Gastrointestinal protozoan and helminth parasite infections in captive chameleons in Germany

Gastrointestinal protozoan and helminth parasite infections in captive chameleons in Germany | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Gastrointestinal protozoan and helminth parasite infections in captive chameleons in Germany Paula Sapion-Miranda, Anja Taubert, Carlos Hermosilla, Malek J. Hallinger This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8902969/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 9 You are reading this latest preprint version Abstract Within lizards, chameleons (family Chamaeleonidae) represent a distinctive and highly specialized clade of insectivorous reptiles. Chameleons have become more popular as reptile pets in recent years worldwide, due to their feasible husbandry and attractiveness. To date, not only reptile pet numbers but also health-compromising endoparasite infections are rising. In the current study, we analyzed a total of 670 faecal samples of 25 different chameleon species kept as pet animals all over Germany. In total, 223 animals (33%) were found to harbor gastrointestinal parasites. In total, 10 different parasite taxa were diagnosed in chameleon fecal samples. The most prevalent parasite stages were oxyurid eggs (14.1%), followed by coccidia oocysts (7.8%) (e. g. Isospora jacarimanni , Isospora spp., Choleoeimeria spp., Eimeria spp.), flagellated protozoan trophozoites (6.3%) (e. g. Leptomonas spp., Tritrichomonas spp., Proteromonas spp.), heterakid eggs (3.6%), thin-shelled nematode eggs (0.6%), digenean trematode eggs (0.6%), pentastomid eggs (0.3%), ascarid eggs (0.2%), Entamoeba spp. cysts (0.2%) and physalopteroid eggs (0.2%). Among current chameleon species, veiled chameleons ( Chamaleo calyptratus ) showed a significant higher prevalence of oxyurid infections (64%) than other chameleon species. Co-infections were diagnosed in 9.8% (22/223) of faecal samples. When performing necropsies, 19/31 chameleons showed parasite-induced pathologies based on histopathological examinations, including one animal being infected with 4 parasite species ( Leptomonas spp., Tritrichomonas spp., Foleyella furcata and Hexametra angusticaecoides) , thereby most likely impacting animal’s wellfare. chameleons parasites exotic pets reptile medicine wildlife Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 1. Introduction To date, reptile husbandry is increasing in popularity since captive-bred reptiles can easily be obtained as exotic pets (Carpenter et al. 2004 ). However, not all reptiles are available from breeders, and enthusiasts seeking for exotic options may turn to wild-caught specimen with potentially unknown origin (Hallinger et al. 2018 ; Marshall et al. 2020 ; Toland et al. 2020 ). This ambiguity contributes to the introduction of numerous potentially invasive (neozoan) pathogens of these reptiles, including both ecto- and endoparasites (Rataj et al. 2011 ). Both wild and captive reptiles are frequently infected with endoparasites, such as apicomplexans, flagellated protozoa, nematodes, cestodes, trematodes, and pentastomids (Frank 1985 ; Paré 2008 ; Pasmans et al. 2008 ; Pantchev and Beck 2013 ; Hallinger et al. 2018 ). While many of these parasites typically own low pathogenicity in their original habitats, they can significantly impact the animal’s health if environmental conditions change (Wilson and Carpenter; 1996 ). Hence, several risk factors like inadequate temperatures, stress, and high-density stocking affect the reptile's immune system (Bower et al.; 2019 ). In this context, enhanced stress during trapping and transport of wild-caught reptiles often drives an increase in morbidity and mortality rates (Bower et al. 2019 ; Stets 2019 ). In captive reptiles, a notable prevalence of gastrointestinal parasitoses is common, especially referring to parasites with monoxenous lifecycles (Schneller and Pantchev 2008; Pantchev and Beck 2013 ; Hallinger et al. 2019 ). This phenomenon can be attributed to the high environmental tenacity of certain endoparasite stages (eggs) and short prepatencies of related parasites (Pasmans et al. 2008 ; Cervone et al. 2016 ; Hallinger et al. 2018 ). Distinct intrinsic factors, such as age, gender, genus and species or extrinsic factors like poor hygiene, stress and nutrition can significantly contribute to the development of clinical diseases in reptiles (Pasmans et al. 2008 ; Pantchev and Beck 2013 ; Hallinger et al. 2018 ). Moreover, inadequate housing conditions create a perfect environment for high parasite burdens and frequent reinfections (Pasmans et al. 2008 ; Pantchev and Beck 2013 ; Wolf et al. 2014 ; Hallinger et al. 2019 ). Recent studies have suggested that feeding insects also represents a significant risk factor of endoparasite transmission, requiring further investigations (Gałęcki and Sokół 2019 ; Müller et. al 2019 ). Within lizards, chameleons (family Chamaelonidae) represent a distinctive and highly specialized clade of insectivorous reptiles. Chameleons, renowned for their attractive appearance, have been successfully bred in captivity for over four decades, while their trading dates to the early 1800s (Carpenter et al. 2004 ; Diaz et al. 2015 ). Similar to other captive reptiles, chameleons often show a high prevalence of endoparasites, a health-compromising condition that has raised considerable concern among herpetologists (Okulewicz et al. 2015 ; Rom et al. 2018 ). Despite this fact, there remains a gap of understanding on parasite transmission, parasite-driven pathogenesis and clinical manifestations (Nash et al. 2023 ). Moreover, many chameleon species, such as the Belalanda chameleon ( Furcifer belalanda ) from Madagascar or the Nguru spiny pygmy chameleon ( Rhampholeon acuminatus ) from Tanzania, fall into categories of critically endangered (4%) animals, others as endangered (21%), vulnerable (12.2%) or nearly threatened (18%), due to factors like habitat loss, illegal exotic pet trade and others (The IUCN Red List of Threatened Species 2019 ). In the context of parasite infections, chameleon ectoparasitism seems relatively rare when compared to other reptiles, since no cases of infestations have been described so far in captive animals. In contrast, infestations with mites belonging to the family Trombiculidae have been described in wild chameleons (Pantchev 2011 ; Eckhardt et al. 2019 ). However, it is important to note that more detailed research is needed to clarify the occurrence of ectoparasite infestations in chameleons kept as pets. In Germany, there are only few studies on parasites affecting chameleons. Consistently, Biallas ( 2013 ) conducted an extensive study on 212 chameleon faecal samples to investigate the prevalence and detrimental effects of endoparasites in captive chameleons, considering animal risk factors like origin, age and sex. Further, Pantchev ( 2011 ) examined 66 chameleon faecal samples and provided an overview on most common parasite infections and related clinical symptoms. Stets ( 2019 ) analyzed 646 feacal samples of Panther chameleons ( Furcifer pardalis ), thereby comparing endoparasite infections in wild-caught and captive-kept animals in the Ukraine. Additionally, some case reports on distinct parasitoses and related treatments in captive chameleons (e. g. F. pardalis, Chamaleo calyptratus ) were recorded (Damiani et al. 2006 ; Reitl et al. 2021 ; Heng et al. 2023 ). Some reports documented endoparasitoses of wild-caught chameleons (Dillehay et al. 1986 ; Lhermitte-Vallarino et al. 2008 ; Heckmann 2014 ; Pellett et al. 2014 ; Collicutt et al. 2017 ; Eckhardt et al. 2019 ) mostly focusing on parasite species with heteroxenous lifecycles. Occasionally, studies on reptile endoparasites also included individual findings on the most commonly kept chameleon species ( Chamaelo calyptratus ) (Wilson and Carpenter, 1996 ; Goldberg et al. 2009; Rataj et al. 2011 ; Okulewicz et al. 2015 ; Rom et al. 2018 ; Satour and Dewir 2018 ), but lacking both large sample numbers and representative data. In general, the identification of pathogenic endoparasite species, including anthropozoonotic ones, is crucial for adequate targeted parasitological treatments, thereby minimizing both chameleon morbidity/mortality and transmission to pet owners (Pantchev 2011 , Mendoza-Roldan et al., 2020 ). Therefore, the current study aimed to provide novel insights into the occurrence of gastrointestinal endoparasites in captive-kept chameleons in Germany by investigating a substantial sample size ( n = 670) but also involving 25 different chameleon species to ensure a representative analysis. In addition to faecal analyses, we performed histopathological examinations on 31 deceased chameleons, providing a new baseline dataset on the presence of endoparasites in these popular herpetological pet animals. Figure 1. Exemplary images of different chameleon species investigated in this study a. a male Panther chameleon ( Furcifer pardalis ), b. a male Parson’s chameleon ( Calumma parsonii ), c. a male Malagasy giant chameleon ( Furcifer oustaleti ), d. a female Veiled chameleon ( Chamaelo calyptratus ). Table 1 Number and origin of chameleon fecal samples (total n = 670) Chameleon species Common name No. examined Origin (private/ vet / zoo) Furcifer pardialis Panther chameleon 250 223/16/11 Chamaeleo calyptratus Veiled chameleon 232 200/30/2 unknown species 103 84/14/5 Calumma parsonii Parson’s chameleon 23 12/0/11 Trioceros melleri Meller’s chameleon 12 11/0/1 Chamaeleo dilepis Flap-necked chameleon 10 9/1/0 Furcifer lateralis Carpet chameleon 6 6/0/0 Trioceros jacksonii Jackson’s chameleon 6 6/0/0 Furcifer oustaleti Malagasy giant chamaleon 5 5/0/0 Trioceros hoehnelii Höhnels chameleon 5 4/1/0 Triceros rudis Coarse chameleon 3 3/0/0 Bradypodion thamnobates Natal Midlands dwarf chameleon 2 2/0/0 Bradypodion fischeri Fischer’s chameleon 2 0/2/0 Trioceros Bitaeniatus Side-striped chameleon 2 2/0/0 Trioceros j. xantholophus Yellow-crested Jackson’s chameleon 2 2/0/0 Furcifer verrucosus Warty chameleon 1 0/1/0 Kinyongia boehmei Böhme’s two-horned chamaleon 1 1/0/0 Rampholeon acuminatus Nguru pygmy chameleon 1 0/1/0 Bradypodion setaroi Setaro’s dwarf chameleon 1 1/0/0 Trioceros johnstoni Johnston’s chameleon 1 0/1/0 Trioceros sternfeldi Crater Highlands side-striped chameleon 1 1/0/0 Trioceros ellioti Elliot’s chameleon 1 1/0/0 2. Materials and Methods 2.1. Faecal samples In total, 670 faecal samples of captive chameleons representing 25 species of different genera (e. g. Chameleo , Furcifer , Triocerus , Calumma , Bradypodion , Rhampholeon , Kinyongia , see Fig. 1) were sent to exomed GmbH laboratory, Marburg, Germany (Table 1 ) between 2012–2019 and examined coprologically. Of these, 573 (85.8%) came from private owners, 67 (10%) from veterinary surgeons, and 30 (4.5%) from zoological gardens across Germany (Fig. 2). Each faecal sample was accompanied by a submission form stating the signalement of the animal (i. e. species, sex, weight, and age) and a short anamnesis (husbandry type, previous parasitological diagnoses, and antiparasitic treatments, see Online Resource 1). To identify different parasites and their stages, direct saline fecal smears (DSFS) were performed according to Cooper ( 2009 ). For DSFS preparation, faeces (10 g) were mixed at a ratio of 1:1 with 0.9% NaCl solution to a uniform suspension of the sample, placed on a microscope slide, covered with a coverslip (22 x 22mm; Nunc) and examined by a light microscope (100x or 400x magnification, Axio Imager M1®, Zeiss, Jena). Oxyurid, heterakid, ascarid, Strongyloides , trematode and penstatomid eggs as well protozoan oocysts, cysts and trophozoites were identified based on previous description guidelines (Barnard and Upton 1994 ; Wilson and Carpenter 1996 ; Schneller and Pantchev 2008; Pantchev 2011 ; Hallinger et al. 2019 ). Faecal samples were listed as positive (Table 2 ), if at least one stage of the above described endoparasites posing a health-compromising risk for chameleons or other lizards was diagnosed. Samples lacking any pathogenic parasitic stages or showing non-pathogenic genera (e.g. Nyctotherus ) as well as pseudoparasites, were here classified as negative. 2.2 Chameleon necropsies A total of 31 chameleon individuals ( n = 31) were necropsied at exomed GmbH laboratory (Table 3 ). In these cases, the signalement (i. e. species, age, sex) and basic medical history, [e. g. clinical signs, husbandry, food, cause of death (natural death/euthanization by veterinary surgeons)] was reported (Online Resource 2). Additionally, tissue samples were collected and fixed in formalin for pathohistological examinations as previously described (Stacy 2021 ). In detail, the chameleon necropsy was performed on its body side by an incision along the ventral midline from the cloaca to the forelimb and shoulder blade and up to the dorsal side. By this incision procedure, the coelomic cavity was exposed without obstructions (Farris et al. 2015 ). Primarily, the gastrointestinal tract was visually inspected for the presence of metazoan endoparasites. Additionally, intestinal content was collected for DSFS. For metazoan endoparasite diagnosis, morphological identification was performed in accordance to Pantchev ( 2011 ) and Hallinger et al. ( 2019 ) using a light microscope supplied with a digital camera (Axio Imager M1®, Zeiss, Jena). 2.3 Microbiological examination of necropsies For further microbial identification, faeces and/or coelom swabs were taken from necropsied animals. For bacterial and fungal cultivation in vitro , both the Columbia/MacConkey sheep blood agar (5%) as well as the Sabouraud dextrose agar (SDA) were used (BioMerieux, Charbonnier les Bains, France). In addition, bacterial isolates were identified by gram staining, oxidase, and catalase tests (BioMerieux, Charbonnier les Bains, France), together with a commercially available API 20 E/NE kit (BioMerieux, Charbonnier les Bains, France), as formerly described in reptiles (Marenzoni et al. 2015 ; Hallinger et al. 2019 ). Figure 2. Geographical origin of captive chameleon fecal samples ( n = 670) according to postcode districts in Germany. 