Table 4.

Taenia spp. and Vesrsteria sp. cysticercoses and Taenia spp. coenuroses in non-human primates.

Taenia sp., case no., citation	Primate species, patient, origin, and case history	Pathological findings	Etiological diagnosis: morphology and DNA analysis (PCR, sequencing)a
Taenia crassiceps Case 1, Baer and Scheidegger (1946)	Hamadryas baboon (Papio hamadryas) “no individual data available”, lived in a group in the Zoo Basel, Switzerland, together with a couple of foxes.The animal was necropsied after a history of tetraplegia.	Necropsy. Larval stages subcutaneously and in the smooth muscles, penetrating to the retroperitoneal cavity, with parasite infiltration into the spinal cord.	Morphology. Diagnosis based on number (n = 30–35) and size (large 182–200, small 138 < 150 μm) of rostellar hooks and budding of the cysticerci. DNA analysis. Nd.
T. crassiceps Case 2, Dyer and Greve (1998)	Adult female black lemur (Eulemur macaco macaco) with a clinical history of lethargy, anorexia, and depression of a week's duration was submitted for necropsy. The animal was housed outdoors in a group.	Necropsy. Fluctuant swelling measuring 10 cm by 6 cm on the left back. A well-defined cystic structure contained hundreds of ellipsoidal to spherical cysts of 1–4 mm in diameter. Infiltration of both the peritoneal and pleural cavities by large numbers of cysticerci were found replacing approximately 90% of the left lung.	Morphology. Cysticerci with calcareous corpuscles, a scolex with a rostellum with hooks, spinous tegument, and exogenous budding. DNA analysis. Nd.
T. crassiceps Case 3, Young et al. (2000)	6-year-old female red ruffed lemur (Varecia variegata rubra) with 2 days history of lethargy and anorexia was presented with a large, fluctuant subcutaneous swelling extending from the dorsal aspect of the skull to the ventral cervical region.	Radiographs of the cervical region showed the mass containing discrete areas of mineralization. Surgical exploration “revealed a multiloculated mass with each individual cyst-like structure containing hundreds of bead-like nodules (<1 mm)”.	Morphology. A wet-prep of the nodules revealed cestode larvae. Histopathology. A single mass was described as granulomatous cellulitis with intralesional larval cestodes. DNA analysis. Nd.
T. crassiceps Case 4, Luzón et al. (2010)	5-year-old male, ring-tailed lemur (Lemur catta) in the Madrid Zoo-Aquarium, Spain, presented a large swelling in its abdomen (15 cm in length × 10 cm in width × 4 cm in height) and was anesthetized for a complete medical evaluation.	US image of the swelling suggested a septated subcutaneous mass. Exploratory surgery revealed an irregular fibrous cystic structure, containing numerous small transparent vesicles (ca. 3 mm in diameter) and after the peritoneal cavity was opened more vesicles were extirpated.	Morphology. Based on the scolex and hooks and budding was observed. DNA analysis. cox1 [P1] and nd1 [P3]: Identities 99–100% with corresponding published T. crassiceps sequences.
T. crassiceps Case 5, Alić et al. (2017)	15-year-old female ring-tailed lemur (L. catta) from Sarajevo Zoo, Bosnia and Herzegovinawas was presented after sudden death	Necropsy. “large multicystic structure, subdivided with fibrous septa and filled with numerous translucent, oval to ellipsoid bladder-like cysts, almost completely replacing right lung lobe “.	Morphology. Cysticerci with single inverted scolex, a rostelum with two rows of hooks and exogenous buds. DNA analysis. cox1 [P2]: Identity 880/880bp, 100% with T. crassiceps (GenBank AF216699, AB033411).
