Maggi et al. Parasites & Vectors (2022) 15:415 https://doi.org/10.1186/s13071-022-05553-8 Parasites & Vectors REVIEW Open Access Vector‑borne and other pathogens of potential relevance disseminated by relocated cats Ricardo Guillermo Maggi1*†, Vicky Halls2†, Friederike Krämer3†, Michael Lappin4†, Maria Grazia Pennisi5†, Andrew S. Peregrine6†, Xavier Roura7†, Bettina Schunack8†, Valeria Scorza4†, Séverine Tasker9,10†, Gad Baneth11†, Patrick Bourdeau12†, Dwight D. Bowman13†, Edward B. Breitschwerdt1†, Gioia Capelli14†, Luís Cardoso15†, Filipe Dantas‑Torres16†, Gerhard Dobler17†, Lluís Ferrer18†, Luigi Gradoni19†, Peter Irwin20†, Frans Jongejan21†, Volkhard A. J. Kempf22†, Barbara Kohn23†, Susan Little24†, Maxime Madder25†, Carla Maia26†, Mary Marcondes27†, Guadalupe Miró28†, Torsten Naucke29†, Gaetano Oliva30†, Domenico Otranto31†, Barend L. Penzhorn21†, Martin Pfeffer32†, Ángel Sainz28†, SungShik Shin33†, Laia Solano‑Gallego18†, Reinhard K. Straubinger34†, Rebecca Traub35† and Ian Wright36† Abstract Large populations of unowned cats constitute an animal welfare, ecological, societal and public health issue world‑ wide. Their relocation and homing are currently carried out in many parts of the world with the intention of relieving suffering and social problems, while contributing to ethical and humane population control in these cat populations. An understanding of an individual cat’s lifestyle and disease status by veterinary team professionals and those work‑ ing with cat charities can help to prevent severe cat stress and the spread of feline pathogens, especially vector-borne pathogens, which can be overlooked in cats. In this article, we discuss the issue of relocation and homing of unowned cats from a global perspective. We also review zoonotic and non-zoonotic infectious agents of cats and give a list of practical recommendations for veterinary team professionals dealing with homing cats. Finally, we present a consen‑ sus statement consolidated at the 15th Symposium of the Companion Vector-Borne Diseases (CVBD) World Forum in 2020, ultimately to help veterinary team professionals understand the problem and the role they have in helping to prevent and manage vector-borne and other pathogens in relocated cats. Keywords: Adoption, Animal welfare, Feline, Homing, Importation, Parasites, Bacteria, Viruses, Prevention, Relocation, Shelter, Zoonosis † Ricardo G. Maggi and all co-authors contributed equally to this work *Correspondence: rgmaggi@ncsu.edu 1 Department of Clinical Sciences, North Carolina State University, Raleigh, NC, USA Full list of author information is available at the end of the article © The Author(s) 2022. 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The Creative Commons Public Domain Dedication waiver (http://​creat​iveco​ mmons.​org/​publi​cdoma​in/​zero/1.​0/) applies to the data made available in this article, unless otherwise stated in a credit line to the data. Maggi et al. Parasites & Vectors (2022) 15:415 Page 2 of 15 Graphical Abstract Background complaints from the community. If strategic, well- At the 15th Symposium of the Companion Vector-Borne planned and executed population management programs Diseases (CVBD) World Forum in 2020, and subse- (i.e. high-volume TNR) existed throughout the world it quently at the International Society of Feline Medicine is likely that many cat relocation decisions would not be (ISFM) Symposium in 2021, the global and public health required. impact of the spread of vector-borne pathogens in relo- The relocation and homing1 of cats are linked to eco- cated cats was discussed. Although this topic has received nomic, cultural and environmental factors. They are attention in dogs [1], it can be overlooked in cats. In this also very much linked to how populations of unowned article, we discuss the issue of relocation and homing of cats are managed [4], which in turn is influenced by the unowned cats from a global perspective. We also review lifestyle of the cats involved [5]. Globally, a spectrum zoonotic and non-zoonotic infectious agents of cats and of cat lifestyle categories exists, i.e. feral, street, inbe- present a list of practical recommendations for veterinary tweener and pet cat (see Fig. 1) [6]. Categorization of a team professionals dealing with homing cats. Finally, we cat’s lifestyle depends on its desire to live with people and present a consensus statement on this topic to help veter- whether it can live independently of people or within a inary team professionals understand the problem and the household. role they have in helping to prevent and manage vector- Unowned cats from all categories can be found in hom- borne and other pathogens in relocated cats. ing centers, where they are housed awaiting adoption or taken into foster care. However, awareness of the needs of Relocation and homing of cats these cats is important for ensuring that homing, includ- Global pet cat populations are tracked by pet product ing relocation, is truly an appropriate outcome for these industries, with the current world pet cat population cats. Homing cats that are unsuitable to be kept as pets exceeding 373 million cats [2]. Providing an equivalent (particularly feral and street cats) can cause severe dis- estimate for unowned cats is more complex, but this tress for these animals. Additionally, the stress involved number has been suggested to be as high as 600 million in travel and relocation of cats, including those that are [3]. Intact free-roaming cats reproduce rapidly, and, in adapted to live in households as pets, is also important to response, homing organizations and individuals look 1 The term homing is used rather than rehoming in recognition that not all for ways to remove these cats from their current situa- cats in need of homing have previously had homes as pets and because the tion with the intention