3. Results 3.1 Faecal samples DSFS-based coproscopical analyses of pet chameleon faecal samples from Germany revealed a wide spectrum of gastrointestinal protozoan and metazoan parasites. In total, 33.2% (223/670) faecal samples proved positive for parasite infections. Among the positive samples from DSFS testing the following parasitic stages were diagnosed: oxyurid eggs (14.1%)(Fig. 3a), coccidian oocysts (7.8%)(Fig. 3b), flagellated protozoan trophozoites (6.3%)(Fig. 3c), heterakid eggs (3.57%), Strongyloides spp. eggs (2.9%), thin-shelled nematode eggs (0.6%), trematode eggs (0.6%)(Fig. 3d), pentastomid eggs (0.3%)(Fig 3e), ascarid eggs (0.15%) (Fig. 3f), Entamoeba cysts (0.15%) and Physalopteroides spp. eggs (0.15%) (please refer to Table 2). Neither cestode eggs nor Cryptosporidium oocysts were found in the current faecal samples. A total of 9.8% of the 223 parasite-positive chameleon samples showed co-infections (Fig. 4). Among these, 4.5% (10/223) of animals exhibited an ‘coccidia-nematode’ co-infection, while 3.6% (8/223) of chameleons had a ‘nematode-flagellated trophozoite’ co-infection. Additionally, 1.7% (4/223) of animals showed an ‘coccidia-flagellated trophozoite’ co-infection. Notably, in the faecal samples, no evidence of multiple infections of tri-, tetra- or higher-orders were detected. Oxyurid nematodes were the most prevalent metazoan parasites with 14.1%. Interestingly, the oxyurid prevalence varied with the chameleon genus, since it proved significantly higher in Veiled chameleons ( C. calyptratus : 64%) when compared to other chameleon species (36%) of this study (Chi-Square test: χ2=31.061, df= 1, P < 0.0001 ) . Overall, no significant correlation between parasite infection and age (i. e. juvenile 1 year of age) was found. The same holds true for other factors, such as sex or origin, since no significant correlations were observed. Other parasitic nematode stages like heterakid eggs (3.6%), embryonated Strongyloides spp. eggs (2.9%), and thin-shelled nematode eggs (0.6%) were found at a lower prevalence. Only one sample proved positive for ascarid eggs (0.1%) and another for Physalopteroides spp. eggs (0.1%). Intestinal trematode eggs were found in 4 samples (0.6%) whilst pentastomid eggs were present in two faecal samples (0.3%). When considering intestinal protozoan parasites, these revealed as the second most frequently diagnosed endoparasites. Among the protozoans, coccidia were the most prevalent in analyzed fecal samples. In this context, the genus Isospora spp. showed the highest prevalence among the identified apicomplexans with 5.1% (34/670). whilst Isospora jaracimmani (1.2%), Choleoeimeria spp. (0.3%) and un-sporulated apicomplexan oocysts (1.3%) were found at lower prevalences. Besides coccidia, Entamoeba cysts (0.15%) and cysts of intestinal flagellates (0.9%) were also diagnosed. Among the flagellated protozoa, Leptomonas spp. reached the highest prevalence (2.2%) followed by Trichomonas spp. (1.5%) whereas other flagellates like Tritrichomonas spp. (1.1%), Proteromonas spp. (0.3%) and Monocercomonas spp. (0.1%) ranked lower. We observed a statistical significant difference in the prevalence of coccidia between adult and juvenile animals since juveniles were more frequently affected ( p < 0.0009). 3.2 Chameleon necropsies A total of 31 animals originating from 10 different chameleons species were subjected to necropsy. Here, most animals (13/31, 41 %) belonged to the Panther chameleon ( F. pardalis ), species (details of origin Table 1). In necropsies, 19 chameleons (61%) tested positive for endoparasite infections (Table 3). Pathological examination revealed that severe parasitoses signified the cause of death in at least 13 animals (40.6%), including isosporosis ( n = 3; 9.3%) (Fig. 5a), choleoeimeriosis (n = 3; 9.3%) (Fig. 5b), heterakiosis ( n = 2; 6.2%)(Fig. 5c), oxyuridiosis ( n = 1;3.1%). An animal (3.1%) died due to a tetra-parasitic infection showing leptomonosis, trichomonosis, foleyellosis ( Foleyella furcata ) and hexametrosis (Fig. 5d). One Jackson’s chameleon ( T. jacksonii ) died due to severe protozoa-driven enteritis caused by Leptomonas spp. and Trichomonas spp. infections. Furthermore, there were five cases of co-infections ( n = 5, 15.6%). Other etiological causes identified for chameleon death were gout (15.6%), mycosis (9.37%), salmonellosis (9.37%), other bacterial infections (9.37%), intestinal obstipation (6.25%), mechanical trauma (6.25%), adenocarcinoma (6.25%), malnutrition (3.1%) and hypervitaminosis D3 (3.1%). In two chameleons, the cause of death remained unknown even though showing enteritis and hepatitis (see Online Resource 2). Table 2. Parasitological findings in chameleon faecal samples [(total: n = 670; positive for parasite stages: 223/670 (33.3%); negative: 447/670 (66.7%)] Parasite species No. positive samples Host species Oxyurid nematodes (Pharyngodonidae ) 95/670 (14.1%) C. calyptratus (61) F. pardalis (16) Unknown species (15) C. parsonii (2) R. acuminatus (1) Coccidia (e.g. Isospora jaracimmani, Isospora spp., Choloerimeria spp.) 53/670 (7.8%) F. pardalis (25) C. calyptratus (21) Unknown species (7) C. dilepsis (2) F. lateralis (1) T. jacksonii (1) Flagellated protozoa (e.g. Leptomonas spp., Tritrichomonas spp., Proteromonas spp.) 42/670 (6.3%) F. pardalis (13) C. calyptratus (12) C. dilepis (3) T. melleri (3) C. parsonii (2) F. oustaleti (1) T. jacksonii (1) Heterakids 24/670 (3.57%) F. pardalis (7) C. parsonii (7) Unknown species (4) F. oustaleti (3) C. calyptratus (2) B. thamnobates (1) Strongyloides spp. 20/670 (2.98%) C. calyptratus (12) Unknown species (9) F. pardalis (5) T. hoehnelii (3) T. rudis (1) Thin-shelled nematode eggs 4/670 (0.6%) T. melleri (1) T. j. xantholophus (1) C. dilepis (1) Unknown species (1) Trematode eggs (Diginea) 4/670 (0.6%) F. pardalis: (3) C. parsonii (1) Pentastomids 2/670 (0.30%) F. pardalis: (1) F. oustaleti (1) Ascarids 1/670 (0.15%) F. pardalis: (1) Amoeba 1/670 (0.15%) F. pardalis: (1) Physalopteroidea 1/670 (0.15%) F. oustaleti: (1) Figure 3. Exemplary photomicrographs of different parasitic stages found in chameleon faecal samples. a. operculated oxyurid egg (Veiled chameleon) b. sporulated Isospora sp. oocyst containing two sporocysts with four sporozoites each (Veiled chameleon), c. flagellated Leptomonas spp. trophozoites (Meller's chameleon), d. trematode egg with miracidium (Jackson's chameleon) e. embryonated pentastomid egg with larva (Malagasy giant chameleon), f. Hexametra angusticaecoides egg (Panther chameleon). Figure 4. Endoparasite mono- and co-infections in faecal samples of chameleons ( n = 670) kept in captivity in Germany Figure 5. Exemplary images from chameleon necropsies and related histological findings. a. Coccidian meronts (black arrows) in the intestinal epithelium of a Veiled chameleon (400x magnification), b. close-up of a dilated gallbladder (black arrow) caused by bile duct obstruction associated with a severe Choleoeimeria sp. infection, c. cross section of a Spinicauda sp. nematode (black arrows) in the small intestine mucosa of a Panther chameleon (200x magnification), d. adult Hexametra angusticaecoides nematode (indicated by black arrow) removed from the subcutaneous tissue, co-infected with Foleyella furcata in the serosa, exposed after removal of the skin (asterisk) of a Panther chameleon. Table 3. Parasitological findings in chameleon necropsies [(total: n = 31; positive for parasite stages: 19/31 (61.2%)] Parasite Species No. of positive (%) Host species n. Flagellated protozoa (e.g. Leptomonas spp., Tritrichomonas spp., Proteromonas spp.) Total: 6 (19.3) F. pardalis (3) C. calyptratus (1) T. melleri (1) T. jacksonii xantholopus (1) Isospora spp. Total: 5 (16.1) C. calyptratus (1) F. pardalis (3) T. melleri (1) Oxyurid nematodes (Pharyngodonidae ) Total: 4 (12.9) C. calyptratus (4) Choleoeimeria spp. Total: 3 (9.6) F. pardalis (1) T. melleri (2) Heterakids (e.g. Spinicauda spp.) Total: 2 (6.4) F. pardalis (1) B. thamnobates (1) Hexametra angusticercoida Total: 1 (3.2) F. pardalis (1) Foleyella furcata Total: 1 (3.2) F. pardalis (1) Strongyloides sp. Total: 1(3.2) Calumma parsonii (1) 4. Discussion The present large-scale epidemiological study on gastrointestinal parasites of chameleons kept in captivity in Germany reveals a high diversity of protozoan and metazoan endoparasite species, thereby confirming findings from previous prevalence studies (Pantchev 2011 ; Biallas 2013 ). Referring to chameleon oxyurid species, Parapharyngodon kenyaensis (Goldberg and Bursey 2008 ), Pharyngodon dimorpha and Thelandros meridionalis (Brygoo 1963 ) have been so far described in wild- and captivity-kept chameleons (Okulewicz et al. 2015 ; Eckhardt et al. 2019 ; Stets 2019 ). In the current study, oxyurid infections represented the most frequently found parasitosis (14.1%). However, earlier publications reported on a considerably higher prevalence (27.3% in Pantchev 2011 and 25.5% in Biallas 2013 ). This might rely on an increasing awareness of owners leading to a more frequent testing for gastrointestinal parasite infections and subsequent treatment in the last years. Among the various chameleon species assessed for oxyuridosis, we observed a notable difference in prevalence. Hence, C. calyptratus showed a significantly higher oxyurid prevalence than F. pardalis and less commonly kept exotic chameleon species. This finding may be explained by its overrepresentation in the sample pool due to its broad availability and low purchase costs, unfortunately often resulting in suboptimal husbandry conditions given by unexperienced owners. In contrast, other, more expensive species like the Panther chameleon or more exotic taxa are typically kept by experienced owners, who may also be more aware on infection monitoring measures. In contrast to captivity, the chances of reinfections are lower in the wild, and a delicate equilibrium between reptile hosts and their oxyurid species has been described. Thus, certain oxyurid nematodes efficiently support the intestinal microbiome to keep the animal in balance (Schneller and Pantchev 2008; Hallinger et al. 2018 ; Wellehan and Walden 2018 ; Eckhardt et al. 2019 ). In captivity, however, chameleons are confined to limited space and often suffer from suboptimal husbandry conditions, disrupting this equilibrium and leading to massive infections with these monoxenous tenacious parasites, finally resulting in (sub)clinical disease (Schneller and Pantchev 2008; Rinaldi et al. 2012 ; Pantchev and Beck 2013 ; Hallinger et al. 2019 ). Oxyuridosis in chameleons is usually of low pathogenicity, though severe infections can cause malabsorption, weight loss and nonspecific signs, especially in young, gravid or immunocompromised animals (Pantchev 2011 ; Biallas 2013 ; Pantchev and Beck 2013 ; Hallinger et al. 2018 ; Kubiak 2020 ). In the current study, oxyurid mono-infection was restricted to solely one necropsied Veiled chameleon, however this parasitosis was accompanied by bacterial co-infections with Salmonella enterica and Stenotrophomonas maltophila . Consequently, it is challenging to attribute the cause of death solely to oxyuridosis. This assumption aligns well with former findings on low intrinsic pathogenicity of oxyurids, reporting that mono-infections did not reveal as single cause of death (Biallas 2013 ). Strongyloides spp. nematodes are widespread parasites affecting captive and wild animals, including chameleons (Biallas 2013 ). The current study revealed a Strongyloides spp. prevalence of 2.9% in chameleons, which is consistent with previous findings (1.5%, Pantchev 2009 , 2011 ). Strongyloides infections may cause both respiratory and gastrointestinal signs due to larval migration and can be fatal (Schneller and Pantchev 2008; Biallas 2013 ; Heng et al. 2023 ). The prevalence of heterakids and ascarids in the current study relatively low, when compared to formerly recorded prevalences of 6–7% in chameleons (Pantchev 2009 ; Biallas 2013 ). Spinicauda sonsinoi (Brygoo 1963 ) and S. inglisi (Chabaud and Brygoo 1962 ) are the most common heterakid species in Malagasi chameleons. These parasites show a rather low pathogenicity if the parasitic burden is low (Schneller and Pantchev 2008). However, ascarid species like Hexametra angusticaercoides bear high pathogenic potential and severe hexametriosis often leads to animal death (Schneller and Pantchev 2008; Pantchev 2011 ; Barton et al. 2020 ; Reitl et al. 2021 ). Of note, H anguisticaercoides has a dual transmission mode in chameleons, an indirect infection route via consumption of H. angusticaercoides -infected arthropod intermediate hosts and a direct transmission path by the ingestion of embryonated eggs from the environment (Chabaud et al. 1962 ; Reitl et al. 2021 ). Remarkably, this nematode has been reported to infect at least two different chameleon species (i. e. F. pardalis and C. calyptratus ) without an involvement of intermediate hosts (Coke 1997 ). Referring to clinical signs, larval migration-driven subcutaneous swellings are commonly reported in chameleons (Pantchev and Beck 2013 ; Reitl et al. 2021 ). Concerning coccidian parasites in chameleons, the genus Isospora is represented by ten species, which have been identified in both wild- and captivity-kept animals (Brygoo 1963 ; Modrý et al. 1997 , 2000 , 2001a ; Abdel-Wasae 2004 ; McAllister 2012 ; Ekawasti et al. 2021). Moreover, nine tetra-sporocystic coccidian species (i. e. Eimeria spp. and Choleoeimeria spp.) have been characterized within the Chamaleonidae family (Sergent 1902 ; Modrý et al. 2001a , b; Sloboda and Modrý, 2006 ). In the current study, the coccidia prevalence (7.8%) was lower compared to oxyurids (14.1%). However, current necropsies revealed a higher level of mortality driven by coccidiosis (due to Isospora spp. or Choleoeimeria spp. infections) when compared to oxyuridosis indicating a higher pathogenicity of these protozoa. Hence, six animals exhibited severe intestinal mucosal damage caused by massive coccidian intracellular replication leading to a massive presence of meront and oocyst stages in the small intestine lumina. Here, coccidia prevalence reached 7.8%, a level similar to findings reported by Pantchev ( 2011 ) with 10.6%. In contrast, Biallas ( 2013 ) recorded a much higher prevalence with 21.7% in a smaller sample size. Eventually, this discrepancy may be explained by a higher proportion of juveniles in former study, the lack of prior antiparasitic treatment or other, unassessed factors like housing and hygiene conditions. As expected, juvenile chameleons (3.7%) showed a higher rate of coccidian infections than adults (2.5%) in the current study. This age-related susceptibility for intestinal coccidia infections is in line with previous reports from Biallas ( 2013 ) and Pantchev ( 2011 ) and is also commonly recorded for mammalian hosts (Foreyt 1990 ; Bangoura and Bardsley 2020 ). Interestingly, coccidiosis frequently occurred in co-infections. Hence, co-infections of Isospora spp. and oxyurids were the most common in the current study, thereby agreeing with findings of Biallas ( 2013 ) and re-confirming that these parasitic infections represent the two most prevalent ones in captive reptile husbandry, as both parasites have direct life cycles and short prepatent periods (Walden and