Kamila Cajiao-Mora

Proctocaecum coronarium and Proctocaecum diploporum (Digenea: Cryptogonimidae): Morphologically- and phylogenetically-distinct but closely related and sympatric crocodilian trematodes with identical life cycles

We describe Catadiscus marielaosornae Cajiao-Mora and Bullard n. sp. (Diplodiscidae) from the intestine of Peters’ thin-toed frog, Leptodactylus petersii (Steindachner) (Anura: Leptodactylidae) from the Yahuarcaca Lake System, Colombia. We assigned our specimens to Catadiscus Cohn, 1904 because they have a smooth-surfaced, pyriform body, centrally-constricted ventro-terminal acetabulum that lacks an accessory sucker, pharynx with extramural sacs, oesophageal bulb, and single testis. The new species differs from all but Catadiscus rodriguezi Caballero, 1955, Catadiscus marinholutzi Freitas and Lent, 1939, and Catadiscus propinquus Freitas and Dobbin, 1956 by having a vitellarium that is confluent anteriorly (vs. two non-confluent fields). It differs from the aforenamed congeners by having a ventral common genital pore that opens anterodextral to the oesophageal bifurcation, round vitelline follicles that become confluent anterior to the testis, and a 1: 2–3 ratio of acetabulum: testis width. Our 28S phylogenetic analysis recovered the new species sister to C. marinholutzi, and Diplodiscidae as paraphyletic, with Catadiscus sister to a clade comprising a sequence ascribed to Chiorchis fabaceus (Diesing, 1838) Fischoeder, 1901 (Cladorchiidae), and sequences of three species of Diplodiscus Diesing, 1836. The species description herein includes several features pertaining to the acetabulum, female genitalia, and excretory system that could comprise useful character states with which to more readily classify and diagnose diplodiscid species and genera. To our knowledge, this is the first record of a digenean infecting Peters’ thin-toed frog, and it is also the first record of any parasite infecting any ditch frog (Leptodactylus spp.) from Colombia.

The synonymies of the echinochasmid genera Mesorchis Dietz, 1909 and Monilifer Dietz, 1909 with Stephanoprora Odhner, 1902 remain contentious and unresolved with morphology. To explore the matter, we herein provide a supplemental description of the type species of Stephanoprora, Stephanoprora ornata Odhner, 1902, based on specimens we collected from the intestine of a Nile crocodile Crocodylus niloticus Laurenti, 1768 (Crocodylia: Crocodylidae) captured in the Kavango River, Namibia. No nucleotide information was available previously for S. ornata. Morphology plus 28S and ITS2 phylogenetic analyses suggested Stephanoprora is a monotypic genus that can be differentiated from other genera by having 26 collar spines. Stephanoprora differs from Mesorchis by the number and distribution of collar spines (26 [2 dorsal spines, 12 lateral spines, 12 corner spines] vs. 22 [2 dorsal spines, 12 lateral spines, 8 corner spines] in Mesorchis), length of the pre-pharyngeal oesophagus, pharynx position posterior to collar (vs. pre-pharyngeal oesophagus short, pharynx anterior to or at level of corner spines), and testes shape (elongate-ovoid, irregular in outline vs. ovoid, atypically elongated). Our 28S and ITS2 phylogenies recovered the new sequence of S. ornata (having 26 collar spines) sister to all remaining echinochasmid sequences, representing species that have 20–24 collar spines. Echinochasmus Dietz, 1909 and Mesorchis were recovered as paraphyletic. We retain Monilifer as a junior subjective synonym of Echinochasmus based a suite of morphological features related to body shape and genitalia and because the designated type for Monilifer was reassigned to Echinochasmus. We accept 25 species of Mesorchis.