to relieve suffering and resolve appropriate solution for some cats is to provide an alternative lifestyle out- doors for them rather than homing them as a pet. Maggi et al. Parasites & Vectors (2022) 15:415 Page 3 of 15 Fig. 1 Categorization of cat lifestyles [6], which exist as a spectrum (figure with courtesy of International Cat Care) consider; recent data from the UK suggest that increasing non-zoonotic as well as zoonotic, commonly occur in numbers of cats are imported from abroad to be pets [4]. cats. For some feline infections, there are known signifi- Important disease implications also exist, such as impor- cant differences in prevalence levels based on the type of tation of vector-borne pathogens, which are not native to cat. For example, infection of the domestic cat with the the geographical relocation destination. Some of these three main retroviruses, feline leukemia virus (FeLV), infectious agents also have zoonotic potential [7]. feline immunodeficiency virus (FIV) and feline foamy Education of veterinary team professionals, as well as virus, is generally highest in cats allowed outdoors and individuals engaged in relocation activities, in under- in contact with other cats [11–13]. We herein provide in standing the risks involved in inappropriate homing and Table 1 a detailed listing of potentially occurring vector- relocation to different parts of the world and in how borne pathogens and their corresponding vectors in cats. to recognize, diagnose, treat and prevent infections is However, the relevance and relative risk for each patho- important. One charity has outlined a possible approach gen may vary broadly, e.g. depending on geographical to testing imported cats [8]. Additionally, online location of the homed cat. resources (https://​cvbd.​elanco.​com; https://​www.​esc- cap.​org; https://​capcv​et.​org/; https://​www.​trocc​ap.​com/; http://​www.​abcdc​atsve​ts.​org/) and open access papers Non‑zoonotic infectious agents [9, 10], including descriptions of infection manifestations Most viral agents of cats are non-zoonotic (e.g., feline and maps documenting risks in different geographical retroviruses; feline coronaviruses; feline herpesvirus-1 areas, are freely available. [FHV]; feline caliciviruses [FCV]). These agents are easily transmitted by direct contact with other cats, their secre- tions or vertical transmission. Thus, most feline viral Infectious agents to consider in homed cats infections are most common in populations of cats with Many infectious disease agents carried by cats are direct contact. This is particularly true for the respira- zoonotic [7]. Viral, bacterial (including rickettsial), tory tract agents, FHV and FCV [14–17]. These agents mycotic and parasitic agents are most common, with can be carried by cats even if vaccinated, and stressful many of the bacterial and parasitic agents being vector- conditions such as homing can induce repeated shed- borne. A common misconception is that cats are less ding of FHV, potentially leading to infection of additional infested with vectors (such as ticks, mites, fleas) and are cats. Hypervirulent FCVs are one of the most danger- also less frequented by temporary vectors (mosquitoes, ous examples of this problem as these variants can cause sand flies and flies) than other companion animals such fatal infection even in adult cats previously vaccinated as dogs. However, vectors and vector-borne pathogens, for other FCVs [14, 16]. While the feline retroviruses are Table 1 Feline vector-borne pathogens (including modes of transmission, potential clinical and laboratory findings and zoonotic potential) (*suspected in cats; **reported in other host species, but not yet in cats) Genus Species Modes of transmission Clinical and laboratory findings (most common Zoonotic potential findings in case of illness, generally often subclinical infection) Bacteria Anaplasma A. bovis Tick bites Unknown Unknown Maggi et al. Parasites & Vectors A. phagocytophilum Tick bites Fever, thrombocytopenia, suspected suppura‑ Yes tive polyarthritis A. platys Tick bites suspected fever and thrombocytopenia Yes Bartonella B. clarridgeiae Flea feces Fever, lymphadenomegaly, suspected endo‑ Yes Animal bites, scratches carditis (2022) 15:415 B. elizabethae Flea feces Suspected endocarditis Yes Animal bites, scratches B. henselae Flea feces, tick bites** Fever, lymphadenomegaly, endocarditis, Yes animal bites, scratches myocarditis B. koehlerae Flea feces Fever, lymphadenomegaly, suspected endo‑ Yes Animal bites, scratches carditis B. quintana Human body louse bites**, flea bites* Fever, endocarditis Yes Animal bites, scratches B. vinsonii berkhoffii Tick bites** Recurrent osteomyelitis Yes Animal bites* Borrelia B. burgdorferi Tick bites Fever, lethargy, suspected suppurative polyar‑ Yes thritis B. hispanica Tick bites, sucking lice bites Weight loss, regenerative anemia, monocytosis Yes B. miyamotoi Tick bites, sucking lice bites Unknown Yes B. persica Tick bites, sucking lice bites Lethargy, anorexia, anemia, thrombocytopenia, Yes icterus Coxiella C. burnetii Flea bites*, tick bites* Abortion, stillbirth Yes Direct contact*, fomites* Ehrlichia E. canis Tick bites Fever, thrombocytopenia, lymphadenomegly Yes E. chaffeensis Tick bites Few data available Yes E. ewingii Tick bites Few data available Yes Francisella F. tularensis Tick bites**, blood sucking vectors (mechani‑ Fever, lymphadenomegaly, cough, dyspnea Yes cal)** Animal bites*, ingestion**, scratches* Haemotropic Mycoplasma spp. ’Candidatus M. haemominutum’ Flea bites*, tick bites* Fever, hemolytic anemia No Animal bites*, vertical** ’Candidatus M. turicensis’ Flea bites*, tick bites* Fever, hemolytic anemia No Animal bites*, vertical** M. haemofelis Flea bites*, tick bites* Fever, hemolytic anemia Yes Animal bites*, vertical** Page 4 of 15 Table 1 (continued) Genus