Mitchell 2021 ; Pike et al. 2023 ). Of note, captive-bred chameleons like C. calyptratus and F. pardalis revealed a twice higher coccidia prevalence compared to their wild-caught counterparts; a phenomenon attributed to the aforementioned factors. Reptilian coccidiosis, like in mammals and birds, is a self-limiting disease (Long 1982 ; Quiroz-Castañeda and Dantán-González 2015 ; Hallinger et al. 2019 ). While typically being asymptomatic and self-limited in the wild, captivity settings allow for reinfections with varying degrees of severity, depending on various extrinsic and intrinsic factors (Mutschmann 2006 ; Bangoura and Bardsley 2020 ). Furthermore, infections with coccidian parasites have a higher likelihood of causing mortality and morbidity in captive lizards (Modrý and Sloboda 2004 ). During Isospora infections of the intestinal mucosa, a wide spectrum of clinical signs may be observed, ranging from watery to hemorrhagic diarrhea, to general symptoms like anorexia, lethargy, weight loss, cachexia, and, in some cases, even gut invagination and obstipation (Mutschmann 2006 ; Schneller and Pantchev 2008; Biallas 2013 ). In contrast, Choleoeimeria species exclusively parasitize the epithelial mucosa of the gallbladder, eventually causing epithelial cell destruction and resulting in gall bladder obstruction with similar symptoms as mentioned before (Modrý and Sloboda 2004 ). Most enteric flagellated protozoa bear low pathogenicity and are often referred to as opportunistic parasites since the development of related clinical symptoms is multifactorial and based on several parameters like stress, overcrowding, nutritional deficiencies, and host specificity (Zwart et al. 1984 ; Mutschmann 2006 ). Nonetheless, in the case of Leptomonas spp., bloody colitis was consistently reported in parasitized chameleons (Schneller and Pantchev 2008; Pantchev 2011 ). Overall, a potential role of feeder insects as paratenic hosts for Leptomonas spp. infections in chameleons has been proposed (Bayon 1915 ; Wenyon 1920 ) but remains unconfirmed. In the current study, most animals with leptomonosis suffered from enteritis but additionally showed co-infections with other parasites. Moreover, the presence of flagellated Leptomonas trophozoites in the gut lumen was mostly linked to intestinal inflammation in vivo . In the current study, Entamoeba stages were diagnosed in only one sample, thereby emphasizing the rareness of this infection in captive chameleons. Nevertheless, Entamoeba invadens is widely recognized for its high pathogenicity in snakes and lizards, but only a single case of amoebiasis has been reported in a chameleon co-infected with Mycobacterium simiae complex (Barnard and Upton 1994 ; Schneller and Pantchev 2008; Wellehan and Walden 2018 ; Fuentes et al. n. d.). Thus, the real impact of dysenteric amoebiasis in chameleons remains uncertain, rendering prevention essential for health sustainment (Barnard and Upton 1994 ; Hatt 2010 ). Pentastomids have been recorded in a wide range of reptile species, including snakes, geckos, agamas, skinks, varanids and crocodiles (Deakins 1973 ; Paré 2008 ; Pantchev and Tappe 2011 ; Rataj et al. 2011 ). Importantly, certain pentastomid genera like Armillifer and Porocephalus own zoonotic potential, especially those parasitizing snakes (Pantchev and Tappe 2011 ; Wellehan and Walden 2018 ; Hallinger et al. 2020 ). Lizards are the most common final hosts for the genus Raillietiella (Paré 2008 ; Barton 2009 ; Mendoza-Roldan et al. 2022 ), which can also infect chameleons (Sapion-Miranda et al. 2025 ; Hellebuyck et al. 2025 ). The zoonotic potential of R. orientalis has not been confirmed yet, however, one case report diagnosed Raillietiella sp. larvae in a human (Tappe et al. 2016). Even though reflecting a rare event, this case calls for further research on the zoonotic potential of pentastomids. These arthropod endoparasites own a highly complex, malunderstood lifecycle that may involve or not intermediate hosts (Bosch 1986 ). Cockroaches serve as intermediate hosts for Raillietiella spp., raising severe concerns on parasite transmission in regions where these synanthropic insects commonly occur in household or zoos (Lavoipierre and Lavoipierre 1965 ; Lavoipierre and Rajamanickam 1973 ; Ali et al. 1985 ; Bosch 1986 ; Palmisano et al. 2022 ). Notably, neither intestinal cestode adults, proglottids nor cestode eggs were found in this parasitological study. 5. Conclusion In recent years, keeping chameleons as exotic pets has become more and more popular. Their fascinating phenotype and the chance of housing in small-spaced terraria renders chameleons as attractive pets for apartment dwellers. However, responsible breeding practices and appropriate husbandry conditions are essential to maintain animal health and to control parasite infections. This includes a routine maintenance of clean enclosures, monitoring for parasite infections by regular coproscopy and the implementation of effective parasite control measures, whenever necessary. Regular medical checks are essential to maintain the overall health and resilience of captive chameleon populations. Moreover, it seems essential to monitor and control wild-caught reptile imports and to assess their infection status, especially with respect to zoonotic pathogens (e. g. pentastomids). In addition, the potential role of arthropods as mechanical vectors or intermediate hosts need to be further investigated to better understand their role in reptile parasite epidemiology. Overall, this study offers a base line data set that will provide not only veterinarians but also herpetologists with valuable insights into the neglected field of reptile medicine. Declarations Compliance with ethical standards Conflict of interest the authors declare that they have no conflict of interest. Funding Declaration this research did not receive funding. Ethical approval was not required for the study in accordance with the local and institutional requirements because the study involved naturally deceased animals and fecal samples. Fecal samples were submitted by owners and veterinarians and analyzed at exomed GmbH laboratories without any disturbance to the animals. Tissue samples and parasites were exclusively collected postmortem from deceased animals and examined at exomed GmbH. Ethics, Consent to Participate, and Consent to Publish declarations: not applicable. Author Contribution P.SM. wrote the main manuscript text and prepared the figures and tables. A.T. supervised, reviewed, and edited the manuscript. C.H. conceptualized, supervised, reviewed, and edited the manuscript. M.H. conceptualized, reviewed, and edited the manuscript. All authors reviewed the manuscript. Acknowledgement AcknowledgmentsThis research was supported by exomed GmbH Laboratory in Marburg, Germany, and the Institute of Parasitology of the Justus Liebig University in Giessen, Germany. Further, we would like to thank Dr. Alexandra Negro for kindly sharing some of the pictures of Malagasy chameleons. Data Availability The datasets generated and analysed during the current study are available as supplementary material accompanying this article. References Abdel-Wasae BM (2004) Isospora taizii (Apicomplexa: Eimeriidae), a new coccidian parasite from the Yemen chameleon ( Chamaeleo calyptratus ) (Sauria: Chamaeleonidae) in Taiz City, Yemen Republic. Assiut Univ Bull Environ Res 7(2):29–34. Ali JH, Riley J, Self JT (1985) A review of the taxonomy and systematics of the pentastomid genus Raillietiella Sambon, 1910 with a description of a new species. Syst Parasitol 7(2):111–123. https://doi.org/10.1007/BF00009814 Bangoura B, Bardsley KD (2020) Ruminant coccidiosis. 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Exp Parasitol 130(3):282–284. https://doi.org/10.1016/j.exppara.2012.01.011 Rom B, Kornas S, Basiaga M (2018) Endoparasites of pet reptiles based on coproscopic methods. Ann Parasitol 64(2). https://doi.org/10.17420/ap6402.142 Sapion-Miranda P, Ebmer D, Kniha E, et al. (2025) First identification of a patent pentastomid pulmonary ( Raillietiella orientalis ) infection in a captive Meller's chameleon ( Trioceros melleri ) in Germany. Int J Parasitol Parasites Wildl 26:101045. Satour NS, Dewir AW (2018) Some internal parasites of reptiles in Alexandria Province, Egypt. Egypt Vet Med Soc Parasitol J 14(1):98–114. https://doi.org/10.21608/evmspj.2018.33974 Schneller P, Pantchev N, Norden N (2008) Parasitology in snakes, lizards and chelonians: A husbandry guide. Edition Chimaira Frankfurt am Main. Scullion FT, Scullion MG (2009) Gastrointestinal protozoal diseases in reptiles. J Exot Pet Med 18(4):266–278. https://doi.org/10.1053/j.jepm.2009.09.004 Sergent ME (1902) Sur une coccidie nouvelle parasite du cameleon vulgaire. Comptes Rendus Soc Biol 54:1260–1261. Sloboda M, Modrý D (2006) New species of Choleoeimeria (Apicomplexa: Eimeriidae) from the veiled chameleon, Chamaeleo calyptratus (Sauria: Chamaeleonidae), with taxonomic revision of eimerian coccidia from chameleons. Folia Parasitol 53:91–97. https://doi.org/10.14411/fp.2006.012 Stacy BA (2021) Reptile necropsy techniques. In: Jacobson ER, Garner MM, eds. Infectious diseases and pathology of reptiles: Color atlas and text. 2nd ed. CRC Press. Stets O (2019) Parasites of panther chameleons ( Furcifer pardalis ) grown in captivity and brought from the wild. J Vet Med Biotechnol Biosaf 5:15–17. https://doi.org/10.36016/JVMBBS-2019-5-4-4 Széll Z, Sréter T, Varga I (2001) Ivermectin toxicosis in a chameleon ( Chamaeleo senegalensis ) infected with Foleyella furcata . J Zoo Wildl Med 32(1):115–117. https://doi.org/10.1638/1042-7260(2001)032[0115:ITIACC]2.0.CO;2 Tappe, D., Sulyok, M., Riu, T., R ́ozsa, L., Bod ́o, I., Schoen, C., Muntau, B., Babocsay, G., Hardi, R., 2016. Co-infections in visceral pentastomiasis, Democratic Republic of the Congo. Emerg. Infect. Dis. 22, 1333. The IUCN Red List of Threatened Species (2019) IUCN Red List of Threatened Species. https://www.iucnredlist.org/en. Accessed 01 Mar 2023. Toland E, Bando M, Hamers M, Cadenas V, Laidlaw R, Martínez-Silvestre A, van der Wielen P (2020) Turning negatives into positives for pet trading and keeping: A review of positive lists. Animals 10(12):2371. https://doi.org/10.3390/ani10122371 Walden M, Mitchell MA (2021) Pathogenesis of Isospora amphiboluri in bearded dragons ( Pogona vitticeps ). Animals 11(2):438. https://doi.org/10.3390/ani11020438 Wellehan JFX, Walden HDS (2018) Parasitology (including hemoparasites). In: Divers SJ, Stahl SJ, eds. Mader’s Reptile and Amphibian Medicine and Surgery. Elsevier Health Sciences. Wenyon CM (1920) Observations on the intestinal protozoa of three Egyptian lizards, with a note on a cell-invading fungus. Parasitology 12(4):350–365. https://doi.org/10.1017/S0031182000014347 Wilson SC, Carpenter JW (1996) Endoparasitic diseases of reptiles. Semin Avian Exot Pet Med 5(2):64–74. https://doi.org/10.1016/S1055-937X(96)80019-3 Wolf D, Vrhovec MG, Failing K, Rossier C, Hermosilla C, Pantchev N (2014) Diagnosis of gastrointestinal parasites in reptiles: Comparison of two coprological methods. Acta Vet Scand 56:44. https://doi.org/10.1186/s13028-014-0044-4 Zwart P, Teunis SFM, Cornelissen JMM (1984) Monocercomoniasis in reptiles. J Zoo Anim Med 15(3):129–134. Additional Declarations No competing interests reported. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-8902969","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":600009536,"identity":"adbe17d7-a611-4410-9814-7c06b0a487a4","order_by":0,"name":"Paula Sapion-Miranda","email":"data:image/png;base64,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","orcid":"","institution":"Justus Liebig University Giessen","correspondingAuthor":true,"prefix":"","firstName":"Paula","middleName":"","lastName":"Sapion-Miranda","suffix":""},{"id":600009537,"identity":"d3e3c7b3-92c6-4d5d-98f6-fbfd8c1f0ed9","order_by":1,"name":"Anja Taubert","email":"","orcid":"","institution":"Justus Liebig University Giessen","correspondingAuthor":false,"prefix":"","firstName":"Anja","middleName":"","lastName":"Taubert","suffix":""},{"id":600009539,"identity":"35b04fea-be69-4808-acea-212808cd3726","order_by":2,"name":"Carlos Hermosilla","email":"","orcid":"","institution":"Justus Liebig University Giessen","correspondingAuthor":false,"prefix":"","firstName":"Carlos","middleName":"","lastName":"Hermosilla","suffix":""},{"id":600009541,"identity":"32082e01-183b-4a41-b20f-e76a69fdec85","order_by":3,"name":"Malek J. Hallinger","email":"","orcid":"","institution":"exomed GmbH","correspondingAuthor":false,"prefix":"","firstName":"Malek","middleName":"J.","lastName":"Hallinger","suffix":""}],"badges":[],"createdAt":"2026-02-17 16:39:14","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-8902969/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-8902969/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":104181364,"identity":"e297aa76-5281-48e5-a62c-f20ec989e615","added_by":"auto","created_at":"2026-03-08 17:27:28","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":8994386,"visible":true,"origin":"","legend":"\u003cp\u003eExemplary images of different chameleon species investigated in this study \u003cstrong\u003ea. \u003c/strong\u003ea male Panther chameleon (\u003cem\u003eFurcifer pardalis\u003c/em\u003e), \u003cstrong\u003eb. \u003c/strong\u003ea male Parson’s chameleon (\u003cem\u003eCalumma parsonii\u003c/em\u003e), \u003cstrong\u003ec. \u003c/strong\u003ea male Malagasy giant chameleon (\u003cem\u003eFurcifer oustaleti\u003c/em\u003e), \u003cstrong\u003ed. \u003c/strong\u003ea female\u003cstrong\u003e \u003c/strong\u003eVeiled chameleon (\u003cem\u003eChamaelo calyptratus\u003c/em\u003e).\u003c/p\u003e","description":"","filename":"Fig1.png","url":"https://assets-eu.researchsquare.com/files/rs-8902969/v1/022697b2f8a08aef027adb6d.png"},{"id":104181360,"identity":"d40e91df-da3e-4d9d-a6d9-7a5a6af6b51b","added_by":"auto","created_at":"2026-03-08 17:27:28","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":1171709,"visible":true,"origin":"","legend":"\u003cp\u003eGeographical origin of captive chameleon fecal samples (\u003cem\u003en\u003c/em\u003e = 670) according to postcode districts in Germany.\u003c/p\u003e","description":"","filename":"Fig2.png","url":"https://assets-eu.researchsquare.com/files/rs-8902969/v1/705c9e0146eda2f77e2ca44a.png"},{"id":104181361,"identity":"4e722611-bfc5-46cd-becf-1eaacaa7927d","added_by":"auto","created_at":"2026-03-08 17:27:28","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":2809830,"visible":true,"origin":"","legend":"\u003cp\u003eExemplary photomicrographs of different parasitic stages found in chameleon faecal samples. \u003cstrong\u003ea.