DESCRIPTION OF FEMALE DENDRITOBILHARZIA PULVERULENTA (BRAUN, 1901) SKRJABIN, 1924 FROM TWO NEW AVIAN HOSTS IN NAMIBIA WITH PHYLOGENETIC ANALYSES AND COMMENTS ON SEVERAL TAXONOMICALLY UNCERTAIN AVIAN SCHISTOSOME SEQUENCES.

During a 2021 parasitological survey of birds in the Nyae Nyae-Khaudum Dispersal Area (Kavango-Zambezi Transfrontier Conservation Area, Namibia), we collected 9 specimens of Dendritobilharzia pulverulenta (Braun, 1901) Skrjabin, 1924 infecting the blood (heart lumen) of a white-backed duck, Thalassornis leuconotus (Eyton, 1838) (Anseriformes: Anatidae), and a fulvous whistling duck, Dendrocygna bicolor (Vieillot, 1816) (Anatidae). These flukes were fixed for morphology and preserved for DNA extraction. We assigned our specimens to DendritobilharziaSkrjabin and Zakharow, 1920 because they were strongly dorso-ventrally flattened in both sexes and had an intestinal cyclocoel with a zig-zag common cecum with lateral dendritic ramifications, numerous testes posterior to the cyclocoel and flanking the dendritic ramifications, and a tightly compacted convoluted ovary as well as lacking an oral sucker, ventral sucker, and gynaecophoric canal. Further, our specimens were morphologically identical to previously published descriptions of D. pulverulenta. Sequences of the 28S from our specimens were nearly identical to those identified as D. pulverulenta from North America (New Mexico), and our 28S phylogenetic analysis recovered D. pulverulenta within a polytomy of other Gigantobilharziinae spp. The CO1 phylogenetic analysis recovered a monophyletic Dendritobilharzia and, with low taxon sampling, a monophyletic Gigantobilharzia. This is the first record of a species of Dendritobilharzia infecting these ducks as well as the first record of an adult Dendritobilharzia from sub-Saharan Africa. The original description of adult D. pulverulenta (type locality: northern Sudan) was based on 2 males only, and hence the present study is the first description of female D. pulverulenta from Africa (the continent of the type locality). We reassign 2 Gigantobilharziinae spp. based on morphology and nucleotide evidence: Gigantobilharzia ensenadense (Lorenti, Brant, Gilardoni, Diaz, and Cremonte, 2022) Dutton and Bullard, n. comb., and Gigantobilharzia patagonense (Lorenti, Brant, Gilardoni, Diaz, and Cremonte, 2022) Dutton and Bullard, n. comb. We also comment on several avian schistosome sequences whose identities need confirmation or that likely have been misidentified.

First report of a fish blood fluke from sub-Saharan Africa: Nomasanguinicola dentata (Paperna, 1964) Warren and Bullard, 2023 infecting African sharptooth catfish, Clarias gariepinus (Burchell, 1822) Teugles, 1982 in the Kavango River, Namibia, and a revised phylogeny for Sanguinicolidae Poche, 1926.

We herein provide a supplemental description of Nomasanguinicola dentata (Paperna, 1964) Warren and Bullard, 2023 (Digenea: Sanguinicolidae) and provide a revised 28S phylogeny to test relationships among freshwater fish blood flukes. We examined the heart of three African sharptooth catfish, Clarias gariepinus (Burchell, 1822) Teugles, 1982 from the Kavango River (northeastern Namibia) that was infected with adults of N. dentata. This blood fluke differs from N. canthoensis by having a body 5.3-6.7 longer than wide (vs. 3.5-4.6), an anterior esophageal swelling 7-8% (vs. 14-24%) of total esophageal length, a posterior esophageal swelling 3-5% (vs. 8-10%) of total esophageal length, a pre-cecal (vs. wholly post-cecal) testis, and an ovary that does not extend laterally beyond the nerve cords. The 28S sequence for N. dentata differed from that of N. canthoensis by 144 bp (9% difference). The phylogenetic analysis recovered these species as sister taxa and Sanguinicolidae as monophyletic. This is the first report of a fish blood fluke from sub-Saharan Africa, and the first report of a species of Nomasanguinicola from Africa in ∼40 yrs.

