a period of 10 months (February to November 2005), were isolated using fractionation techniques. Collected data were analysed by χ²‐test. A total of 60 of the 600 goats examined (10.00%) were infected with ectoparasites. The prevalent ectoparasites observed were species of Rhipicephalus (28 goats; 4.67%), Boophilus (8 goats; 1.33%), Amblyomma (6 goats; 1.00%), Hyalomma (6 goats; 1.00%), Linognathus (18 goats; 3.00%) and Damalina (1 goat; 0.17%). The infection rates according to age and sex of goats were not significantly different (P > 0.05). Goats were observed to have single or multiple infections of ectoparasites. More goats (46) had a single infection than a multiple infection (14). Regarding distribution of the ectoparasites on the various sites of the animal body, it was observed that lice preferred the head, back, ears and belly, while ticks were found in the hooves and interdigital spaces as well as at the sites listed for lice.     

Ectoparasites of northern fulmars Fulmarus glacialis (Procellariiformes: Procellariidae) from the Canadian Arctic

We studied the prevalence and intensity of infestation of ectoparasites on northern fulmars (Fulmarus glacialis L.) from a breeding colony in Arctic Canada in June–August 2003. No fleas or ticks were found on any fulmars, but three species of chewing lice (Phthiraptera) were recorded: Ischnocera: Perineus nigrolimbatus (Giebel 1874), Ischnocera: Saemundssonia occidentalis (Kellogg 1896), and Amblycera: Ancistrona vagelli (Fabricius 1787). Non-breeding birds had a higher prevalence of lice than breeding birds, and prevalence varied markedly among louse species. Our study is an important baseline for the occurrence of ectoparasites on northern fulmars in the high Arctic, a region undergoing extensive environmental change due to global warming, and an area where parasites are expected to extend ranges or increase in prevalence under changing annual temperature regimes.     

Determinants of ectoparasite assemblage structure on rodent hosts from South American marshlands: the effect of host species,locality and season

The relative effects of host species identity, locality and season on ectoparasite assemblages (relative abundances and species richness) harboured by four cricetid rodent hosts (Akodon azarae, Oligoryzomys flavescens, Oxymycterus rufus and Scapteromys aquaticus) were assessed across six closely located sites in Buenos Aires province, Argentina. Relative abundances of ectoparasites (14 species including gamasid mites, an ixodid tick, a trombiculid mite, lice and fleas), as well as total ectoparasite abundance and species richness, were determined mainly by host species and to a lesser extent by locality (despite the small spatial scale of the study), whereas seasonal effect was weak, albeit significant. The abundances of some ectoparasites were determined solely by host, whereas those of other ectoparasites (sometimes belonging to the same higher taxon) were also affected by locality and/or season. In gamasids, there was a significant effect of locality for some species, but not for others. In fleas and lice, the effect of locality was similar in different species, suggesting that this effect is related to the characteristic life history strategy.     

Ectoparasites of gray squirrels in two different habitats and screening of selected ectoparasites for bartonellae

Gray squirrels, Sciurus carolinensis, were livetrapped in 2 different habitat types, woodland (67 squirrels) and parkland (53 squirrels), in southeastern Georgia. Ectoparasites were recovered from anesthetized squirrels and compared between hosts from the 2 habitats. Because of the absence of low vegetation in parkland habitats, it was hypothesized that the ectoparasite fauna, especially ticks and chiggers, would be more diverse on woodland squirrels. The results were generally in agreement with this hypothesis. Seventeen species of ectoparasites were recovered from woodland squirrels, compared with 6 species from parkland squirrels. Five species of ticks and 3 species of chiggers parasitized the woodland squirrels compared with no ticks or chiggers on the parkland squirrels. Significantly higher infestation prevalences were recorded on woodland compared with parkland squirrels for the flea Orchopeas howardi, the tick Amblyomma americanum, and the mesostigmatid mite Androlaelaps fahrenholzi. The mean intensity for O. howardi also was significantly higher on woodland than on parkland squirrels. Because a new strain of Bartonella sp. was isolated recently from S. carolinensis in Georgia, selected ectoparasites from this study were screened for bartonellae by polymerase chain reaction (PCR). Some of the fleas and lice, but none of the mites tested, were PCR positive, suggesting that fleas, or lice, or both, might be vectors of bartonellae between squirrels. Six distinct strains of Bartonella sp. were detected, 2 in fleas and 4 in lice.     