\u003c/strong\u003e operculated\u003cstrong\u003e \u003c/strong\u003eoxyurid\u003cstrong\u003e \u003c/strong\u003eegg (Veiled chameleon)\u003cstrong\u003e b. \u003c/strong\u003esporulated\u003cstrong\u003e \u003c/strong\u003e\u003cem\u003eIsospora \u003c/em\u003esp. oocyst containing two sporocysts with four sporozoites each (Veiled chameleon), \u003cstrong\u003ec.\u003c/strong\u003e flagellated\u003cem\u003e Leptomonas\u003c/em\u003e spp. trophozoites (Meller's chameleon), \u003cstrong\u003ed. \u003c/strong\u003etrematode egg with miracidium (Jackson's chameleon)\u003cstrong\u003e e. \u003c/strong\u003eembryonated pentastomid egg with larva (Malagasy giant chameleon), \u003cstrong\u003ef. \u003c/strong\u003e\u003cem\u003eHexametra angusticaecoides \u003c/em\u003eegg\u003cem\u003e \u003c/em\u003e(Panther chameleon).\u003c/p\u003e","description":"","filename":"Fig3.png","url":"https://assets-eu.researchsquare.com/files/rs-8902969/v1/2652bee5cb6681ac446845c9.png"},{"id":104404074,"identity":"75cf262c-b423-44c6-bc8e-ba1658627c8f","added_by":"auto","created_at":"2026-03-11 12:19:42","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":596658,"visible":true,"origin":"","legend":"\u003cp\u003eEndoparasite\u003cstrong\u003e \u003c/strong\u003emono- and co-infections in faecal samples of chameleons (\u003cem\u003en\u003c/em\u003e = 670) kept in captivity in Germany\u003c/p\u003e","description":"","filename":"Fig4.png","url":"https://assets-eu.researchsquare.com/files/rs-8902969/v1/a53a54d394ff526341f8ace5.png"},{"id":104181367,"identity":"9a267077-a1a8-488c-915d-05b06feaebed","added_by":"auto","created_at":"2026-03-08 17:27:29","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":6584857,"visible":true,"origin":"","legend":"\u003cp\u003eExemplary images from\u003cstrong\u003e \u003c/strong\u003echameleon necropsies and related histological findings. \u003cstrong\u003ea. \u003c/strong\u003eCoccidian meronts (black arrows) in the intestinal epithelium of a Veiled chameleon (400x magnification), \u003cstrong\u003eb. \u003c/strong\u003eclose-up of a dilated gallbladder (black arrow) caused by bile duct obstruction associated with a severe \u003cem\u003eCholeoeimeria\u003c/em\u003e sp. infection, \u003cstrong\u003ec. \u003c/strong\u003ecross section of a \u003cem\u003eSpinicauda \u003c/em\u003esp. nematode (black arrows) in the small intestine mucosa of a Panther chameleon (200x magnification),\u003cstrong\u003e d. \u003c/strong\u003eadult\u003cstrong\u003e \u003c/strong\u003e\u003cem\u003eHexametra angusticaecoides \u003c/em\u003enematode (indicated by black arrow)\u003cem\u003e \u003c/em\u003eremoved from the subcutaneous tissue, co-infected with \u003cem\u003eFoleyella furcata\u003c/em\u003e in the serosa, exposed after removal of the skin (asterisk) of a Panther chameleon.\u003c/p\u003e","description":"","filename":"Fig5.png","url":"https://assets-eu.researchsquare.com/files/rs-8902969/v1/0f1efd1d04afe7c381b020ea.png"},{"id":104409015,"identity":"49c5931d-6de8-410d-98e2-234e61ec8208","added_by":"auto","created_at":"2026-03-11 12:43:55","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":29393632,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8902969/v1/f4ec7a1c-8eeb-45f4-b83b-40bef78649c9.pdf"},{"id":104404368,"identity":"dcbb1fd7-187d-4bee-8cd2-c5efee9df645","added_by":"auto","created_at":"2026-03-11 12:20:08","extension":"xlsx","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":31448,"visible":true,"origin":"","legend":"","description":"","filename":"AD2.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-8902969/v1/5dcefbe4dc020e6b17f5f2bd.xlsx"},{"id":104181363,"identity":"0c76f5b1-b48f-48c3-aeb2-8c06ba3d3d87","added_by":"auto","created_at":"2026-03-08 17:27:28","extension":"xlsx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":168389,"visible":true,"origin":"","legend":"","description":"","filename":"AD1.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-8902969/v1/191c6a44610328949d433aab.xlsx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Gastrointestinal protozoan and helminth parasite infections in captive chameleons in Germany","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003eTo date, reptile husbandry is increasing in popularity since captive-bred reptiles can easily be obtained as exotic pets (Carpenter et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). However, not all reptiles are available from breeders, and enthusiasts seeking for exotic options may turn to wild-caught specimen with potentially unknown origin (Hallinger et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Marshall et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Toland et al. \u003cspan citationid=\"CR98\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). This ambiguity contributes to the introduction of numerous potentially invasive (neozoan) pathogens of these reptiles, including both ecto- and endoparasites (Rataj et al. \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2011\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBoth wild and captive reptiles are frequently infected with endoparasites, such as apicomplexans, flagellated protozoa, nematodes, cestodes, trematodes, and pentastomids (Frank \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e1985\u003c/span\u003e; Par\u0026eacute; \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Pasmans et al. \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Pantchev and Beck \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Hallinger et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). While many of these parasites typically own low pathogenicity in their original habitats, they can significantly impact the animal\u0026rsquo;s health if environmental conditions change (Wilson and Carpenter; \u003cspan citationid=\"CR102\" class=\"CitationRef\"\u003e1996\u003c/span\u003e). Hence, several risk factors like inadequate temperatures, stress, and high-density stocking affect the reptile's immune system (Bower et al.; \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). In this context, enhanced stress during trapping and transport of wild-caught reptiles often drives an increase in morbidity and mortality rates (Bower et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Stets \u003cspan citationid=\"CR92\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn captive reptiles, a notable prevalence of gastrointestinal parasitoses is common, especially referring to parasites with monoxenous lifecycles (Schneller and Pantchev 2008; Pantchev and Beck \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Hallinger et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). This phenomenon can be attributed to the high environmental tenacity of certain endoparasite stages (eggs) and short prepatencies of related parasites (Pasmans et al. \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Cervone et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Hallinger et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Distinct intrinsic factors, such as age, gender, genus and species or extrinsic factors like poor hygiene, stress and nutrition can significantly contribute to the development of clinical diseases in reptiles (Pasmans et al. \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Pantchev and Beck \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Hallinger et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Moreover, inadequate housing conditions create a perfect environment for high parasite burdens and frequent reinfections (Pasmans et al. \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Pantchev and Beck \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Wolf et al. \u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Hallinger et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Recent studies have suggested that feeding insects also represents a significant risk factor of endoparasite transmission, requiring further investigations (Gałęcki and Sok\u0026oacute;ł \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; M\u0026uuml;ller et. al \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eWithin lizards, chameleons (family Chamaelonidae) represent a distinctive and highly specialized clade of insectivorous reptiles. Chameleons, renowned for their attractive appearance, have been successfully bred in captivity for over four decades, while their trading dates to the early 1800s (Carpenter et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Diaz et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2015\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSimilar to other captive reptiles, chameleons often show a high prevalence of endoparasites, a health-compromising condition that has raised considerable concern among herpetologists (Okulewicz et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Rom et al. \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Despite this fact, there remains a gap of understanding on parasite transmission, parasite-driven pathogenesis and clinical manifestations (Nash et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Moreover, many chameleon species, such as the Belalanda chameleon (\u003cem\u003eFurcifer belalanda\u003c/em\u003e) from Madagascar or the Nguru spiny pygmy chameleon (\u003cem\u003eRhampholeon acuminatus\u003c/em\u003e) from Tanzania, fall into categories of critically endangered (4%) animals, others as endangered (21%), vulnerable (12.2%) or nearly threatened (18%), due to factors like habitat loss, illegal exotic pet trade and others (The IUCN Red List of Threatened Species \u003cspan citationid=\"CR97\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn the context of parasite infections, chameleon ectoparasitism seems relatively rare when compared to other reptiles, since no cases of infestations have been described so far in captive animals. In contrast, infestations with mites belonging to the family Trombiculidae have been described in wild chameleons (Pantchev \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Eckhardt et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). However, it is important to note that more detailed research is needed to clarify the occurrence of ectoparasite infestations in chameleons kept as pets.\u003c/p\u003e \u003cp\u003eIn Germany, there are only few studies on parasites affecting chameleons. Consistently, Biallas (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) conducted an extensive study on 212 chameleon faecal samples to investigate the prevalence and detrimental effects of endoparasites in captive chameleons, considering animal risk factors like origin, age and sex. Further, Pantchev (\u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) examined 66 chameleon faecal samples and provided an overview on most common parasite infections and related clinical symptoms. Stets (\u003cspan citationid=\"CR92\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) analyzed 646 feacal samples of Panther chameleons (\u003cem\u003eFurcifer pardalis\u003c/em\u003e), thereby comparing endoparasite infections in wild-caught and captive-kept animals in the Ukraine. Additionally, some case reports on distinct parasitoses and related treatments in captive chameleons (e. g. \u003cem\u003eF. pardalis, Chamaleo calyptratus\u003c/em\u003e) were recorded (Damiani et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Reitl et al. \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Heng et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Some reports documented endoparasitoses of wild-caught chameleons (Dillehay et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e1986\u003c/span\u003e; Lhermitte-Vallarino et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Heckmann \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Pellett et al. \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Collicutt et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Eckhardt et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) mostly focusing on parasite species with heteroxenous lifecycles. Occasionally, studies on reptile endoparasites also included individual findings on the most commonly kept chameleon species (\u003cem\u003eChamaelo calyptratus\u003c/em\u003e) (Wilson and Carpenter, \u003cspan citationid=\"CR102\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Goldberg et al. 2009; Rataj et al. \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Okulewicz et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Rom et al. \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Satour and Dewir \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), but lacking both large sample numbers and representative data.\u003c/p\u003e \u003cp\u003eIn general, the identification of pathogenic endoparasite species, including anthropozoonotic ones, is crucial for adequate targeted parasitological treatments, thereby minimizing both chameleon morbidity/mortality and transmission to pet owners (Pantchev \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2011\u003c/span\u003e, Mendoza-Roldan et al., \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Therefore, the current study aimed to provide novel insights into the occurrence of gastrointestinal endoparasites in captive-kept chameleons in Germany by investigating a substantial sample size (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;670) but also involving 25 different chameleon species to ensure a representative analysis. In addition to faecal analyses, we performed histopathological examinations on 31 deceased chameleons, providing a new baseline dataset on the presence of endoparasites in these popular herpetological pet animals.\u003c/p\u003e \u003cp\u003e \u003cb\u003eFigure 1.\u003c/b\u003e Exemplary images of different chameleon species investigated in this study \u003cb\u003ea.\u003c/b\u003e a male Panther chameleon (\u003cem\u003eFurcifer pardalis\u003c/em\u003e), \u003cb\u003eb.\u003c/b\u003e a male Parson\u0026rsquo;s chameleon (\u003cem\u003eCalumma parsonii\u003c/em\u003e), \u003cb\u003ec.\u003c/b\u003e a male Malagasy giant chameleon (\u003cem\u003eFurcifer oustaleti\u003c/em\u003e), \u003cb\u003ed.