Species Modes of transmission Clinical and laboratory findings (most common Zoonotic potential findings in case of illness, generally often subclinical infection) Neorickettsia N. mikurensis Tick bites** Unknown; suspected lymphadenomegaly and Yes fever Maggi et al. Parasites & Vectors N. risticii Ingestion of infected trematodes** Unknown; suspected anorexia, lymphadeno‑ Unknown megaly and diarrhea Rickettsia R. conorii Tick bites Suspected fever Yes R. felis Flea bites Unknown; suspected fever Yes R. massiliae Tick bites Suspected fever Yes (2022) 15:415 R. rickettsii Tick bites Suspected fever Yes R. typhi Flea bites Suspected fever Yes Yersinia Y. pestis Flea bites Fever, necrotic stomatitis, lymphadenomegaly, Yes direct contact, ingestion of infected host/prey cough, dyspnea Protozoa Babesia B. canis Tick bites** Mild chronic anemia (few data available) Unknown vertical** B. felis Tick bites* Hemolytic anemia, icterus, increased ALT Unknown B. lengau Tick bites* Anemia (few data available) Unknown B. leo Tick bites* Anemia (few data available) Unknown B. vogeli Tick bites** Unknown No B. panickeri Tick bites* Anorexia, lethargy, hemolytic anemia (few data Unknown available) B. hongkongensis Tick bites* Unknown Unknown B. presentii Tick bites* Anemia, fever, icterus, lethargy (few data avail‑ Unknown able) Cytauxzoon Cytauxzoon felis Tick bites Fever, icterus, splenomegaly, hepatomegaly, Unknown fatal infection Cytauxzoon sp. Tick bites* Sub-clinical to rare severe fatal infection with Unknown anemia and fever Hepatozoon H. canis Ingestion of intermediate hosts (ticks)**, inges‑ Unknown Unknown tion of paratenic hosts**, vertical** H. felis Unknown, vertical* Icterus, pancytopenia, increased SDMA (sym‑ Unknown metric dimethylarginine) H. silvestris Unknown Myocarditis Unknown Page 5 of 15 Table 1 (continued) Genus Species Modes of transmission Clinical and laboratory findings (most common Zoonotic potential findings in case of illness, generally often subclinical infection) Leishmania L. amazonensis Sand fly bites* Skin and mucosal lesions (nodular, ulcerative) Yes L. braziliensis Sand fly bites* Skin and mucosal lesions (nodular, ulcerative) Yes Maggi et al. Parasites & Vectors L. infantum Sand fly bites Skin and mucosal lesions (nodular, ulcerative), Yes Animal bites**, blood transfusion**, vertical** lymphadenopathy, ocular lesions (conjunc‑ tivitis, uveitis), hyperglobulinemia, anemia, proteinuria L. mexicana Sand fly bites* Skin and mucosal lesions (nodular, ulcerative) Yes (2022) 15:415 (L. venezuelensis) Trypanosoma T. brucei (brucei) Tsetse fly bites, tabanids and other biting flies Fever, emaciation, anorexia, edema and Yes (mechanical) erythema (head, face), alopecia, lymphadeno‑ megaly, ocular lesions (keratitis, conjunctivitis, photophobia, epiphora, pannus, hypopyon), eventual lethal outcome T. congolense Tsetse fly bites, tabanids and other biting flies Emaciation, anemia, no ocular signs, fatal Yes (mechanical) outcome T. cruzi Triatomine feces Seizures, transient posterior paralysis (few data Yes available) T. evansi Tabanids and stable flies (mechanical), vampire Fever, weight loss, diarrhea and vomiting, Yes bats bites** edema (head, face), lymphadenomegaly, ocular lesions (corneal opacity), regenerative anemia, icterus, hindlimb instability, muscle atrophy, potential fatal outcome Viruses Feline calicivirus (FCV) Flea bites* Fever, stomatitis, upper respiratory signs, pol‑ No direct contact, fomites yarthritis, vasculitis with systemic variants Dabie bandavirus (formerly: Severe Fever Tick bites Fever, anorexia, lethargy, vomiting, thrombocy‑ Yes with Thrombocytopenia Syndrome Virus Animal bites, direct contact, scratches topenia, leukopenia [SFTSV]) West Nile virus (WNV) Mosquito bites Neurologic signs Yes Helminths Acanthocheilonema A. dracunculoides Flea bites** Unknown Unknown A. reconditum Flea bites**, lice (sucking and chewing) bites/ Unknown Unknown ingestion** Aelurostrongylus A. abstrusus Ingestion of intermediate hosts (slugs and Cough, dyspnea No snails), ingestion of paratenic hosts, ingestion of free larvae in snail/slug slime Page 6 of 15 Maggi et al. Parasites & Vectors Table 1 (continued) Genus Species Modes of transmission Clinical and laboratory findings (most common Zoonotic potential findings in case of illness, generally often (2022) 15:415 subclinical infection) Brugia B. malayi Mosquito bites Lymphadenomegaly, lymphedema (few data Yes available) B. pahangi Mosquito bites Lymphadenomegaly, lymphedema (few data Yes available) Cercopithifilaria C. grassii Tick bites Unknown Unknown Dirofilaria D. immitis Mosquito bites Cough, dyspnea, sudden death Yes D. repens Mosquito bites Subcutaneous nodules, dermatitis Yes Troglostrongylus T. brevior Ingestion of intermediate hosts (slugs and Cough, dyspnea, nasal discharge No snails), ingestion of paratenic hosts, inges‑ tion of free larvae in snail/slug slime, vertical, transmammary T. subcrenatus Unknown Cough, dyspnea, nasal discharge No Thelazia T. californiensis Deposition by secretophagous house flies Blepharospasm, epiphora, conjunctivitis, Yes (Fannia canicularis, Fannia benjamini) keratitis T. callipaeda Deposition by secretophagous fruit flies (Phor- Blepharospasm, epiphora, conjunctivitis, Yes tica variegata, Amiota okadai) keratitis Dipylidium D. caninum Ingestion of intermediate hosts (fleas, lice) Unthrifty, rarely vomiting or diarrhea Yes Page 7 of 15 Maggi et al. Parasites & Vectors (2022) 15:415 Page 8 of 15 non-zoonotic, if they cause immune deficiency, the risk Even though there is little information regarding cats as of shedding of other infectious agents, including zoonotic reservoirs for zoonotic agents, owners and animal han- ones, may increase [18, 19]. As the early stages of retrovi- dlers are at risk of infection with some viruses, bacteria, ral infections can be