T. crassiceps Case 6, Bleyer et al. (2018)	28-year-old female, captive-born Nilgiri langur (Semnopithecus johnii) from a German zoo, with a right ankle joint age-related myxosarcoma developed an edematous swelling of the left thigh and also suffered from lethargy and anorexia. Animal was euthanized because of poor general condition.	Necropsy. Skeletal muscle of the left thigh had been replaced by a multilocular cystic mass containing numerous sand-grain–sized whitish structures; similar cysts were also present in the lung and the myocardium.	Morphology. Based on hook morphology. DNA analysis. cox1 [P1]: Identities 98–100% (450 bp) with T. crassiceps sequences.
Taenia serialis Case 1, Schwartz (1927)	Gelada baboon (Theropithecus gelada) from the National Zoological Park at Washington, D. C, with a large subcutaneous tumor in the right thoracic region.	Necropsy. Large subcutaneous tumor (1.5 kg). The majority of the mature cysts contained several hundred scoleces.	Morphology based on larval hooks and of adult worms from experimentally infected dogs. DNA analysis. Nd.
T. serialis Case 2, Sandground (1937)	Juvenile (<2 years) spotted-nose monkey (Cercopithecus nictitans) imported from the West African Coast to the USA as a pet animal. Slight torticollis for 3–4 weeks, soft swelling at the back of the head. Died.	Necropsy.Many discolored bodies (1–2 cm) attached to mesentery. Scalp with bone erosion and protrusion of a mass of translucent, white cysts.	Morphology. Microscopical examination corresponds to Coenurus serialis. DNA analysis. Nd.
T. serialis Case 3, Elek and Finkelstein (1939)	4-year-old gelada baboon (T. gelada), borne in captivity in Germany and imported to USA (zoological park). Spastic paralysis in right lower extremity before death.	Necropsy. Multiple subcutaneous nodules in upper and lower extremities (1–12 cm), and a large intra-abdominal cystic mass infiltrating underlying muscles.	Morphology. Diagnosis of cysts (no microscopic details provided). DNA analysis. Nd.
T. serialis Case 4, Clark (1969)	Male gelada baboon (T. gelada), transported between two zoological parks in the USA. Nodular enlargement on right thigh. Euthanized because surgery was not successful.	Necropsy. Subcutaneous nodular enlargement (11–25 cm) with small ulcerations.	Morphology. Diagnosis of cysts (no microscopic details provided). Infection of 2 dogs was not successful. DNA analysis. Nd.
T. serialis Case 5, Schneider-Crease et al. (2013)	13-year-old male, wild Ethiopian gelada baboon (T. gelada) from a national park. Tissue was extracted from a protuberant coenurus on the left ventral forelimb.	Small to large swellings in various parts of the animal's body.	Morphology. Not reported. DNA analysis.Its2 [P9] and 12S [P5]: 99% and 99% identity with T. serialis sequences.
T. multiceps (probably misdiagnosed) Leith and Satterfield (1974)	6-year-old female gelada baboon (T. gelada) from a zoological park in the USA. Animal, which was imported from Ethiopia with 2 years, presented multiple subcutaenous cysts.	Necropsy. Multilocular easily ruptured cystic masses in the left masseter and temporal muscle region. Further subcutaneous cysts in the scapular region extending into the abdomen.	Morphology. Size, number, and shape of hooklets provided by the author, but inconclusive according to Table 5. DNA analysis. Nd.
T. braunib Railliet and Marullaz (1919)	Rhesus monkey (Macaca mulatta), kept in France but unknown origin, infected with Leishmania tropica, died in experiment.	Necropsy. Abdominal tumor at left perineum (2.7–5 cm).	Morphology. Microscopical measurements performed.b DNA analysis. Nd.
Taenia brauni Fain (1956)	Monkey (Cercopithecus mitis doggetti), Rwanda	Cysts in brain (parietal lobe; 2 cm), heart (apex; 1 cm), and parotid gland	Morphology. Performed but no details provided by the author. DNA analysis. Nd.
Taenia sp. coenurosis Lau et al. (1973)	Whitehanded gibbon (Hylobates lar) at zoological park in the USA. Left eye progressing gradually for 3–4 weeks. Retrobulbar mass at radiography. Euthanasia.	Cyst behind the left eyeball.	Morphology. Size and shape analysis of hooklets inconclusive. DNA analysis. Nd.