\u003c/b\u003e a female Veiled chameleon (\u003cem\u003eChamaelo calyptratus\u003c/em\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eNumber and origin of chameleon fecal samples (total \u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;670)\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"8\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eChameleon species\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCommon name\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eNo. examined\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colspan=\"3\" nameend=\"c8\" namest=\"c6\"\u003e \u003cp\u003eOrigin (private/ vet / zoo)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eFurcifer pardialis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003ePanther chameleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e250\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e223/16/11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eChamaeleo calyptratus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eVeiled chameleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e232\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e200/30/2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003eunknown species\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e103\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e84/14/5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eCalumma parsonii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eParson\u0026rsquo;s chameleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e12/0/11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTrioceros melleri\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMeller\u0026rsquo;s chameleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e11/0/1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eChamaeleo dilepis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFlap-necked chameleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e9/1/0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eFurcifer lateralis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCarpet chameleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e6/0/0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTrioceros jacksonii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eJackson\u0026rsquo;s chameleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e6/0/0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eFurcifer oustaleti\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMalagasy giant chamaleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e5/0/0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTrioceros hoehnelii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eH\u0026ouml;hnels chameleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e4/1/0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTriceros rudis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCoarse chameleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e3/0/0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eBradypodion thamnobates\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNatal Midlands dwarf chameleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2/0/0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eBradypodion fischeri\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFischer\u0026rsquo;s chameleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0/2/0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTrioceros Bitaeniatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSide-striped chameleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2/0/0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTrioceros j. xantholophus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eYellow-crested Jackson\u0026rsquo;s chameleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2/0/0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eFurcifer verrucosus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eWarty chameleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0/1/0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eKinyongia boehmei\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eB\u0026ouml;hme\u0026rsquo;s two-horned chamaleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1/0/0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eRampholeon acuminatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eNguru pygmy chameleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0/1/0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eBradypodion setaroi\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSetaro\u0026rsquo;s dwarf chameleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1/0/0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTrioceros johnstoni\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eJohnston\u0026rsquo;s chameleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0/1/0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTrioceros sternfeldi\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCrater Highlands side-striped chameleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1/0/0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTrioceros ellioti\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eElliot\u0026rsquo;s chameleon\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1/0/0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"1\" nameend=\"c8\" namest=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e"},{"header":"2. Materials and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003e2.1. Faecal samples\u003c/h2\u003e \u003cp\u003eIn total, 670 faecal samples of captive chameleons representing 25 species of different genera (e. g. \u003cem\u003eChameleo\u003c/em\u003e, \u003cem\u003eFurcifer\u003c/em\u003e, \u003cem\u003eTriocerus\u003c/em\u003e, \u003cem\u003eCalumma\u003c/em\u003e, \u003cem\u003eBradypodion\u003c/em\u003e, \u003cem\u003eRhampholeon\u003c/em\u003e, \u003cem\u003eKinyongia\u003c/em\u003e, see Fig.\u0026nbsp;1) were sent to exomed GmbH laboratory, Marburg, Germany (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) between 2012\u0026ndash;2019 and examined coprologically. Of these, 573 (85.8%) came from private owners, 67 (10%) from veterinary surgeons, and 30 (4.5%) from zoological gardens across Germany (Fig.\u0026nbsp;2). Each faecal sample was accompanied by a submission form stating the signalement of the animal (i. e. species, sex, weight, and age) and a short anamnesis (husbandry type, previous parasitological diagnoses, and antiparasitic treatments, see Online Resource 1).\u003c/p\u003e \u003cp\u003eTo identify different parasites and their stages, direct saline fecal smears (DSFS) were performed according to Cooper (\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). For DSFS preparation, faeces (10 g) were mixed at a ratio of 1:1 with 0.9% NaCl solution to a uniform suspension of the sample, placed on a microscope slide, covered with a coverslip (22 x 22mm; Nunc) and examined by a light microscope (100x or 400x magnification, Axio Imager M1\u0026reg;, Zeiss, Jena). Oxyurid, heterakid, ascarid, \u003cem\u003eStrongyloides\u003c/em\u003e, trematode and penstatomid eggs as well protozoan oocysts, cysts and trophozoites were identified based on previous description guidelines (Barnard and Upton \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Wilson and Carpenter \u003cspan citationid=\"CR102\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Schneller and Pantchev 2008; Pantchev \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Hallinger et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Faecal samples were listed as positive (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e), if at least one stage of the above described endoparasites posing a health-compromising risk for chameleons or other lizards was diagnosed. Samples lacking any pathogenic parasitic stages or showing non-pathogenic genera (e.g. \u003cem\u003eNyctotherus\u003c/em\u003e) as well as pseudoparasites, were here classified as negative.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003e2.2 Chameleon necropsies\u003c/h2\u003e \u003cp\u003eA total of 31 chameleon individuals (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;31) were necropsied at exomed GmbH laboratory (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). In these cases, the signalement (i. e. species, age, sex) and basic medical history, [e. g. clinical signs, husbandry, food, cause of death (natural death/euthanization by veterinary surgeons)] was reported (Online Resource 2). Additionally, tissue samples were collected and fixed in formalin for pathohistological examinations as previously described (Stacy \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). In detail, the chameleon necropsy was performed on its body side by an incision along the ventral midline from the cloaca to the forelimb and shoulder blade and up to the dorsal side. By this incision procedure, the coelomic cavity was exposed without obstructions (Farris et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Primarily, the gastrointestinal tract was visually inspected for the presence of metazoan endoparasites. Additionally, intestinal content was collected for DSFS. For metazoan endoparasite diagnosis, morphological identification was performed in accordance to Pantchev (\u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) and Hallinger et al. (\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) using a light microscope supplied with a digital camera (Axio Imager M1\u0026reg;, Zeiss, Jena).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003e2.3 Microbiological examination of necropsies\u003c/h2\u003e \u003cp\u003eFor further microbial identification, faeces and/or coelom swabs were taken from necropsied animals. For bacterial and fungal cultivation \u003cem\u003ein vitro\u003c/em\u003e, both the Columbia/MacConkey sheep blood agar (5%) as well as the Sabouraud dextrose agar (SDA) were used (BioMerieux, Charbonnier les Bains, France). In addition, bacterial isolates were identified by gram staining, oxidase, and catalase tests (BioMerieux, Charbonnier les Bains, France), together with a commercially available API 20 E/NE kit (BioMerieux, Charbonnier les Bains, France), as formerly described in reptiles (Marenzoni et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Hallinger et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cb\u003eFigure 2.\u003c/b\u003e Geographical origin of captive chameleon fecal samples (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;670) according to postcode districts in Germany.\u003c/p\u003e \u003c/div\u003e"},{"header":"3. Results","content":"\u003cp\u003e\u003cstrong\u003e3.1 Faecal samples\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eDSFS-based coproscopical analyses of pet chameleon faecal samples from Germany revealed a wide spectrum of gastrointestinal protozoan and metazoan parasites. In total, 33.2% (223/670) faecal samples proved positive for parasite infections. Among the positive samples from DSFS testing the following parasitic stages were diagnosed: oxyurid eggs (14.1%)(Fig. 3a), coccidian oocysts (7.8%)(Fig. 3b), flagellated protozoan trophozoites (6.3%)(Fig. 3c), heterakid eggs (3.57%), \u003cem\u003eStrongyloides\u003c/em\u003e spp. eggs (2.9%), thin-shelled nematode eggs (0.6%), trematode eggs (0.6%)(Fig. 3d), pentastomid eggs (0.3%)(Fig 3e), ascarid eggs (0.15%) (Fig. 3f), \u003cem\u003eEntamoeba\u003c/em\u003e cysts (0.15%) and \u003cem\u003ePhysalopteroides\u0026nbsp;\u003c/em\u003espp. eggs (0.15%) (please refer to Table 2). Neither cestode eggs nor \u003cem\u003eCryptosporidium\u0026nbsp;\u003c/em\u003eoocysts were found in the current faecal samples.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eA total of 9.8% of the 223 parasite-positive chameleon samples showed co-infections (Fig. 4). Among these, 4.5% (10/223) of animals exhibited an \u0026lsquo;coccidia-nematode\u0026rsquo; co-infection, while 3.6% (8/223) of chameleons had a \u0026lsquo;nematode-flagellated trophozoite\u0026rsquo; co-infection. Additionally, 1.7% (4/223) of animals showed an \u0026lsquo;coccidia-flagellated trophozoite\u0026rsquo; co-infection. Notably, in the faecal samples, no evidence of multiple infections of tri-, tetra- or higher-orders were detected.\u003c/p\u003e\n\u003cp\u003eOxyurid nematodes were the most prevalent metazoan parasites with 14.1%. Interestingly, the oxyurid prevalence varied with the chameleon genus, since it proved significantly higher in Veiled chameleons (\u003cem\u003eC. calyptratus\u003c/em\u003e: 64%) when compared to other chameleon species (36%) of this study (Chi-Square test:\u0026nbsp;\u0026chi;2=31.061, \u003cem\u003edf= 1, P\u0026nbsp;\u003c/em\u003e\u0026lt; 0.0001\u003cem\u003e)\u003c/em\u003e. Overall, no significant correlation between parasite infection and age (i. e. juvenile \u0026lt;1 year of age versus adult \u0026gt;1 year of age) was found. The same holds true for other factors, such as sex or origin, since no significant correlations were observed.\u003c/p\u003e\n\u003cp\u003eOther parasitic nematode stages like heterakid eggs (3.6%), embryonated \u003cem\u003eStrongyloides\u003c/em\u003e spp. eggs (2.9%), and thin-shelled nematode eggs (0.6%) were found at a lower prevalence. Only one sample proved positive for ascarid eggs (0.1%) and another for \u003cem\u003ePhysalopteroides\u0026nbsp;\u003c/em\u003espp. eggs (0.1%). Intestinal trematode eggs were found in 4 samples (0.6%) whilst pentastomid eggs were present in two faecal samples (0.3%). When considering intestinal protozoan parasites, these\u0026nbsp;revealed as the second most frequently diagnosed endoparasites. Among the protozoans, coccidia were the most prevalent in analyzed fecal samples. In this context, the genus \u003cem\u003eIsospora\u003c/em\u003e spp. showed the highest prevalence among the identified apicomplexans with 5.1% (34/670). whilst \u003cem\u003eIsospora jaracimmani\u0026nbsp;\u003c/em\u003e(1.2%), \u003cem\u003eCholeoeimeria\u003c/em\u003e spp. (0.3%) and un-sporulated apicomplexan oocysts (1.3%) were found at lower prevalences. Besides coccidia, \u003cem\u003eEntamoeba\u003c/em\u003e cysts (0.15%) and cysts of intestinal flagellates (0.9%) were also diagnosed. Among the flagellated protozoa, \u003cem\u003eLeptomonas\u003c/em\u003e spp. reached the highest prevalence (2.2%) followed by \u003cem\u003eTrichomonas\u003c/em\u003e spp. (1.5%) whereas other flagellates like \u003cem\u003eTritrichomonas\u003c/em\u003e spp. (1.1%), \u003cem\u003eProteromonas\u003c/em\u003e spp. (0.3%) and \u003cem\u003eMonocercomonas\u003c/em\u003e spp. (0.1%) ranked lower. We observed a statistical significant difference in the prevalence of coccidia between adult and juvenile animals since juveniles were more frequently affected (\u003cem\u003ep\u003c/em\u003e \u0026lt; 0.0009).