subclinical, testing for FeLV and FIV fungi and numerous endo- and ectoparasites. However, in all cats being homed is strongly recommended. misinformation is common, e.g. overestimation of the Among deep not zoonotic mycotic infections, Crypto- risk of cat ownership for toxoplasmosis [26]. Veterinary coccus spp. and Aspergillus spp. can cause severe feline team professionals therefore must be well informed diseases, but they are non-contagious [20]. Cats with about zoonotic agents associated with cats and be able cryptococcosis are good sentinel species for environmen- to communicate such information effectively to cat own- tal contamination and the consequent risk of exposure ers, particularly about routes, modes of transmission and for humans and animals. prevention. Transmission of zoonotic pathogens may Feline Mycoplasma spp. commonly colonize mucus occur through feces (either by direct contact or indirectly membranes of cats, and the respiratory agents in this through contaminated soil, water or raw produce); hair group are occasionally associated with clinical disease (e.g., dermatophytes, either directly or indirectly); oral, and are contagious among cats and thus likely more com- eye, respiratory, skin and urogenital secretions/exudates; mon in cats housed together [21–23]. bites and scratches; shared environmental exposure (e.g., The type of cat is also potentially associated with inhalation of spores for systemic fungi like Blastomyces increased prevalence rates for several non-zoonotic dermatitidis); and shared vectors such as fleas, ticks, parasitic agents. For example, cats allowed outdoors are mites, sand flies, mosquitoes or flies (see Table 1). The inclined to hunt and have increased risk of exposure to intensive self-grooming of cats increases the risk for the infectious agents carried by intermediate or paratenic potential spread of pathogens from anal, genital and ocu- hosts. Examples include the protozoan Cystoisospora lar mucous membranes to the mouth of the cat and fur- felis, several cestodes such as Taenia taeniaeformis and ther spread with saliva to the fur [27–29]. the lungworms Aelurostrongylus abstrusus and Troglos- Scratches and bites are common transmission modes trongylus spp. Most other common parasitic agents of for zoonotic pathogens. Indeed, an average of 1% of all cats are zoonotic and are listed below in the correspond- emergency room visits (per year) in the US are to evaluate ing section. Periodic deworming is critical to avoiding people bitten by animals [30], and an estimated 400,000 environmental contamination with eggs and larvae of cat bites and 4.5 million dog bites occur in the US every helminths. Cats exposed to the feces of other cats may year [31]. Other high-income countries such as Australia, also be more likely infected by other non-zoonotic proto- Canada and France have comparable annual incidence zoal agents such as Tritrichomonas blagburni (previously rates for dog bites; worldwide cat bites account for 2–50% referred to as feline isolates of Tritrichomonas foetus). of injuries related to animal bites [31], of which a large Again, other important protozoal agents in this context percentage become infected. The risk of human infection are zoonotic and are listed below. from scratches and bites is increased when the cat owner/ Fleas, mosquitoes, sand flies and ticks are the com- handler is immunocompromised and/or the cat they are mon vectors for feline vector-borne pathogens. However, in contact with is showing clinical signs of a disease [7]. only a few of these are non-zoonotic or possess unknown The risk of zoonotic pathogen transmission should be zoonotic potential, of which the latter is represented by evaluated by the attending veterinarian and discussed the protozoan genera Babesia, Cytauxzoon and Hepato- with the owner, including appropriate deworming, vector zoon (see Table 1). While indoor cats are still at risk, cats control and handling recommendations. with access outdoors are more commonly exposed to Zoonotic agents occur across all relevant patho- vectors in general, emphasizing the importance of vector gen types, i.e. parasites, viruses, fungi and bacteria. control. Regarding parasites, there are a few that need particu- lar zoonotic attention: Toxoplasma gondii is still one of Zoonotic infectious agents the most important zoonotic pathogens. Toxoplasmosis Zoonoses are infections that are naturally transmitted in people is a multisystemic disease that causes granu- between animals and humans, or between humans and lomatous inflammation in several tissues. Especially at animals [24]. Thus, in contrast to the terms ‘anthropo- risk are immunocompromised individuals, where infec- zoonosis’ (transmission from animals to humans) and tion frequently presents with pulmonary disease or dif- ‘zooanthroponosis’ (transmission from humans to ani- fuse encephalitis [32], and seronegative, naïve, pregnant mals) [25], the term zoonosis is non-directional and can women, as intrauterine infection causing congenital refer to either route of transmission [24]. toxoplasmosis may cause abortion, neonatal death or fetal abnormalities with detrimental consequences for Maggi et al. Parasites & Vectors (2022) 15:415 Page 9 of 15 the new-born child [33–36]. Additionally, infection by contamination with triatomine bug feces. Other Tryp- with T. gondii has also been recognized as an important anosoma spp. such as Trypanosoma evansi (primarily cause of retinochoroiditis in humans [37], as a result transmitted by biting flies), Trypanosoma brucei brucei of either prenatal infection or an infection that was and Trypanosoma congolense (both mainly transmitted acquired postnatally [38]. Cats are the definitive host of by tsetse flies) have been reported in cats with clinical T. gondii and typically shed oocysts for only 2 weeks in signs, but to a