Record of five new endemic small mammal hosts for four ectoparasite species from Sri Lanka

Both ticks and fleas are medically important arthropods, which act as vectors for a number of zoonotic diseases. In Sri Lanka, 35 species of ticks and 20 species of fleas have been reported from mammals, birds and reptiles. However, records of these parasites from small mammals are scanty. Here, we report ticks and fleas infesting muroid rodents and shrews including endemic species collected from around the country. Of the 149 small mammals collected, 48 individuals were infested with ticks, fleas or both. Immature stages of three tick species, Ixodes ceylonensis, Haemaphysalis hystricis and Hyalomma brevipunctata and unidentified species of Rhipicephalus were recorded from ten small mammal species. Four species of fleas, Stivalius phoberus, Stivalius aporus, Lentistivalius ferinus and Nosopsyllus tamilanus were identified from five species of small mammals. We report new host species for I. ceylonensis (Crocidura miya, Suncus montanus and Solisorex pearsoni), S. aporus (S. montanus and Mus fernandoni), N. tamilanus (S. pearsoni) and Rhipicephalus spp. (Crocidura hikmiya, S. montanus, S. pearsoni, M. fernandoni). All new host species reported here are endangered or critically endangered endemics to Sri Lanka.     

Host-ectoparasite associations; the role of host traits,season and habitat on parasitism interactions of the rodents of northeastern Iran

Rodents play a significant role as reservoirs of zoonotic diseases. Nevertheless, in general their ectoparasite assemblage and host-ectoparasite associations are poorly known. This study intended to provide new insights into the relationships between ectoparasites and rodents in northeastern Iran. Rodents were captured using live traps during the years 2016–2020, and their ectoparasites were collected. Parasitological indices such as infestation rate, prevalence and mean intensity of infestation were analyzed. A total of 284 rodents, belonging to 17 species, were trapped and found to be infested by 178 ectoparasites from five orders Siphonaptera, Phthiraptera, Ixodida, Mesostigmata and Trombidiformes. The overall infestation rate was 50.3%. The flea Nosopsyllus fasciatus and the louse Polyplax asiatica dominated among all fleas and lice, respectively. Haemaphysalis punctata and Haemolaelaps sp. were recorded as the most abundant tick and mite, respectively. Nosopsyllus fasciatus exhibited low and Polyplax asiatica moderate host specificity. Approximately 64.2% of ectoparasites shared more than one host, and others were singletons. Seasonal fluctuations were found in the occurrence of ectoparasite; fleas and lice were more abundant in spring and winter, respectively. Ticks demonstrated high abundance in spring and summer and mites were more common in autumn. The overall prevalence of ectoparasite on male rodents was greater than that on females (56.4% vs. 44.4%), while similar mean intensities were detected for both sexes. This study extends the knowledge on the distribution, seasonality and host choice of four main groups of ectoparasites in association with rodents. Further studies are needed to provide deep insight into how relationships and interactions between ectoparasite and rodents are formed, and how they can be applied in epidemiology.     

Neopsylla musseri n. sp. (Siphonaptera-Ctenophthalmidiae), a new flea from central Sulawesi (Indonesia)]

This flea is a montane flea, known from elevations of 1,700-2,300 m, that parasitizes endemic murid rodents. It represents a newly recorded genus and family of fleas for Sulawesi. Morphologically, the closest taxon to N. musseri n. sp. is N. sondaica Jordan, 1931 from Java; it is not closely related to N. luma Traub, 1954 from Borneo.     