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e3.2 Chameleon necropsies\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eA total of 31 animals originating from 10 different chameleons species were subjected to necropsy. Here, most animals (13/31, 41 %) belonged to the Panther chameleon (\u003cem\u003eF. pardalis\u003c/em\u003e), species (details of origin Table 1). In necropsies, 19 chameleons (61%) tested positive for endoparasite infections (Table 3). Pathological examination revealed that severe parasitoses signified the cause of death in at least 13 animals (40.6%), including isosporosis (\u003cem\u003en\u003c/em\u003e = 3; 9.3%) (Fig. 5a), choleoeimeriosis (n = 3; 9.3%) (Fig. 5b), heterakiosis (\u003cem\u003en\u003c/em\u003e = 2; 6.2%)(Fig. 5c), oxyuridiosis (\u003cem\u003en\u003c/em\u003e = 1;3.1%). An animal\u003cem\u003e\u0026nbsp;\u003c/em\u003e(3.1%) died due to a tetra-parasitic infection showing leptomonosis, trichomonosis, foleyellosis (\u003cem\u003eFoleyella furcata\u003c/em\u003e) and hexametrosis (Fig. 5d). One Jackson\u0026rsquo;s chameleon (\u003cem\u003eT.\u003c/em\u003e \u003cem\u003ejacksonii\u003c/em\u003e) died due to severe protozoa-driven enteritis caused by \u003cem\u003eLeptomonas\u0026nbsp;\u003c/em\u003espp. and \u003cem\u003eTrichomonas\u0026nbsp;\u003c/em\u003espp. infections. Furthermore, there were five cases of co-infections (\u003cem\u003en\u003c/em\u003e = 5, 15.6%).\u003c/p\u003e\n\u003cp\u003eOther etiological causes identified for chameleon death were gout (15.6%), mycosis (9.37%), salmonellosis (9.37%), other bacterial infections (9.37%), intestinal obstipation (6.25%), mechanical trauma (6.25%), adenocarcinoma (6.25%), malnutrition (3.1%) and hypervitaminosis D3 (3.1%). In two chameleons, the cause of death remained unknown even though showing enteritis and hepatitis (see Online Resource 2).\u0026nbsp;\u003cbr\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 2.\u0026nbsp;\u003c/strong\u003eParasitological findings in chameleon faecal samples [(total:\u003cem\u003e\u0026nbsp;n\u003c/em\u003e = 670; positive for parasite stages: 223/670 (33.3%);\u003cem\u003e\u0026nbsp;\u003c/em\u003enegative: 447/670 (66.7%)]\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" align=\"\" width=\"564\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 31.2057%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eParasite species\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 32.9787%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eNo. positive samples\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 35.8156%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eHost species\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 31.2057%;\"\u003e\n \u003cp\u003eOxyurid nematodes (Pharyngodonidae\u003cem\u003e)\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 32.9787%;\"\u003e\n \u003cp\u003e95/670 (14.1%)\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 35.8156%;\"\u003e\n \u003cp\u003e\u003cem\u003eC. calyptratus\u0026nbsp;\u003c/em\u003e(61)\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eF. pardalis\u0026nbsp;\u003c/em\u003e(16)\u0026nbsp;\u003c/p\u003e\n \u003cp\u003eUnknown species (15)\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eC. parsonii\u0026nbsp;\u003c/em\u003e(2)\u003c/p\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;R. acuminatus\u003c/em\u003e (1)\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 31.2057%;\"\u003e\n \u003cp\u003eCoccidia (e.g. \u003cem\u003eIsospora jaracimmani,\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eIsospora\u0026nbsp;\u003c/em\u003espp., \u003cem\u003eCholoerimeria\u0026nbsp;\u003c/em\u003espp.)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 32.9787%;\"\u003e\n \u003cp\u003e53/670 (7.8%)\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 35.8156%;\"\u003e\n \u003cp\u003e\u003cem\u003eF. pardalis\u0026nbsp;\u003c/em\u003e(25)\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eC. calyptratus\u0026nbsp;\u003c/em\u003e(21)\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003eUnknown species (7)\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eC. dilepsis\u0026nbsp;\u003c/em\u003e(2)\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eF. lateralis (1)\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eT. jacksonii\u0026nbsp;\u003c/em\u003e(1)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 31.2057%;\"\u003e\n \u003cp\u003eFlagellated protozoa (e.g. \u003cem\u003eLeptomonas\u0026nbsp;\u003c/em\u003espp., \u003cem\u003eTritrichomonas\u0026nbsp;\u003c/em\u003espp., \u003cem\u003eProteromonas\u003c/em\u003e spp.)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 32.9787%;\"\u003e\n \u003cp\u003e42/670 (6.3%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 35.8156%;\"\u003e\n \u003cp\u003e\u003cem\u003eF. pardalis\u0026nbsp;\u003c/em\u003e(13)\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eC. calyptratus\u0026nbsp;\u003c/em\u003e(12)\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eC. dilepis\u0026nbsp;\u003c/em\u003e(3)\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eT. melleri\u0026nbsp;\u003c/em\u003e(3)\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eC. parsonii\u0026nbsp;\u003c/em\u003e(2)\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eF. oustaleti\u0026nbsp;\u003c/em\u003e(1)\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eT. jacksonii\u0026nbsp;\u003c/em\u003e(1)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 31.2057%;\"\u003e\n \u003cp\u003eHeterakids\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 32.9787%;\"\u003e\n \u003cp\u003e24/670 (3.57%)\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 35.8156%;\"\u003e\n \u003cp\u003e\u003cem\u003eF. pardalis\u0026nbsp;\u003c/em\u003e(7)\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eC. parsonii\u0026nbsp;\u003c/em\u003e(7)\u003c/p\u003e\n \u003cp\u003eUnknown species (4)\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eF. oustaleti\u0026nbsp;\u003c/em\u003e(3)\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eC. calyptratus\u0026nbsp;\u003c/em\u003e(2)\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eB. thamnobates\u0026nbsp;\u003c/em\u003e(1)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 31.2057%;\"\u003e\n \u003cp\u003e\u003cem\u003eStrongyloides\u0026nbsp;\u003c/em\u003espp.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 32.9787%;\"\u003e\n \u003cp\u003e20/670 (2.98%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 35.8156%;\"\u003e\n \u003cp\u003e\u003cem\u003eC. calyptratus\u0026nbsp;\u003c/em\u003e(12)\u0026nbsp;\u003c/p\u003e\n \u003cp\u003eUnknown species (9)\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eF. pardalis\u0026nbsp;\u003c/em\u003e(5)\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eT. hoehnelii\u0026nbsp;\u003c/em\u003e(3)\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eT. rudis\u0026nbsp;\u003c/em\u003e(1)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 31.2057%;\"\u003e\n \u003cp\u003eThin-shelled nematode eggs\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 32.9787%;\"\u003e\n \u003cp\u003e4/670 (0.6%)\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 35.8156%;\"\u003e\n \u003cp\u003e\u003cem\u003eT. melleri\u0026nbsp;\u003c/em\u003e(1)\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eT. j. xantholophus\u0026nbsp;\u003c/em\u003e(1)\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eC. dilepis\u0026nbsp;\u003c/em\u003e(1) \u0026nbsp;\u003c/p\u003e\n \u003cp\u003eUnknown species (1)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 31.2057%;\"\u003e\n \u003cp\u003eTrematode eggs (Diginea)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 32.9787%;\"\u003e\n \u003cp\u003e4/670 (0.6%)\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 35.8156%;\"\u003e\n \u003cp\u003e\u003cem\u003eF. pardalis:\u0026nbsp;\u003c/em\u003e(3)\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eC. parsonii\u0026nbsp;\u003c/em\u003e(1)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 31.2057%;\"\u003e\n \u003cp\u003ePentastomids\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 32.9787%;\"\u003e\n \u003cp\u003e2/670 (0.30%)\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 35.8156%;\"\u003e\n \u003cp\u003e\u003cem\u003eF. pardalis:\u0026nbsp;\u003c/em\u003e(1)\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eF. oustaleti\u0026nbsp;\u003c/em\u003e(1)\u003cem\u003e\u0026nbsp; \u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 31.2057%;\"\u003e\n \u003cp\u003eAscarids\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 32.9787%;\"\u003e\n \u003cp\u003e1/670 (0.15%)\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 35.8156%;\"\u003e\n \u003cp\u003e\u003cem\u003eF. pardalis:\u0026nbsp;\u003c/em\u003e(1)\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 31.2057%;\"\u003e\n \u003cp\u003eAmoeba\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 32.9787%;\"\u003e\n \u003cp\u003e1/670 (0.15%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 35.8156%;\"\u003e\n \u003cp\u003e\u003cem\u003eF. pardalis:\u0026nbsp;\u003c/em\u003e(1)\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 31.2057%;\"\u003e\n \u003cp\u003ePhysalopteroidea\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 32.9787%;\"\u003e\n \u003cp\u003e1/670 (0.15%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 35.8156%;\"\u003e\n \u003cp\u003e\u003cem\u003eF. oustaleti:\u0026nbsp;\u003c/em\u003e(1)\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cstrong\u003eFigure 3.\u0026nbsp;\u003c/strong\u003eExemplary photomicrographs of different parasitic stages found in chameleon faecal samples. \u003cstrong\u003ea.\u003c/strong\u003e operculated\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003eoxyurid\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003eegg (Veiled chameleon)\u003cstrong\u003e\u0026nbsp;b.\u0026nbsp;\u003c/strong\u003esporulated\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003cem\u003eIsospora\u0026nbsp;\u003c/em\u003esp. oocyst containing two sporocysts with four sporozoites each (Veiled chameleon), \u003cstrong\u003ec.\u003c/strong\u003e flagellated\u003cem\u003e\u0026nbsp;Leptomonas\u003c/em\u003e spp. trophozoites (Meller\u0026apos;s chameleon), \u003cstrong\u003ed.\u0026nbsp;\u003c/strong\u003etrematode egg with miracidium (Jackson\u0026apos;s chameleon)\u003cstrong\u003e\u0026nbsp;e.\u0026nbsp;\u003c/strong\u003eembryonated pentastomid egg with larva (Malagasy giant chameleon), \u003cstrong\u003ef.\u0026nbsp;\u003c/strong\u003e\u003cem\u003eHexametra angusticaecoides\u0026nbsp;\u003c/em\u003eegg\u003cem\u003e\u0026nbsp;\u003c/em\u003e(Panther chameleon).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFigure 4.\u0026nbsp;\u003c/strong\u003eEndoparasite\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003emono- and co-infections in faecal samples of chameleons (\u003cem\u003en\u003c/em\u003e = 670) kept in captivity in Germany\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFigure 5.\u0026nbsp;\u003c/strong\u003eExemplary images from\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003echameleon necropsies and related histological findings. \u003cstrong\u003ea.\u0026nbsp;\u003c/strong\u003eCoccidian meronts (black arrows) in the intestinal epithelium of a Veiled chameleon (400x magnification), \u003cstrong\u003eb.\u0026nbsp;\u003c/strong\u003eclose-up of a dilated gallbladder (black arrow) caused by bile duct obstruction associated with a severe \u003cem\u003eCholeoeimeria\u003c/em\u003e sp. infection, \u003cstrong\u003ec.\u0026nbsp;\u003c/strong\u003ecross section of a \u003cem\u003eSpinicauda\u0026nbsp;\u003c/em\u003esp. nematode (black arrows) in the small intestine mucosa of a Panther chameleon (200x magnification),\u003cstrong\u003e\u0026nbsp;d.\u0026nbsp;\u003c/strong\u003eadult\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003cem\u003eHexametra angusticaecoides\u0026nbsp;\u003c/em\u003enematode (indicated by black arrow)\u003cem\u003e\u0026nbsp;\u003c/em\u003eremoved from the subcutaneous tissue, co-infected with \u003cem\u003eFoleyella furcata\u003c/em\u003e in the serosa, exposed after removal of the skin (asterisk) of a Panther chameleon.\u003cstrong\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 3.\u0026nbsp;\u003c/strong\u003eParasitological findings in chameleon necropsies [(total:\u003cem\u003e\u0026nbsp;n\u003c/em\u003e = 31; positive for parasite stages: 19/31 (61.2%)]\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"869\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 21.7241%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eParasite Species\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 18.5057%;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 26.092%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eNo. of positive (%)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 21.7241%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eHost species n.\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 2.18391%;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 7.58621%;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 2.18391%;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"2\" valign=\"top\" style=\"width: 40.2299%;\"\u003e\n \u003cp\u003eFlagellated protozoa (e.g. \u003cem\u003eLeptomonas\u0026nbsp;\u003c/em\u003espp., \u003cem\u003eTritrichomonas\u0026nbsp;\u003c/em\u003espp., \u003cem\u003eProteromonas\u003c/em\u003e spp.)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 26.092%;\"\u003e\n \u003cp\u003eTotal: 6 (19.3)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"4\" valign=\"top\" style=\"width: 33.6782%;\"\u003e\n \u003cp\u003e\u003cem\u003eF. pardalis\u0026nbsp;\u003c/em\u003e(3) \u003cem\u003eC. calyptratus\u0026nbsp;\u003c/em\u003e(1)\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eT. melleri\u0026nbsp;\u003c/em\u003e(1)\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eT. jacksonii xantholopus\u0026nbsp;\u003c/em\u003e(1)\u003c/p\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"2\" valign=\"top\" style=\"width: 40.2299%;\"\u003e\n \u003cp\u003e\u003cem\u003eIsospora\u0026nbsp;\u003c/em\u003espp.\u003cstrong\u003e\u003cem\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;\u0026nbsp;\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 26.092%;\"\u003e\n \u003cp\u003eTotal: 5 (16.1)\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"4\" valign=\"top\" style=\"width: 33.6782%;\"\u003e\n \u003cp\u003e\u003cem\u003eC. calyptratus\u0026nbsp;\u003c/em\u003e(1)\u003cem\u003e\u0026nbsp;F. pardalis\u0026nbsp;\u003c/em\u003e(3)\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eT. melleri\u0026nbsp;\u003c/em\u003e(1)\u003c/p\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"2\" valign=\"top\" style=\"width: 40.2299%;\"\u003e\n \u003cp\u003eOxyurid nematodes (Pharyngodonidae\u003cem\u003e)\u003c/em\u003e\u003cstrong\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 26.092%;\"\u003e\n \u003cp\u003eTotal: 4 (12.9)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"4\" valign=\"top\" style=\"width: 33.6782%;\"\u003e\n \u003cp\u003e\u003cem\u003eC. calyptratus\u0026nbsp;\u003c/em\u003e(4)\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"2\" valign=\"top\" style=\"width: 40.2299%;\"\u003e\n \u003cp\u003e\u003cem\u003eCholeoeimeria\u0026nbsp;\u003c/em\u003espp.