lesser extent than in dogs. These three a cat’s life [39]. Humans can become infected after acci- protozoans, common in many animal species, have been dental ingestion of oocysts (which have had time [days] very rarely associated with atypical human infections to sporulate, as they are not immediately infective after [44]. While vector-borne, for all these protozoans an excretion/shedding) from cat feces, feces-contaminated additional risk of mechanical infection exists, especially soil, water, fruits or vegetables, or through the ingestion via direct blood contact (e.g. needle injury when collect- of raw meat containing tissue cysts. Due to the restricted ing a blood sample). time of oocyst shedding in cats and the time needed for Toxocara cati, Toxocara malaysiensis (in Southeast oocysts to sporulate to reach an infective stage, food- Asia) and hookworms (Ancylostoma braziliense, Ancy- borne sources as well as soil and water contact may be lostoma tubaeforme, Ancylostoma ceylanicum and greater human risk factors for toxoplasmosis rather than Uncinaria stenocephala) are common helminthic endo- direct contacts with cats. Thus, while cat ownership parasites of cats with proven or suspected zoonotic is often top of mind when considering risk reduction potential. Toxocara cati can cause visceral, ocular and measures for toxoplasmosis, an awareness of the other neural larva migrans in humans, with children at rela- described sources of infection for people, which demand tively higher risk of T. cati infection due to behavio- different control measures, for example, ensuring meat ral predilections like geophagia, pica and coprophagia. is well cooked, raw produce is well washed and hands Hookworm, in particular A. braziliense, is the most com- are thoroughly washed following gardening or outdoor mon cause of ‘creeping eruptions’ or chronic cutaneous play is required [26]. Further potential zoonotic proto- larva migrans in people [45]. Ancylostoma ceylanicum is zoan feline parasites are Giardia duodenalis and Crypto- an important emerging zoonosis and now recognized as sporidium spp. Giardia duodenalis, also named Giardia the second most common hookworm infecting humans intestinalis and Giardia lamblia, which is the species in the Asia Pacific [46, 47]. Echinococcus multilocula- that infects mammals. Giardia duodenalis is consid- ris is another relevant zoonotic endoparasite causing ered a species complex that comprises several genotypes severe alveolar echinococcosis in people. Cats can serve or assemblages; assemblages A and B frequently infect as a definitive host for E. multilocularis, but their role in humans and other mammals, while others are host-spe- maintaining the life cycle and their true zoonotic risk is cific [40]. Most G. duodenalis isolates from cats are typed under debate [48, 49]. However, cats have been shown as assemblage F [40, 41]. However, besides this cat-spe- as a possible source of infection for humans in Europe, cific assemblage, cats harbor other genotypes that can be though to a lesser extent than dogs [50, 51]. Cats become transmitted to humans such as assemblages A and B [40]. infected with Dipylidium caninum by ingestion of Regarding Cryptosporidium spp., cats are rarely infected, infected fleas or lice when grooming [52]. Humans, par- and typically with host-specific Cryptosporidium felis, ticularly young children, can acquire D. caninum infec- while humans are usually infected with Cryptosporidium tion by accidental ingestion of infected fleas. Dirofilaria hominis and Cryptosporidium parvum [42]. However, spp. and Brugia spp. are zoonotic filarial nematodes C. felis infections have also been detected in humans transmitted by mosquito bites to cats in endemic areas. [43]. People generally acquire Cryptosporidium spp. and Human lymphatic filariosis caused by Brugia malayi G. duodenalis infections by drinking contaminated water occurs in South India, Sri Lanka and some foci in South- during recreation or by direct contact with infected cat- east Asia. In endemic areas of Europe and the Americas, tle. Cryptosporidium felis oocysts and G. duodenalis cysts Dirofilaria immitis is mainly responsible for unifocal are immediately infective after excretion/shedding, so or multifocal pulmonary nodules in people. Dirofilaria people with immunosuppression should be careful when repens is distributed in Europe and in Asia and may cause handling cats with diarrhea. a wide range of symptoms in people based on the tissue Vector-borne zoonotic protozoans include the agents involved during migration and the location of the final of some severe human diseases such as zoonotic leish- subcutaneous nodular lesions [53]. Human thelaziosis maniosis, caused by Leishmania infantum and transmit- caused by the two feline eyeworm species, Thelazia cal- ted by phlebotomine sand fly bites in both the Old and lipaeda and Thelazia californiensis, may manifest as con- New Worlds, and American trypanosomosis or Chagas junctivitis, but keratitis and corneal ulcers can also occur disease, caused by Trypanosoma cruzi and transmitted [54–56]. Maggi et al. Parasites & Vectors (2022) 15:415 Page 10 of 15 To avoid oral infections with parasites, cats should arthritis, osteoarthritis and septic shock [66, 76–83]. be prevented from hunting and ingesting raw meat, as Besides a general risk of transmission via cat bites with infected intermediate hosts or contaminated raw meat these pathogens, there is an additional exposure risk to may represent sources of infection for endoparasites. antimicrobial-resistant (AMR) bacteria, which have been Daily removal of feces from the soil or litter tray will demonstrated in the feline oral cavity [84]. Urogenital decrease the risk of environmental contamination and (Coxiella, Leptospira) or ocular and respiratory (Bor- infection in animals and humans by