云南省吸虱昆虫区系及物种多样性

吸虱是寄生于真兽类哺乳动物体表的专性吸血寄生虫,广布于世界各地。云南省已知吸虱昆虫9科13属44种,分别占中国已知吸虱科、属、种的81.82%,59.09%,45.83%。文章参考大量相关文献,从分类阶元、特有物种、动物地理区划和宿主动物4个方面分析云南省吸虱的物种多样性。云南省吸虱特有种有13种,占云南省已知吸虱种类的29.55%,27种为东洋种,15种为古北和东洋两界兼有种,广布种9种。吸虱在5个地理小区的分布,以横断山中部和横断山南部2个地理小区的吸虱物种多样性较高,其它3个区的物种多样性较低。相对于全国而言,云南省吸虱物种多样性较高,吸虱的宿主动物种类丰富。但蚤、恙螨和革螨等其它体表寄生虫相比较,兽类宿主动物体表吸虱的物种多样性明显低于其它体表寄生虫,1科(属)阶元的吸虱其宿主多为相对一致的1个科(属)动物阶元,反映了吸虱宿主特异性较高的事实,吸虱昆虫与其对应的宿主动物已经形成了比较稳定的"一对一"的寄生关系,这是吸虱昆虫与其宿主动物协同进化的生态学表现。     

Ticks (Acari: Ixodidae) within various phytophysiognomies of a Cerrado reserve in Uberlandia, Minas Gerais, Brazil

Cerrado biome, the South American savannah, covers about 2 million km² and is very rich in endemic species but threatened by agriculture. In this report free-living tick species are presented, and their seasonal and relative distribution within the various phytophysiognomies in a small Cerrado reserve in Minas Gerais State, Brazil. Overall 2,694 free-living ticks were found during a 2 years sampling period with CO₂ traps and cloth dragging. Of these, 73.5% were Amblyomma cajennense and 0.6% Amblyomma dubitatum. All other ticks (25.9%) were retained as Amblyomma spp. Adults of A. cajennense peaked in spring, the nymphs in winter of both years. Amblyomma larval clusters were found in autumn and winter. Adult ticks (46.7%) and nymphs (39.5%) were most often found in woodlands, whereas most larval clusters were found in valley-side marshes (39%). Amblyomma cajennense, Anocentor nitens, Rhipicephalus (Boophilus) microplusand Rhipicephalus sanguineus ticks were found on domestic animals from neighboring properties. Search for Rickettsia in the hemolymph of 497 A. cajennense and one A. dubitatum ticks yielded negative results. Results confirmed earlier reports on the overwhelming prevalence of A. cajennense ticks in the Cerrado biome of Brazil and added information to habitat preferences of this tick species, a major vector in Brazil of the Rocky Mountain spotted fever.     

Ectoparasites of commensal rodents in Sulawesi Utara, Indonesia, with notes on species of medical importance

Ectoparasite records are presented for four species of commensal murid rodents (Rattus rattus palelae Miller & Hollister, R. argentiventer (Robinson & Kloss), R. exulans (Peale) and Mus musculus castaneus Waterhouse) in Sulawesi Utara, with particular reference to the potential for these arthropods to bite and transmit pathogens to humans. The flea, Xenopsylla cheopis (Rothschild), was most common on R.r. palelae and is capable of transmitting plague and other pathogens to humans although no current foci for these diseases are known in Sulawesi. Hoplopleura pacifica Ewing and Polyplax spinulosa (Burmeister) sucking lice parasitized all three Rattus species although H. pacifica was mainly associated with R. exulans and P. spinulosa with R.r. palelae. These lice do not bite humans but may be intramurid vectors of murine typhus and other zoonoses. The mites Laelaps echidnina Berlese and L. nuttalli Hirst were both collected; the latter was recorded from all four murid species, mainly R. exulans. The mite Ornithonyssus bacoti Hirst was rare. Only one chigger mite, Walchiella oudemansi (Walch), was retrieved from murids (from R. exulans) and a single Leptotrombidium deliense (Walch) chigger was taken from a human subject. Although L. deliense is a significant vector of scrub typhus, a disease known from Sulawesi, the L. deliense-R. argentiventer relationship frequently noted in the ecology of this rickettsial disease, was not evident in this survey. Other ectoparasites collected from murids were the ticks, Ixodes granulatus Supino (first record for Sulawesi), Haemaphysalis sp. and Dermacentor sp., the mites Myocoptes musculinus (Koch) and Listrophoroides cucullatus (Trouessart), acarids and a uropodid.(ABSTRACT TRUNCATED AT 250 WORDS)     

Seasonal studies on commensal rats and their ectoparasites in a rural area of Egypt: the relationship of ectoparasites to the species, locality, and relative abundance of the host.