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 26.092%;\"\u003e\n \u003cp\u003eTotal: 3 (9.6)\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"4\" valign=\"top\" style=\"width: 33.6782%;\"\u003e\n \u003cp\u003e\u003cem\u003eF. pardalis\u0026nbsp;\u003c/em\u003e(1)\u003c/p\u003e\n \u003cp\u003e\u003cem\u003eT. melleri\u0026nbsp;\u003c/em\u003e(2)\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"2\" valign=\"top\" style=\"width: 40.2299%;\"\u003e\n \u003cp\u003eHeterakids (e.g. \u003cem\u003eSpinicauda\u0026nbsp;\u003c/em\u003espp.)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 26.092%;\"\u003e\n \u003cp\u003eTotal: 2 (6.4)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"4\" valign=\"top\" style=\"width: 33.6782%;\"\u003e\n \u003cp\u003e\u003cem\u003eF. pardalis\u003c/em\u003e (1)\u003cem\u003e\u0026nbsp;B. thamnobates\u0026nbsp;\u003c/em\u003e(1)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"2\" valign=\"top\" style=\"width: 40.2299%;\"\u003e\n \u003cp\u003eHexametra angusticercoida\u003cstrong\u003e\u003cem\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;\u0026nbsp;\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 26.092%;\"\u003e\n \u003cp\u003eTotal: 1 (3.2)\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"4\" valign=\"top\" style=\"width: 33.6782%;\"\u003e\n \u003cp\u003e\u003cem\u003eF. pardalis\u0026nbsp;\u003c/em\u003e(1)\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"2\" valign=\"top\" style=\"width: 40.2299%;\"\u003e\n \u003cp\u003eFoleyella furcata\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 26.092%;\"\u003e\n \u003cp\u003eTotal: 1 (3.2)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"4\" valign=\"top\" style=\"width: 33.6782%;\"\u003e\n \u003cp\u003e\u003cem\u003eF. pardalis\u0026nbsp;\u003c/em\u003e(1)\u003c/p\u003e\n \u003cp\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"2\" valign=\"top\" style=\"width: 40.2299%;\"\u003e\n \u003cp\u003eStrongyloides sp.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 26.092%;\"\u003e\n \u003cp\u003eTotal: 1(3.2)\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"4\" valign=\"top\" style=\"width: 33.6782%;\"\u003e\n \u003cp\u003e\u003cem\u003eCalumma parsonii\u0026nbsp;\u003c/em\u003e(1)\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e"},{"header":"4. Discussion","content":"\u003cp\u003eThe present large-scale epidemiological study on gastrointestinal parasites of chameleons kept in captivity in Germany reveals a high diversity of protozoan and metazoan endoparasite species, thereby confirming findings from previous prevalence studies (Pantchev \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Biallas \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eReferring to chameleon oxyurid species, \u003cem\u003eParapharyngodon kenyaensis\u003c/em\u003e (Goldberg and Bursey \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), \u003cem\u003ePharyngodon dimorpha\u003c/em\u003e and \u003cem\u003eThelandros meridionalis\u003c/em\u003e (Brygoo \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1963\u003c/span\u003e) have been so far described in wild- and captivity-kept chameleons (Okulewicz et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Eckhardt et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Stets \u003cspan citationid=\"CR92\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). In the current study, oxyurid infections represented the most frequently found parasitosis (14.1%). However, earlier publications reported on a considerably higher prevalence (27.3% in Pantchev \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2011\u003c/span\u003e and 25.5% in Biallas \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). This might rely on an increasing awareness of owners leading to a more frequent testing for gastrointestinal parasite infections and subsequent treatment in the last years. Among the various chameleon species assessed for oxyuridosis, we observed a notable difference in prevalence. Hence, \u003cem\u003eC. calyptratus\u003c/em\u003e showed a significantly higher oxyurid prevalence than \u003cem\u003eF. pardalis\u003c/em\u003e and less commonly kept exotic chameleon species. This finding may be explained by its overrepresentation in the sample pool due to its broad availability and low purchase costs, unfortunately often resulting in suboptimal husbandry conditions given by unexperienced owners. In contrast, other, more expensive species like the Panther chameleon or more exotic taxa are typically kept by experienced owners, who may also be more aware on infection monitoring measures.\u003c/p\u003e \u003cp\u003eIn contrast to captivity, the chances of reinfections are lower in the wild, and a delicate equilibrium between reptile hosts and their oxyurid species has been described. Thus, certain oxyurid nematodes efficiently support the intestinal microbiome to keep the animal in balance (Schneller and Pantchev 2008; Hallinger et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Wellehan and Walden \u003cspan citationid=\"CR100\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Eckhardt et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). In captivity, however, chameleons are confined to limited space and often suffer from suboptimal husbandry conditions, disrupting this equilibrium and leading to massive infections with these monoxenous tenacious parasites, finally resulting in (sub)clinical disease (Schneller and Pantchev 2008; Rinaldi et al. \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Pantchev and Beck \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Hallinger et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Oxyuridosis in chameleons is usually of low pathogenicity, though severe infections can cause malabsorption, weight loss and nonspecific signs, especially in young, gravid or immunocompromised animals (Pantchev \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Biallas \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Pantchev and Beck \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Hallinger et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Kubiak \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In the current study, oxyurid mono-infection was restricted to solely one necropsied Veiled chameleon, however this parasitosis was accompanied by bacterial co-infections with \u003cem\u003eSalmonella enterica\u003c/em\u003e and \u003cem\u003eStenotrophomonas maltophila\u003c/em\u003e. Consequently, it is challenging to attribute the cause of death solely to oxyuridosis. This assumption aligns well with former findings on low intrinsic pathogenicity of oxyurids, reporting that mono-infections did not reveal as single cause of death (Biallas \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cem\u003eStrongyloides\u003c/em\u003e spp. nematodes are widespread parasites affecting captive and wild animals, including chameleons (Biallas \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). The current study revealed a \u003cem\u003eStrongyloides\u003c/em\u003e spp. prevalence of 2.9% in chameleons, which is consistent with previous findings (1.5%, Pantchev \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2009\u003c/span\u003e, \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). \u003cem\u003eStrongyloides\u003c/em\u003e infections may cause both respiratory and gastrointestinal signs due to larval migration and can be fatal (Schneller and Pantchev 2008; Biallas \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Heng et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe prevalence of heterakids and ascarids in the current study relatively low, when compared to formerly recorded prevalences of 6\u0026ndash;7% in chameleons (Pantchev \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Biallas \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). \u003cem\u003eSpinicauda sonsinoi\u003c/em\u003e (Brygoo \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1963\u003c/span\u003e) and \u003cem\u003eS. inglisi\u003c/em\u003e (Chabaud and Brygoo \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e1962\u003c/span\u003e) are the most common heterakid species in Malagasi chameleons. These parasites show a rather low pathogenicity if the parasitic burden is low (Schneller and Pantchev 2008). However, ascarid species like \u003cem\u003eHexametra angusticaercoides\u003c/em\u003e bear high pathogenic potential and severe hexametriosis often leads to animal death (Schneller and Pantchev 2008; Pantchev \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Barton et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Reitl et al. \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Of note, \u003cem\u003eH anguisticaercoides\u003c/em\u003e has a dual transmission mode in chameleons, an indirect infection route via consumption of \u003cem\u003eH. angusticaercoides\u003c/em\u003e-infected arthropod intermediate hosts and a direct transmission path by the ingestion of embryonated eggs from the environment (Chabaud et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1962\u003c/span\u003e; Reitl et al. \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Remarkably, this nematode has been reported to infect at least two different chameleon species (i. e. \u003cem\u003eF. pardalis\u003c/em\u003e and \u003cem\u003eC. calyptratus\u003c/em\u003e) without an involvement of intermediate hosts (Coke \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1997\u003c/span\u003e). Referring to clinical signs, larval migration-driven subcutaneous swellings are commonly reported in chameleons (Pantchev and Beck \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Reitl et al. \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eConcerning coccidian parasites in chameleons, the genus \u003cem\u003eIsospora\u003c/em\u003e is represented by ten species, which have been identified in both wild- and captivity-kept animals (Brygoo \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1963\u003c/span\u003e; Modr\u0026yacute; et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e1997\u003c/span\u003e, \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2000\u003c/span\u003e, \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2001a\u003c/span\u003e; Abdel-Wasae \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; McAllister \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Ekawasti et al. 2021). Moreover, nine tetra-sporocystic coccidian species (i. e. \u003cem\u003eEimeria\u003c/em\u003e spp. and \u003cem\u003eCholeoeimeria\u003c/em\u003e spp.) have been characterized within the Chamaleonidae family (Sergent \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e1902\u003c/span\u003e; Modr\u0026yacute; et al. \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2001a\u003c/span\u003e, b; Sloboda and Modr\u0026yacute;, \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). In the current study, the coccidia prevalence (7.8%) was lower compared to oxyurids (14.1%). However, current necropsies revealed a higher level of mortality driven by coccidiosis (due to \u003cem\u003eIsospora\u003c/em\u003e spp. or \u003cem\u003eCholeoeimeria\u003c/em\u003e spp. infections) when compared to oxyuridosis indicating a higher pathogenicity of these protozoa. Hence, six animals exhibited severe intestinal mucosal damage caused by massive coccidian intracellular replication leading to a massive presence of meront and oocyst stages in the small intestine lumina. Here, coccidia prevalence reached 7.8%, a level similar to findings reported by Pantchev (\u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) with 10.6%. In contrast, Biallas (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) recorded a much higher prevalence with 21.7% in a smaller sample size. Eventually, this discrepancy may be explained by a higher proportion of juveniles in former study, the lack of prior antiparasitic treatment or other, unassessed factors like housing and hygiene conditions. As expected, juvenile chameleons (3.7%) showed a higher rate of coccidian infections than adults (2.5%) in the current study. This age-related susceptibility for intestinal coccidia infections is in line with previous reports from Biallas (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) and Pantchev (\u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) and is also commonly recorded for mammalian hosts (Foreyt \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Bangoura and Bardsley \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eInterestingly, coccidiosis frequently occurred in co-infections. Hence, co-infections of \u003cem\u003eIsospora\u003c/em\u003e spp. and oxyurids were the most common in the current study, thereby agreeing with findings of Biallas (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) and re-confirming that these parasitic infections represent the two most prevalent ones in captive reptile husbandry, as both parasites have direct life cycles and short prepatent periods (Walden and Mitchell \u003cspan citationid=\"CR99\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Pike et al. \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Of note, captive-bred chameleons like \u003cem\u003eC. calyptratus\u003c/em\u003e and \u003cem\u003eF. pardalis\u003c/em\u003e revealed a twice higher coccidia prevalence compared to their wild-caught counterparts; a phenomenon attributed to the aforementioned factors.