Taenia martis De Liberato et al. (2014)	18-year-old male ring-tailed lemur (L. catta), immunocompetent, lives in a zoo in Rome (Italy). Apathy, loss of appetite, abdominal distension and diarrhoea were observed 10 days before death.	Severe exudative fibrinous-purulent peritonitis with numerous adhesions between the abdominal wall and the bowel loops. After intestine removal, two free and viable, 4 cm long, whitish, leaf-like parasitic forms were pinpointed.	Morphology. Macroscopic examination of the two parasites allowed their identification as larval stages of cestodes. DNA analysis. cox1 [P1]: Identity 98% with T. martis sequences.
T. martis Brunet et al. (2014)	3-years-old subadult mail tonkean macaque (Macaca tonkeana), immunocompetent, born and raised at Strasbourg University Centre of Primatology, France.	Abdominal mass (±10 cm × 5 cm) was detected at palpation without clinical signs.	Morphology. Cysticerci and rostellar hooks were most close to the T. martis description. DNA analysis. cox1 [P1]: Identity 382/383 bp or 99.7% with T. martis sequence (GenBank AB731758). nd1 [P3]: Identity 438/439 or 99.8% with EU544606.
Versteria sp. Goldberg et al. (2014)	Juvenile (sex?) Bornean orangutan (Pongo pygmaeus), captive born in Colorado, USA, (no indications of immunosuppression). Loss of appetite and intermittent, moist cough; became increasingly lethargic and was found dead after 2 days.	Necropsy. Diffuse hemorrhages in the lungs, splenomegaly, a pale mottled liver, and thoracic and pericardial effusions. Histopathology of the liver revealed cystic structures containing eukaryotic parasite cells. Cause of death: “acute respiratory distress due to disseminated infection with an unknown parasite”.	Morphology. Not conclusive. DNA analysis. Deep sequencing identified a Taenia spp. 12S [P7] and cox1/nd1 [P1/P3] phylogenetic trees placed the organisme within Versteria sp. (Cestoda: Taeniidae).
Taenia hydatigena Hobbs et al. (2003)	5-year-old male rhesus macaque (M. mulatta), born and raised in a primate colony in China and imported 15 months earlier to the Oregon National Primate Research Centre. An abdominal mass was discovered during routine physical examination.	Necropsy. A pale yellow cyst attached to the greater omentum containing 500 ml of flocculent yellow fluid.	Morphology. Rostellar hooks, the large size of the cysticercus, and its location within the peritoneal cavity were consistent with the diagnosis of a Taenia hydatigena cysticercosis. DNA analysis. Nd.
T. hydatigena Tsubota et al. (2009)	5-years-old male long-tailed macaque (Macaca fascicularis), born and raised in a primate research colony in China in a toxicity study in Osaka (Japan). The monkey showed no clinical signs and was sacrificed at the end of a dosing experiment.	Necropsy. A yellow cyst filled with more than 100 ml of pale yellow fluid was found in the abdominal cavity.	Morphology. A cysticercus with a well developed scolex was found in the cyst. DNA analysis. nd1 [P3]: Identities of 96.7%–98.5% with T. hydatigena sequences.
Taenia sp. Wolff et al. (1989)	Captive-born 1-year-old male red ruffed lemur (V. variegata rubra) with unremarkable routine quarantine examination with radiography (zoo, USA).	Necropsy. Extrapleural calcified larval cestode in the left ventro-caudal thorax and pulmonary nodule with a cysticercus in the left dorso-caudal lung.	Morphology. C ysticercus with armed scolex; species identification not done. DNA analysis. Nd.

Nd: Not done or no data given.

^aPrimer pairs used are given in squared brackets and refer to Table 6.

^bDiagnosed as Multiceps ramosus or M. lemuris by the author, which are considered to be a synonym for T. brauni (Loos-Frank, 2000).