The present study was carried out in 3 villages, namely Kafr Ayoub Soliman, Kafr Ibrahim El-Aidi, and El-Sa'adat, Sharqiya Governorate, Egypt. A total of 519 rats was collected from the 3 study sites: 46.6% Rattus rattus, and 53.4% Rattus norvegicus. A total of 20,643 ectoparasites was recovered from R. rattus: 33.3% mites, 33.8% fleas, and 32.9% lice. From R. norvegicus a total of 40,997 ectoparasites was recovered: 28.9% mites, 31% fleas, and 40.1% lice. Three common mite species were recovered from both rat hosts, i.e., Ornithonyssus bacoti, Radfordia ensifera, and Laelaps nuttalli. Three common flea species were also recovered from both rat hosts, i.e., Echidnophaga gallinacea, Leptopsylla segnis, and Xenopsylla cheopis. Polyplax spinulosa was the only dominant louse species that infested both rat hosts. Rats did not show a definite breeding season, and the seasonal rat indices were generally low in different study sites. There were no significant differences between the prevalence of each of mites, fleas, and lice in both rat species. The total general indices of mites and fleas, on the other hand, was significantly higher in R. norvegicus. The general index of X. cheopis was high and ranged between 5.9 in R. rattus and 14.5 in R. norvegicus. Season-related changes were observed in the general index of each of L. segnis infesting both rat species and R. ensifera and O. bacoti infesting R. norvegicus. The prevalence and general indices of some ectoparasites showed differences related to the locality of their rat hosts. Seasonal changes in the general indices of some ectoparasites paralleled seasonal changes in the relative abundance of their rat hosts.     

Analysis of ectoparasites (chigger mites,gamasid mites,fleas and sucking lice) of the Yunnan red‐backed vole (Eothenomys miletus) sampled throughout its range in southwest China

The Yunnan red‐backed vole Eothenomys miletus (Rodentia: Cricetidae) is an endemic rodent species and reservoir host of zoonoses in southwest China. Based on a large host sample (2463 voles collected from 39 localities between 2001 and 2013), a general analysis of four categories of ectoparasite (fleas, sucking lice, chigger mites and gamasid mites) on E. miletus across its entire range of distribution was made. This analysis identified a total of 71 895 ectoparasites belonging to 320 species (30 species of flea, 9 of sucking louse, 106 of gamasid mite and 175 of chigger mite) with a high prevalence (87%), mean abundance (29.19) and mean intensity (33.69). Of the 18 vector species of zoonoses found on E. miletus, the flea Ctenophthalmus quadratus (Siphonaptera: Hystrichopsyllidae) and chigger mite Leptotrombidium scutellare (Trombidiformes: Trombiculidae) were the dominant species; these are the main vectors of zoonoses in China. All of the dominant parasite species showed an aggregated distribution pattern. Male voles harboured more species of parasite than females. Chigger mites represented the most abundant species group on voles and their prevalence was positively correlated with mean abundance (r = 0.73; P < 0.05). As a single rodent species, E. miletus has a high potential to harbour abundant ectoparasites with high species diversity and high rates of infestation. The sex of the vole affects ectoparasite infestation.     

Tick infestation on birds in an urban Atlantic Forest fragment in north-eastern Brazil

Birds are important hosts for various tick species, playing a significant role in their biological life cycle and dispersion. In this study, we investigated tick infestations on birds trapped in an urban remnant of Atlantic Forest in Pernambuco state, Brazil. From February 2015 to March 2017, 541 birds belonging to 52 species were trapped with mist nets and examined for ectoparasites. Birds trapped in the late successional forest were significantly more infested than birds trapped in the early successional forest. In the same way, ectoparasite infestation varied significantly according to bird weight and collection plot. Overall, 198 birds (36.6%) belonging to 27 species were parasitized by ectoparasites (i.e., ticks, lice and/or mites). Ectoparasites were effectively collected from 111 birds, of which 99 belonging to 20 species were infested by ticks (n?=?261), namely, Amblyomma longirostre (13 nymphs), Amblyomma nodosum (21 nymphs), Amblyomma varium (one nymph), and Amblyomma spp. (five nymphs and 221 larvae). Most of the ticks (>?90%) were collected from Passeriformes. This study provides the second record of A. varium in Pernambuco state and confirms that birds, especially Passeriformes, are important hosts for larvae and nymphs of Amblyomma spp. in the Atlantic Forest biome of Pernambuco.