NEW GENUS AND SPECIES OF CYCLOCOELIDAE STOSSICH, 1902 (PLATYHELMINTHES: DIGENEA) INFECTING THE NASOPHARYNGEAL CAVITY OF CANADA GOOSE, BRANTA CANADENSIS (ANSERIFORMES: ANATIDAE) FROM WESTERN ALABAMA.

While surveying the parasites of birds associated with western Alabama aquaculture ponds, we collected several specimens of Anativermis normdroneni n. gen., n. sp. (Digenea: Cyclocoelidae) from the nasopharyngeal cavity of a Canada goose, Branta canadensis (Linnaeus, 1758) (Anseriformes: Anatidae). These flukes were heat killed and fixed in neutral buffered formalin for morphology or preserved in 95% ethanol for DNA extraction. Anativermis resembles Morishitium (Witenberg, 1928) by having testes that are spheroid with smooth margins and located in the posterior quarter of the body, an anterior testis that is lateral to the midline and abuts the respective cecum, a posterior testis that is medial (testes diagonal) and abuts the cyclocoel, a genital pore that is immediately postpharyngeal, and a vitellarium that is discontinuous posteriorly. The new genus differs from Morishitium and is unique among all other cyclocoelid genera by having the combination of a body that is broadest in the anterior body half, a posterior body end that is more sharply tapered than the anterior body end, an ovary that nearly abuts the posterior testis, a vitellarium that is asymmetrical and distributes from the area immediately posterior to the cecal bifurcation posteriad to approximately the level of the ovary, and uterine loops extending dorsolateral to the ceca and filling the space between the ceca and the respective body margin for nearly the entire body length. The new genus was recovered as a distinct lineage in separate 28S, 18S, and ITS2 phylogenetic analyses. This is the first report of a cyclocoelid infecting the Canada goose and of a cyclocoelid from Alabama.

FIRST RECORD OF A PARASITE, DACTYLOGYRUS CF. SKRJABINI (MONOGENOIDEA: DACTYLOGYRIDAE), INFECTING INVASIVE SILVER CARP, HYPOPHTHALMICHTHYS MOLITRIX (VALENCIENNES, 1844) (CYPRINIFORMES: XENOCYPRIDIDAE) IN NORTH AMERICA.

The parasites infecting invasive carps in North America (all Cypriniformes: Xenocyprididae: grass carp, Ctenopharyngodon idella [Valenciennes, 1844]; silver carp, Hypophthalmichthys molitrix [Valenciennes, 1844]; bighead carp, Hypophthalmichthys nobilis [Richardson, 1845]; and black carp, Mylopharyngodon piceus [Richardson, 1846]) are little studied, and no parasite has been reported from silver carp there. We herein surveyed silver carp from Barkley Reservoir and Cheatham Reservoir (Cumberland River, Tennessee; June and December 2021) and the White River (Arkansas; May 2022) and collected numerous monogenoid specimens infecting the pores on the outer face of the gill raker plate. We heat-killed, formalin-fixed, and routinely stained some specimens for morphology and preserved others in 95% ethanol for DNA extraction and sequencing of the large subunit ribosomal DNA (28S). We identified our specimens as Dactylogyrus cf. skrjabini because they had a dorsal anchor deep root that is much longer than the superficial root, an approximately parallel penis and accessory piece, and a relatively large marginal hook pair V. No type specimen of Dactylogyrus skrjabiniAkhmerov, 1954 (type host and locality is silver carp, Amur River, Russia) is publicly available, but we borrowed several vouchers (NSMT-Pl 6393) that infected the gill rakers of silver carp captured in the Watarase River, Japan. The original description of D. skrjabini was highly stylized and diagrammatical, differing from the specimens we studied from North America and Japan by the dorsal anchor having a superficial root and shaft that comprise a strongly C-shaped hook (the superficial root curves toward the dorsal anchor point) (vs. superficial root straight, at ∼45° angle to deep root and directed away from the dorsal anchor point), a single, much reduced transverse bar that is narrow for its entire breadth (vs. dorsal and ventral transverse bars robust and broad, having an irregular outline), an accessory piece that lacks digitiform projections (vs. accessory piece with 4 digitiform projections), and an accessory piece that lacks a half cardioid-shaped process (vs. accessory piece having a half cardioid-shaped process). Our 28S sequences (generated from 4 specimens of D. cf. skrjabini: 2 from Tennessee [763 base pairs (bp)] and 2 from Arkansas [776 bp]) were identical to 1 ascribed to D. skrjabini from Japan. The present study is the first verifiable and credible report of a parasite from silver carp in North America and the first nucleotide information for a parasite from silver carp in North America.