T. gondii oocysts, detella and Chlamydia)-associated pathogens are also hookworm and Toxocara eggs as they are not immedi- common. Coxiella burnetii can cause reproductive dis- ately infective after being excreted in cat feces. This is in orders in animals; clinical signs and symptoms in people contrast to E. multilocularis eggs (morphologically indis- are variable, including febrile illness, pneumonia, hepa- tinguishable from Taenia spp. eggs), which are imme- titis and reproductive disorders. While farm animals are diately infective, thus demanding feces removal with considered the main source for zoonotic infections, cats special care. have also been associated with zoonotic infections [85]. For all vector-transmitted parasites, comprehensive Zoonotic transmission can occur by aerosol contami- vector control is the most relevant approach. For some nation after contact with placenta or amniotic fluids of parasites, e.g. heartworm, specific prophylaxis is essential both aborting and healthy cats [85, 86]. Leptospira spp. [57–60]. can cause infections in mammals, including cats and While several viral infections occur in cats (see above), humans [87]. Infection is transmitted by direct contact only a few viruses, such as rabies virus and several other with infected urine or by contact with infected water or lyssaviruses, are recognized as potential agents of viral soil. Although cats have not been considered as a main zoonoses. Several cases of cat-associated zoonotic cow- source of infection for people, specific antibodies against pox infections, leading to dermal subcutaneous tissue Leptospira spp., Leptospira spp. DNA as well as a posi- necrosis, neurogenic inflammation, colliquative lym- tive bacterial culture in urine and kidneys have been phadenitis or ocular disease, have been described in peo- detected in cats [88, 89]. Bordetella bronchiseptica causes ple exposed to cat scratches or bites [61–65]. In fact, it chronic respiratory infections in cats, dogs and humans is estimated that 50% of human cowpox cases in the UK [15]. However, zoonotic transmission is infrequent, and are due to transmission from cats [66]. Other well-rec- most cases occur in immunocompromised patients. ognized viruses, such as the avian influenza virus H7N2, Chlamydia felis causes respiratory and ocular infections can also be transmitted from cats to humans [67–72]. in cats, particularly in multi-cat environments [90]. Chla- Vector-borne viruses infecting both cats and humans mydia felis is transmitted by direct contact between cats include West Nile virus and, in Asia, Dabie bandavirus, and between humans and cats. Again, the risk of zoonotic a phlebovirus causing the severe fever with thrombocyto- transmission is extremely rare and is highest in immuno- penia syndrome (SFTS) (see Table 1). compromised individuals; infection in humans is mainly Campylobacter, Salmonella, Clostridium and Yersinia asymptomatic or causes acute conjunctivitis [90]. are a few examples of enteric zoonotic bacteria that can The most common fungi that are directly zoonotic are be passed to humans by feces (i.e. ingestion of the infec- the dermatophytes (e.g. Microsporum spp., Trichophy- tious agent in contaminated food, water, other envi- ton spp.) and the genus Sporothrix spp. Both groups are ronmental sources or via hands) or direct contact with transmitted from cat to cat and from cat to people [91]. infected cats. Additionally, enteropathogenic Escheri- Cats can be subclinical carriers of Microsporum canis chia coli (EPEC) and enterohemorrhagic Escherichia with long-haired breeds and kittens particularly associ- coli (EHEC) are reported in cats, presenting a potential ated with infected premises. About half of people living source of human infection [73, 74]. Shiga toxin-produc- in households with dermatophyte-infected cats develop ing Escherichia coli (STEC), responsible for the hemolytic ringworm lesions [92]. In tropical and subtropical regions uremic syndrome in humans, have also been detected in in Latin America, thermodimorphic fungi of the genus dog as well as in cat fecal samples with virulence genes in Sporothrix are responsible for the most frequent subcuta- common with isolates from humans, thus constituting a neous mycosis, with endemic occurrence [93]. Cat-trans- potential additional source of human infection [75]. mitted sporotrichosis caused by Sporothrix brasiliensis Bacteria of the genera Bartonella, Capnocytophaga, has been a zoonosis of the south and southeast regions Francisella, Pasteurella, Rickettsia, Staphylococcus and of Brazil for more than 20 years [94]. The systemic fungi even Yersinia (i.e. Yersinia pestis) are known zoonotic Blastomyces dermatitidis, Coccidioides immitis and His- pathogens of cats that can cause serious skin and sys- toplasma capsulatum, which are common to some coun- temic infections in people and that may result in severe tries like the USA, are acquired from the environment. sequelae including meningitis, endocarditis, septic Thus, outdoor cats in endemic areas are of greatest risk. Maggi et al. Parasites & Vectors (2022) 15:415 Page 11 of 15 These agents are generally not transferred among cats or infestations on relocated cats pose a particular risk in between cats and their owners and handlers [95]. terms of zoonotic exposure and wider spread of infection. Zoonoses with a preferred direction of transmission Measures to prevent vector-borne pathogen transmis- from humans to animals should also be considered. sion and establishment of exotic vectors are therefore an For example, cats have been shown to be susceptible to important consideration, and veterinary team profession- SARS-CoV-2 infection acquired in COVID-19-positive als have a vital role to play in raising awareness among households with mild to severe feline respiratory dis- cat owners and rescue charities, maintaining