     

Detection of Rickettsia spp. and host and habitat associations of fleas (Siphonaptera) in eastern Taiwan

Rickettsia typhi and Rickettsia felis (Rickettsiales: Rickettsiaceae) are two rickettsiae principally transmitted by fleas, but the detection of either pathogen has rarely been attempted in Taiwan. Of 2048 small mammals trapped in eastern Taiwan, Apodemus agrarius Pallas (24.5%) and Mus caroli Bonhote (24.4%) (both: Rodentia: Muridae) were the most abundant, and M. caroli hosted the highest proportion of fleas (63.9% of 330 fleas). Two flea species were identified: Stivalius aporus Jordan and Rothschild (Siphonaptera: Stivaliidae), and Acropsylla episema Rothschild (Siphonaptera: Leptopsyllidae). Nested polymerase chain reaction targeting parts of the ompB and gltA genes showed six fleas to be positive for Rickettsia spp. (3.8% of 160 samples), which showed the greatest similarity to R. felis, Rickettsia japonica, Rickettsia conorii or Rickettsia sp. TwKM01. Rickettsia typhi was not detected in the fleas and Rickettsia co-infection did not occur. Both flea species were more abundant during months with lower temperatures and less rainfall, and flea abundance on M. caroli was not related to soil hardness, vegetative height, ground cover by litter or by understory layer, or the abundance of M. caroli. Our study reveals the potential circulation of R. felis and other rickettsiae in eastern Taiwan, necessitating further surveillance of rickettsial diseases in this region. This is especially important because many novel rickettsioses are emerging worldwide.     

Diversity and Coexistence of Ectoparasites in Small Rodents in a Tropical Dry Forest

To understand the mechanisms driving species diversity is central to community ecology. Here, we explored if habitat partitioning is associated with a species‐rich ectoparasite community in small rodents from a tropical dry forest in western Mexico. We trapped 199 mice in three 0.5 ha‐plots from eight small rodent species for every two months, from July 2011 to April 2012, and collected their ectoparasites. We identified 17 species of mites, two sucking lice species, two phoretic species, and one commensal species. The most abundant ectoparasite species was Steptolaelaps liomydis, representing 42 percent of all ectoparasites collected; seven ectoparasite species had < 10 individuals. Eighteen ectoparasite species (of 22 species) were collected from the most abundant rodent Liomys pictus. C‐score and the number of checkerboard species pairs were significantly higher against a random expectation. Ectoparasite species in L. pictus mice showed host microhabitat partitioning; Fahrenholzia ehrlichi and Fahrenholzia texana were found only in the anterior dorsal area, Ornithonysus sp. occurred along the dorsal part, Ixodes species were restricted to the ears, and Steptolaelaps liomydis was found throughout the body. We also identified ectoparasite communities with distinct species composition in two rodent species that use contrasting macrohabitats (L. pictus, strictly terrestrial; Peromyscus perfulvus, mostly arboreal). The remaining and low abundant rodent species showed a species‐poor ectoparasite community composition. We conclude that habitat partitioning at both macro and microhabitat scales appeared to characterize the species‐rich ectoparasite community. Conversely, most rodent host species with low abundances showed a species‐poor ectoparasite community.     

The Argasidae, Ixodidae and Nuttalliellidae (Acari: Ixodida): A World List of Valid Tick Names

The world's argasid tick fauna comprises 183 species in four genera, namely Argas, Carios, Ornithodoros and Otobius in the family Argasidae. The ixodid tick fauna consists of 241 species in the genus Ixodes and 442 species in the genera Amblyomma, Anomalohimalaya, Bothriocroton, Cosmiomma, Dermacentor, Haemaphysalis, Hyalomma, Margaropus, Nosomma, Rhipicentor and Rhipicephalus in the family Ixodidae, with the genus Boophilus becoming a subgenus of the genus Rhipicephalus. The family Nuttalliellidae is represented by the monospecific genus Nuttalliella. The species names of these ticks, based on seven previous complete or partial listings, as well as those of recently described new species, are presented in tabular format. This revised version was published online in July 2006 with corrections to the Cover Date.     