\u003c/p\u003e \u003cp\u003eReptilian coccidiosis, like in mammals and birds, is a self-limiting disease (Long \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e1982\u003c/span\u003e; Quiroz-Casta\u0026ntilde;eda and Dant\u0026aacute;n-Gonz\u0026aacute;lez \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Hallinger et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). While typically being asymptomatic and self-limited in the wild, captivity settings allow for reinfections with varying degrees of severity, depending on various extrinsic and intrinsic factors (Mutschmann \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Bangoura and Bardsley \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Furthermore, infections with coccidian parasites have a higher likelihood of causing mortality and morbidity in captive lizards (Modr\u0026yacute; and Sloboda \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). During \u003cem\u003eIsospora\u003c/em\u003e infections of the intestinal mucosa, a wide spectrum of clinical signs may be observed, ranging from watery to hemorrhagic diarrhea, to general symptoms like anorexia, lethargy, weight loss, cachexia, and, in some cases, even gut invagination and obstipation (Mutschmann \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Schneller and Pantchev 2008; Biallas \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). In contrast, \u003cem\u003eCholeoeimeria\u003c/em\u003e species exclusively parasitize the epithelial mucosa of the gallbladder, eventually causing epithelial cell destruction and resulting in gall bladder obstruction with similar symptoms as mentioned before (Modr\u0026yacute; and Sloboda \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2004\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eMost enteric flagellated protozoa bear low pathogenicity and are often referred to as opportunistic parasites since the development of related clinical symptoms is multifactorial and based on several parameters like stress, overcrowding, nutritional deficiencies, and host specificity (Zwart et al. \u003cspan citationid=\"CR104\" class=\"CitationRef\"\u003e1984\u003c/span\u003e; Mutschmann \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). Nonetheless, in the case of \u003cem\u003eLeptomonas\u003c/em\u003e spp., bloody colitis was consistently reported in parasitized chameleons (Schneller and Pantchev 2008; Pantchev \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Overall, a potential role of feeder insects as paratenic hosts for \u003cem\u003eLeptomonas\u003c/em\u003e spp. infections in chameleons has been proposed (Bayon \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1915\u003c/span\u003e; Wenyon \u003cspan citationid=\"CR101\" class=\"CitationRef\"\u003e1920\u003c/span\u003e) but remains unconfirmed. In the current study, most animals with leptomonosis suffered from enteritis but additionally showed co-infections with other parasites. Moreover, the presence of flagellated \u003cem\u003eLeptomonas\u003c/em\u003e trophozoites in the gut lumen was mostly linked to intestinal inflammation \u003cem\u003ein vivo\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eIn the current study, \u003cem\u003eEntamoeba\u003c/em\u003e stages were diagnosed in only one sample, thereby emphasizing the rareness of this infection in captive chameleons. Nevertheless, \u003cem\u003eEntamoeba invadens\u003c/em\u003e is widely recognized for its high pathogenicity in snakes and lizards, but only a single case of amoebiasis has been reported in a chameleon co-infected with \u003cem\u003eMycobacterium simiae\u003c/em\u003e complex (Barnard and Upton \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Schneller and Pantchev 2008; Wellehan and Walden \u003cspan citationid=\"CR100\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Fuentes et al. n. d.). Thus, the real impact of dysenteric amoebiasis in chameleons remains uncertain, rendering prevention essential for health sustainment (Barnard and Upton \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1994\u003c/span\u003e; Hatt \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2010\u003c/span\u003e).\u003c/p\u003e \u003cp\u003ePentastomids have been recorded in a wide range of reptile species, including snakes, geckos, agamas, skinks, varanids and crocodiles (Deakins \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e1973\u003c/span\u003e; Par\u0026eacute; \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Pantchev and Tappe \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Rataj et al. \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Importantly, certain pentastomid genera like \u003cem\u003eArmillifer\u003c/em\u003e and \u003cem\u003ePorocephalus\u003c/em\u003e own zoonotic potential, especially those parasitizing snakes (Pantchev and Tappe \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Wellehan and Walden \u003cspan citationid=\"CR100\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Hallinger et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Lizards are the most common final hosts for the genus \u003cem\u003eRaillietiella\u003c/em\u003e (Par\u0026eacute; \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Barton \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Mendoza-Roldan et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), which can also infect chameleons (Sapion-Miranda et al. \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Hellebuyck et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The zoonotic potential of \u003cem\u003eR. orientalis\u003c/em\u003e has not been confirmed yet, however, one case report diagnosed \u003cem\u003eRaillietiella\u003c/em\u003e sp. larvae in a human (Tappe et al. 2016). Even though reflecting a rare event, this case calls for further research on the zoonotic potential of pentastomids. These arthropod endoparasites own a highly complex, malunderstood lifecycle that may involve or not intermediate hosts (Bosch \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1986\u003c/span\u003e). Cockroaches serve as intermediate hosts for \u003cem\u003eRaillietiella\u003c/em\u003e spp., raising severe concerns on parasite transmission in regions where these synanthropic insects commonly occur in household or zoos (Lavoipierre and Lavoipierre \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e1965\u003c/span\u003e; Lavoipierre and Rajamanickam \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e1973\u003c/span\u003e; Ali et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e1985\u003c/span\u003e; Bosch \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1986\u003c/span\u003e; Palmisano et al. \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eNotably, neither intestinal cestode adults, proglottids nor cestode eggs were found in this parasitological study.\u003c/p\u003e"},{"header":"5. Conclusion","content":"\u003cp\u003eIn recent years, keeping chameleons as exotic pets has become more and more popular. Their fascinating phenotype and the chance of housing in small-spaced terraria renders chameleons as attractive pets for apartment dwellers. However, responsible breeding practices and appropriate husbandry conditions are essential to maintain animal health and to control parasite infections. This includes a routine maintenance of clean enclosures, monitoring for parasite infections by regular coproscopy and the implementation of effective parasite control measures, whenever necessary. Regular medical checks are essential to maintain the overall health and resilience of captive chameleon populations. Moreover, it seems essential to monitor and control wild-caught reptile imports and to assess their infection status, especially with respect to zoonotic pathogens (e. g. pentastomids). In addition, the potential role of arthropods as mechanical vectors or intermediate hosts need to be further investigated to better understand their role in reptile parasite epidemiology. Overall, this study offers a base line data set that will provide not only veterinarians but also herpetologists with valuable insights into the neglected field of reptile medicine.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eCompliance with ethical standards\u003c/h2\u003e\n\u003cp\u003eConflict of interest the authors declare that they have no conflict of interest.\u003c/p\u003e\n\u003ch2\u003eFunding Declaration this research did not receive funding.\u003c/h2\u003e\n\u003cp\u003eEthical approval was not required for the study in accordance with the local and institutional requirements because the study involved naturally deceased animals and fecal samples. Fecal samples were submitted by owners and veterinarians and analyzed at exomed GmbH laboratories without any disturbance to the animals. Tissue samples and parasites were exclusively collected postmortem from deceased animals and examined at exomed GmbH.\u003c/p\u003e\n\u003ch2\u003eEthics, Consent to Participate, and Consent to Publish declarations:\u0026nbsp;\u003c/h2\u003e\n\u003cp\u003enot applicable.\u003c/p\u003e\n\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\n\u003cp\u003eP.SM. wrote the main manuscript text and prepared the figures and tables. A.T. supervised, reviewed, and edited the manuscript. C.H. conceptualized, supervised, reviewed, and edited the manuscript. M.H. conceptualized, reviewed, and edited the manuscript. All authors reviewed the manuscript.\u003c/p\u003e\n\u003ch2\u003eAcknowledgement\u003c/h2\u003e\n\u003cp\u003eAcknowledgmentsThis research was supported by exomed GmbH Laboratory in Marburg, Germany, and the Institute of Parasitology of the Justus Liebig University in Giessen, Germany. Further, we would like to thank Dr. Alexandra Negro for kindly sharing some of the pictures of Malagasy chameleons.\u003c/p\u003e\n\u003ch2\u003eData Availability\u003c/h2\u003e\n\u003cp\u003eThe datasets generated and analysed during the current study are available as supplementary material accompanying this article.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAbdel-Wasae BM (2004) \u003cem\u003eIsospora taizii\u003c/em\u003e (Apicomplexa: Eimeriidae), a new coccidian parasite from the Yemen chameleon (\u003cem\u003eChamaeleo calyptratus\u003c/em\u003e) (Sauria: Chamaeleonidae) in Taiz City, Yemen Republic. 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University Park Press, Baltimore, USA, 1982, pp. 502-508\u003c/li\u003e\n\u003cli\u003eMaia JP, Crottini A, Harris DJ (2014) Microscopic and molecular characterization of \u003cem\u003eHepatozoon domerguei\u003c/em\u003e (Apicomplexa) and \u003cem\u003eFoleyella furcata\u003c/em\u003e (Nematoda) in wild endemic reptiles from Madagascar. Parasite 21:47. https://doi.org/10.1051/parasite/2014046\u003c/li\u003e\n\u003cli\u003eMarenzoni ML, Zicavo A, Veronesi F, et al. (2015) Microbiological and parasitological investigation on chelonians reared in Italian facilities. Vet Ital 51(3):173\u0026ndash;178. https://doi.org/10.12834/VetIt.7.21.3\u003c/li\u003e\n\u003cli\u003eMarshall BM, Strine C, Hughes AC (2020) Thousands of reptile species threatened by under-regulated global trade. 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J Zoo Anim Med 15(3):129\u0026ndash;134.\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"parasitology-research","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"pare","sideBox":"Learn more about [Parasitology Research](http://link.springer.com/journal/436)","snPcode":"436","submissionUrl":"https://submission.nature.com/new-submission/436/3","title":"Parasitology Research","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"chameleons, parasites, exotic pets, reptile medicine, wildlife","lastPublishedDoi":"10.21203/rs.3.rs-8902969/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8902969/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eWithin lizards, chameleons (family Chamaeleonidae) represent a distinctive and highly specialized clade of insectivorous reptiles. Chameleons have become more popular as reptile pets in recent years worldwide, due to their feasible husbandry and attractiveness. To date, not only reptile pet numbers but also health-compromising endoparasite infections are rising. In the current study, we analyzed a total of 670 faecal samples of 25 different chameleon species kept as pet animals all over Germany. In total, 223 animals (33%) were found to harbor gastrointestinal parasites. In total, 10 different parasite taxa were diagnosed in chameleon fecal samples. The most prevalent parasite stages were oxyurid eggs (14.1%), followed by coccidia oocysts (7.8%) (e. g. \u003cem\u003eIsospora jacarimanni\u003c/em\u003e, \u003cem\u003eIsospora\u003c/em\u003e spp., \u003cem\u003eCholeoeimeria\u003c/em\u003e spp., \u003cem\u003eEimeria\u003c/em\u003e spp.), flagellated protozoan trophozoites (6.3%) (e. g. \u003cem\u003eLeptomonas\u003c/em\u003e spp., \u003cem\u003eTritrichomonas\u003c/em\u003e spp., \u003cem\u003eProteromonas\u003c/em\u003e spp.), heterakid eggs (3.6%), thin-shelled nematode eggs (0.6%), digenean trematode eggs (0.6%), pentastomid eggs (0.3%), ascarid eggs (0.2%), \u003cem\u003eEntamoeba\u003c/em\u003e spp. cysts (0.2%) and physalopteroid eggs (0.2%). Among current chameleon species, veiled chameleons (\u003cem\u003eChamaleo calyptratus\u003c/em\u003e) showed a significant higher prevalence of oxyurid infections (64%) than other chameleon species. Co-infections were diagnosed in 9.8% (22/223) of faecal samples. When performing necropsies, 19/31 chameleons showed parasite-induced pathologies based on histopathological examinations, including one animal being infected with 4 parasite species (\u003cem\u003eLeptomonas\u003c/em\u003e spp., \u003cem\u003eTritrichomonas\u003c/em\u003e spp., \u003cem\u003eFoleyella furcata\u003c/em\u003e and \u003cem\u003eHexametra angusticaecoides)\u003c/em\u003e, thereby most likely impacting animal\u0026rsquo;s wellfare.\u003c/p\u003e","manuscriptTitle":"Gastrointestinal protozoan and helminth parasite infections in captive chameleons in Germany","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-03-08 17:27:23","doi":"10.21203/rs.3.rs-8902969/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2026-03-30T18:24:21+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-03-26T03:40:15+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-03-23T19:29:48+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"325442381912361043257266873646081623038","date":"2026-03-05T19:09:18+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"207145758267184167125217004947584458286","date":"2026-03-03T12:56:52+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-03-02T09:05:15+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-02-24T04:17:45+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2026-02-24T03:47:38+00:00","index":"","fulltext":""},{"type":"submitted","content":"Parasitology Research","date":"2026-02-17T16:29:13+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"parasitology-research","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"pare","sideBox":"Learn more about [Parasitology Research](http://link.springer.com/journal/436)","snPcode":"436","submissionUrl":"https://submission.nature.com/new-submission/436/3","title":"Parasitology Research","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"4464bed1-daae-4a75-ad4b-caac8d03e08f","owner":[],"postedDate":"March 8th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-05-16T13:38:48+00:00","versionOfRecord":[],"versionCreatedAt":"2026-03-08 17:27:23","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8902969","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8902969","identity":"rs-8902969","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}