biosecurity, ease observed [96, 97]. However, cats shed SARS-CoV-2 reducing zoonotic risk and improving the welfare of relo- for only short periods of time, and to date only one case cated cats. When assessing homing cats, professionals of cat-to-human transmission has been reported [98, should: 99]. Other examples for this direction of transmission include influenza A viruses, Mycobacterium tuberculo- i. Educate the public regarding the risks of adopt- sis, methicillin-resistant Staphylococcus aureus (MRSA), ing cats from abroad or distant regions. Education Helicobacter pylori, Entamoeba histolytica, Streptococcus regarding the benefits of local cat adoption and pneumoniae and other Streptococcus spp. the importance of considering the lifestyle of the Many of the above-mentioned zoonotic pathogens are cat will also enable potential new owners to make vector-borne (see Table 1). Up to 80% of Ctenocephal- informed choices. This communication should be ides felis collected from cats contain the DNA of either compassionate and non-confrontational as most a human or cat pathogen [100], with Bartonella spp. like charities working in this field, and people adopting Bartonella henselae and Bartonella clarridgeiae, Rickett- pets, do so with the best of intentions. sia felis and the hemoplasmas being the most common. ii. Ensure that neutering has been carried out and Thus, any type of cat with increased risk of flea infestation that the cat ideally has a registered microchip or an is more likely to be a carrier of human or feline patho- alternative form of identification in case microchip- gens. Tick-borne disease agents have also been increas- ping is unavailable. As well as playing an important ingly found in the blood of cats as molecular diagnostic role in population control, neutering helps prevent techniques have become more sensitive and available to horizontal (venereal and through fighting) and ver- veterinary health care providers. We now know that tick- tical transmission of pathogens. Microchipping infested cats can harbor several zoonotic as well as non- or alternative identification allows previous treat- zoonotic infectious agents, such as Anaplasma spp. (both ment, vaccination status and testing records to be Anaplasma phagocytophilum and Anaplasma platys), traced. Bartonella spp., Babesia spp., Borrelia spp., Cytauxzoon iii. Ask about origin and travel history for any recently spp., Ehrlichia spp., Hepatozoon spp. and Rickettsia spp. acquired cat. This will allow for the selection of [101–104]. appropriate diagnostic tests and potential treat- Generally, individual cats can be infected subclinically ments depending on pathogens and vectors pre- with feline vector-borne pathogens and therefore may sent in the region of origin and clinicopathological be potentially homed into households or locations previ- alterations. Online maps are available for profes- ously naïve to the corresponding agent. The clinical signs sionals to consult to determine known risks in dif- of disease associated with feline vector-borne pathogens ferent geographical locations (e.g. https://​www.​ have been reviewed in detail [9, 10, 105]. For the flea- and esccap.​org/​guide​lines-​maps/, https://​capcv​et.​org/​ tick-borne agents, fever is generally the most common maps/#/, https://​cvbd.​elanco.​com/​cvbd-​maps). clinical sign (see Table 1). iv. Perform a thorough clinical examination. Particu- lar attention should be paid to the oral cavity, eyes, Screening, testing, education skin, feet and claws/nail beds as these areas are and recommendations particularly likely to be affected by feline vector- When cats are relocated, disease risks are often under- borne diseases. Skin lesions may also indicate the estimated. This is in part due to a lack of data, espe- presence of current arthropod infestations. Fur- cially regarding vector-borne pathogens in cats and, as ther information on clinical signs associated with mentioned earlier, because cats are often perceived to vector-borne and other pathogens can be found be at a lower risk of tick-borne pathogens in particular at the following websites: https://​www.​esccap.​org, or arthropod-borne diseases in general. Vector-borne https://​capcv​et.​org, https://​www.​trocc​ap.​com, pathogens in cats commonly occur, pose disease risks to https://​cvbd.​elanco.​com, http://​www.​abcdc​atsve​ts.​ the individual cat as well as the wider feline populations org/. and have zoonotic potential (see Table 1). Arthropod Maggi et al. Parasites & Vectors (2022) 15:415 Page 12 of 15 v. Thoroughly check the cat for parasites. Check for infections and help charities and potential new owners to fleas, ticks and lice and ensure that preventative assess the long-term disease risks for any individual cat. treatment against these and other ectoparasitic arthropods is in place if ongoing risk is a concern. Consensus statement If allowed to establish, flea household infestations Large numbers of unowned cats continue to consti- represent an interface where zoonotic patho- tute an animal welfare, ecological, societal and public gens such as Bartonella spp., Rickettsia spp. and health problem worldwide. While well-planned and Dipylidium caninum could be transmitted. Rhipi- executed population management programs, such as cephalus spp. and Ixodes spp. ticks are also capable Trap-Neuter-Return (TNR), are a key component in the of establishing infestations in households, the for- long-term control of street cat numbers, other strate- mer indoors and in catteries, and both in gardens, gies are required for managing existing populations. allowing onward transmission of tick-borne path- Relocation and homing of unowned cats is one strat- ogens. The importance of ongoing preventative egy used in many parts of the world. However, a lack arthropod prevention should be emphasized