Presence of Bartonella and Rickettsia spp. in cat fleas and brown dog ticks collected from dogs in American Samoa

Surveys for ectoparasites of dogs on the American Samoa islands of Aunu'u and Tutuila were conducted in June 2012, and was followed by molecular screening of samples. One species of flea, Ctenocephalides felis, and one tick species, Rhipicephalus sanguineus, were collected and tested for Bartonella and Rickettsia species via PCR. Bartonella clarridgeiae and an unnamed spotted fever group Rickettsia were detected in the fleas; the Rickettsia species was previously reported from Ctenocephalides spp. from Egypt, Thailand, USA, and the Republic of the Marshall Islands. None of the ticks were positive for Bartonella or Rickettsia species. This is the first report of flea-borne Rickettsia and B. clarridgeiae, considered an emerging human pathogen in New Zealand, from American Samoa. Ectoparasite-borne infections are easily misdiagnosed or ignored as their symptoms are often vague and similar to other illnesses (e.g., measles, dengue). Our results indicate a potential threat to human and animal health as infected fleas were collected from household pets.     

Seasonal patterns of rodents, fleas and plague status in the Western Usambara Mountains, Tanzania

Field and commensal rodents were live-trapped at three villages in an active focus of plague (Yersinia pseudotuberculosis pestis) in Lushoto District, Western Usambara Mountains, Tanga Region, Tanzania, from December 1983 to November 1984. Their flea ectoparasites were collected, identified and counted. The rodent carcasses were serologically examined for specific plague antibodies and antigens, and bacteriologically examined for bipolar staining bacilli. A total of 1758 traps were set during the 12-month period and 924 animals were caught. From these, 1037 fleas were collected. Rattus rattus (L.), Praomys natalensis (Smith) and Lophuromys flavopunctatus Thomas comprised the largest proportions of the rodent population, while Dinopsyllus lypusus Jordan & Rothschild, Ctenophthalmus calceatus Waterston and Xenopsylla brasiliensis (Baker) were the dominant flea species. Rodents were most abundantly trapped during December and January. Flea indices were highest from December to May. Human plague was most active from November to March. Rodents contained plague antibodies every month except May and July, with a peak in September. Plague antigens and bipolar bacilli were detected in rodent organs during January-April. From the product of abundance and infection rate, the most prevalent rodent hosts of plague appeared to be R. rattus, Otomys angoniensis Wroughton, P. natalensis and Pelomys fallax (Peters). Continuous integrated control of rodents and fleas was recommended, reinforced by quarantine and maintenance of a surveillance service for clinical detection, diagnosis and treatment of patients in the plague endemic area.     

Influence of bill and foot morphology on the ectoparasites of barn owls

Preening is the principle behavioral defense used by birds to combat ectoparasites. Most birds have a small overhang at the tip of their bills that is used to shear through the tough cuticle of ectoparasitic arthropods, making preening much more efficient. Birds may also scratch with their feet to defend against ectoparasites. This is particularly important for removing ectoparasites on the head, which birds cannot preen. Scratching may be enhanced by the comb-like serrations that are found on the claws of birds in many avian families. We examined the prevalence and intensity of ectoparasites of barn owls (Tyto alba pratincola) in southern Idaho in relation to bill hook length and morphological characteristics of the pectinate claw. The barn owls in our study were infested with 3 species of lice (Phthiraptera: Ischnocera): Colpocephalum turbinatum , Kurodaia subpachygaster, and Strigiphilus aitkeni . Bill hook length was associated with the prevalence of these lice. Owls with longer hooks were more likely to be infested with lice. Conventional wisdom suggests that the bill morphology of raptors has been shaped by selection for efficient foraging; our data suggest that hook morphology may also play a role in ectoparasite defense. The number of teeth on the pectinate claw was also associated with the prevalence of lice. Owls that had claws with more teeth were less likely to be infested with lice, which suggests that larger pectinate claws may offer relatively more protection against ectoparasitic lice. Experiments that manipulate the bill hook and pectinate claw are needed to confirm whether these host characters are involved in ectoparasite defense. Finally, we recovered mammalian ectoparasites from 4 barn owls. We recovered species of mammalian lice (Phthiraptera:Anoplura) and fleas (Siphonaptera) that are commonly found on microtine rodents. The owls probably acquired these parasites from recently eaten prey. This represents 1 of the few documented cases of parasites "straggling" from prey to predator.