We propose and describe Alobophora sandrae Cajiao-Mora & Bullard n. gen., n. sp. (Digenea: Caballerotrematidae) for specimens we collected from arapaima, Arapaima gigas sensu lato (Osteoglossiformes: Arapaimidae) in the Amazon River near Leticia, Colombia. Alobophora differs from Caballerotrema Prudhoe, 1960 by lacking head collar projections and by having clustered corner spines and a narrow head collar (4–5× wider than pharynx), whereas Caballerotrema has head collar projections, lacks clustered corner spines, and has a broad head collar (7–8× wider than pharynx). We reassign Caballerotrema annulatum (Diesing, 1850) Ostrowski de Núñez & Sattmann, 2002 to the new genus, as Alobophora annulata (Diesing, 1850) Cajiao-Mora and Bullard n. comb., and provide a supplemental description of Caballerotrema brasiliense Prudhoe, 1960 based on specimens we collected from arapaima. We also examined the holotype and a paratype of Caballerotrema piscicola (Stunkard, 1960) Kostadinova & Gibson, 2001 and concluded that C. piscicola is a junior subjective synonym of C. brasiliense. Our 28S phylogeny recovered A. sandrae sister to A. annulata, with that clade sister to a clade comprising C. brasiliense and an innominate species of Caballerotrema. Caballerotrematidae was recovered sister to Echinostomatidae. We also provide a dichotomous key to caballerotrematids based on head collar projections, corner spine arrangement, proportional pharynx and head collar breadth, testes shape and arrangement, body surface spine shape and distribution, vitellarium distribution, and abundance of prostatic cells.

Herein, we provide a supplemental description of Caballerotrema annulatum (Diesing, 1850) Ostrowski de Núñez and Sattmann, 2002 (Digenea: Caballerotrematidae Tkach, Kudlai, and Kostadinova, 2016) based on specimens collected from the intestine of an electric eel, Electrophorus cf. varii (Gymnotiformes: Gymnotidae) captured in the Amazon River (Colombia). This caballerotrematid can be differentiated from its congeners by the following combination of morphological features: body surface spines forming contiguous transverse rows, concentric (wrapping dorso-ventrally around body), distributing into posterior body half (vs. restricted to anterior body half in Caballerotrema brasiliense Prudhoe, 1960; indeterminate for Caballerotrema aruanense Thatcher, 1980 and Caballerotrema piscicola [Stunkard, 1960] Kostadinova and Gibson, 2001); head collar lacking projections (vs. having them in C. brasiliense, C. aruanense, and C. piscicola), narrow (head collar more narrow than maximum body width vs. the head collar being obviously wider than the body in C. brasiliense, C. aruanense, and C. piscicola); corner spines clustered (vs. corner spines distributing as 2 separated pairs in C. brasiliense, C. aruanense, and C. piscicola); pharynx approximately at level of the corner spines (vs. pharynx far anterior to corner spines in C. brasiliense, C. aruanense, and C. piscicola); and testes ovoid and nonoverlapping (C. aruanense; vs. sinuous and overlapping in C. brasiliense and C. piscicola). Based on our results, we revise the diagnosis of Caballerotrema Prudhoe, 1960 to include features associated with the shape and distribution of body surface spines, orientation and position of head collar spines, cirrus sac, seminal vesicle, oviduct, Laurer's canal, oötype, vitellarium, and transverse vitelline ducts. We performed Bayesian inference analyses using the partial large subunit ribosomal (28S) DNA gene. Our 28S sequence of C. annulatum was recovered sister to that of Caballerotrema sp. (which is the only other caballerotrematid sequence available in GenBank) from an arapaima, Arapaima gigas (Schinz, 1822) (Osteoglossiformes: Arapaimidae) in the Peruvian Amazon. Our sequence of C. annulatum comprises the only caballerotrematid sequenced tethered to a morphological description and a voucher specimen in a lending museum. The present study is a new host record and new locality record for C. annulatum. The phylogeny comprises the most resolved and taxon-rich evolutionary hypothesis for Echinostomatoidea published to date.