to cat of understanding of an individual cat’s lifestyle and dis- owners and handlers, if there is a recognized risk, ease status can lead to animal and/or owner stress, the to limit zoonotic risk and vector-borne pathogen dissemination of feline pathogens and an increased risk transmission. Check for endoparasites and ensure of exposure to zoonotic agents. Raising awareness of that owners are advised on an effective treatment these issues among veterinary team professionals and regime based on origin of cat, diagnosed infections those working with cat charities is therefore essential. and future exposure risk. This includes knowledge of the cat’s lifestyle, includ- vi. Ascertain the cat’s FeLV/FIV status and consider ing how this influences exposure to pathogens, the clinical pathology evaluations. As well as being sig- geographic distribution of cat pathogens, their clinical nificant pathogens in their own right, FeLV and/or signs and/or clinicopathological abnormalities, vectors, FIV are risk factors for several other infections in modes of transmission, vector and parasite control and cats. Biochemistry, hematology profiles and urinal- evaluation of the potential zoonotic risks. Appropriate ysis are also useful to check for thrombocytopenia, testing, surveillance, recording and reporting of infec- anemia and hyperglobulinemia or proteinuria, for tious agents in homed cats is also a vital component in example, because these can be suggestive of vector- tracking the geographic spread and emergence of feline borne diseases. pathogens and zoonoses. vii. Report all relevant findings, especially foreign arthropods and pathogens. Very few vector-borne pathogens in cats are notifiable by law. Therefore, Conclusions reporting unusual findings to local health authori- While large numbers of unowned cats continue to be a ties, universities, independent organizations such welfare issue globally, a multifaceted approach to con- as those mentioned in point IV and peer-reviewed trolling cat numbers and associated pathogen trans- publications will help generate an up-to-date mission is vital. The increased relocation and homing picture of where vectors and pathogens may be of unowned cats to reduce feline suffering and social emerging. Examples of published reports include problems mean that strategies are required to reduce Leishmania spp. and Hepatozoon spp. found in cats accompanying pathogen spread and zoonotic risk. in Germany [106], Rhipicephalus pusillus found on Increasing veterinary education regarding cat lifestyles, cats in France [107] or Haemaphysalis leachi found at risk pathogens and their vectors alongside increased on cats imported from Africa to the UK [108]. testing, surveillance and overpopulation control with TNR systems is key to achieving these aims. Cooperation with cat charities and rescue organiza- Acknowledgements tions is beneficial as the above steps are most effective if We thank Elanco Animal Health for organizing and facilitating our discussion carried out before homing takes place. Adequate arthro- at the 15th Symposium of the CVBD World Forum (online) in November 2020. pod protection, before homing or during the TNR pro- We thank Dr. Claudia Böhm and Dr. Tamara Hofstede, Elanco Animal Health, for critical review and scientific support. gram, will minimize the risk of flea-, tick-, mosquito- or sand fly-borne pathogen transmission. Identification of Author contributions infection and disease before relocation and including this RM, VH, ML, MGP, XR, VS, ST and IW drafted the initial statement and the tables. FK and BS coordinated the publication process and drafted the statement evaluation in any viability assessment for homing will and the tables. All other co-authors worked on the initial manuscript and also help to reduce further transmission of any existing contributed to the final document. All authors read and approved the final manuscript. Maggi et al. Parasites & Vectors (2022) 15:415 Page 13 of 15 Funding Received: 9 September 2022 Accepted: 9 October 2022 FDT received a research fellowship from CNPq (Bolsa de Produtividade; Grant No. 313118/2018-3). FK is working in a project funded by Elanco Animal Health at TransMIT GmbH, Germany. The participation of LC was supported by the projects UIDB/CVT/00772/2020 and LA/P/0059/2020 funded by the Portuguese Foundation for Science and Technology (FCT). References 1. Wright I, Jongejan F, Marcondes M, Peregrine A, Baneth G, Bourdeau Availability of data and materials P, et al. Parasites and vector-borne diseases disseminated by rehomed Not applicable. dogs. Parasit Vectors. 2020;13:546. 2. Statista. Number of dogs and cats kept as pets worldwide in 2018. 2019. https://​www.​stati​sta.​com/​stati​stics/​10443​86/​dog-​and-​cat-​pet-​ Declarations popul​ation-​world​wide/. Accessed 9 Aug 2021. 3. World Atlas. 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Epidemiology of sporotrichosis in Latin America. Myco‑ • thorough peer review by experienced researchers in your field pathol. 1989;108:113–6. • rapid publication on acceptance 94. Lopes-Bezerra LM, Mora-Montes HM, Zhang Y, Nino-Vega G, Rodrigues • support for research data, including large and complex data types AM, de Camargo ZP, et al. Sporotrichosis between 1898 and 2017: the evolution of knowledge on a changeable disease and on emerging • gold Open Access which fosters wider collaboration and increased citations etiological agents. Med Mycol. 2018;56:126–43. • maximum visibility for your research: over 100M website views per year 95. Lloret A, Hartmann K, Pennisi MG, Ferrer L, Addie D, Belák S, et al. Rare systemic mycoses in cats: blastomycosis, histoplasmosis and coccidioi‑ At BMC, research is always in progress. domycosis: ABCD guidelines on prevention and management. J Feline Med Surg. 2013;15:624–7. Learn more biomedcentral.com/submissions