ENCYCLOBREPHUS BREVIVITELLUS N. SP. (DIGENEA: PLAGIORCHIOIDEA, INCERTAE SEDIS) INFECTING THE MEKONG SNAIL-EATING TURTLE, MALAYEMYS SUBTRIJUGA, IN THE MEKONG RIVER, VIETNAM, WITH PHYLOGENETIC ANALYSES BASED ON RDNA.

A new digenean species belonging in EncyclobrephusSinha, 1949 is described, and the generic diagnosis is amended to accommodate variation in several features of the new species. Worms were collected from the intestines of 2 specimens of the Mekong snail-eating turtle, Malayemys subtrijuga (Schlegel and Müller, 1845). Permanent whole-mounted worms were studied using light microscopy, and ribosomal DNA (rDNA) sequences were generated from 3 worms. We investigated the phylogenetic relationship of the new species among some digenea using separate Bayesian inference analyses, 1 based on the 28S rDNA gene and rooted using a species from the Monorchioidea Odhner, 1911, and a second based on the internal transcribed spacer 1 region rooted by a species in the Microphalloidea Ward, 1901. Prior to the analyses, Encyclobrephus was classified in the Encyclometridae Mehra, 1931. Previous studies using rDNA from the type species for the family, Encyclometra colubrimurorum (Rudolphi, 1819) Baylis and Cannon, 1924, have demonstrated that En. colubrimurorum is closely related to species of Polylekithum Arnold, 1934 in the Gorgoderoidea Looss, 1901. Nevertheless, phylograms from both analyses indicated that the new species of Encyclobrephus belongs in the Plagiorchioidea Lühe, 1901, related to species in the families Cephalogonimidae Looss, 1899, Plagiorchiidae Lühe, 1901, Reniferidae Pratt, 1902, and Telorchiidae Looss, 1899. The present results suggest that Encyclobrephus is not closely related to En. colubrimurorum. Familial classification of Encyclobrephus is contingent on molecular data availability for the type species but it should be removed from the Encyclometridae and classified as incertae sedis within the Plagiorchioidea. Encyclometridae belongs in the Gorgoderoidea, not the Plagiorchioidea.

RESOLVING THE PARAPHYLETIC TURTLE BLOOD FLUKES: REVISION OF SPIRORCHIIDAE STUNKARD, 1921 AND PROPOSAL OF CARETTACOLIDAE YAMAGUTI, 1958, HAPALOTREMATIDAE (STUNKARD, 1921) POCHE, 1926, BARACKTREMATIDAE N. FAM., PLATTIDAE N. FAM., AND ATAMATAMIDAE N. FAM.

Turtle blood flukes ("spirorchiids") comprise a paraphyletic assemblage including the monophyletic Schistosomatidae Stiles and Hassall, 1898 as a crown group. We herein morphologically diagnose the natural groups of turtle blood flukes and propose family names for them. Spirorchiidae Stunkard, 1921 (Spirorchis MacCallum, 1919 [type]; Spirhapalum Ejsmont, 1927; Plasmiorchis Mehra, 1934; MonticelliusMehra, 1939; VasotremaStunkard, 1928; provisionally UterotremaPlatt and Pichelin, 1994) has a ventral sucker (lost in Spirorchis), an esophageal gland surrounding the entire esophagus, lateral esophageal diverticula (plicate organ and medial esophageal diverticulum present or absent), a glandular mass at the esophagus base, paired ceca surrounded by vitelline follicles for their entire length, a non-filamented, ovoid egg, and typically a Manter's organ. Baracktrematidae n. fam. (Baracktrema Roberts, Platt, and Bullard, 2016 [type]; Unicaecum Stunkard, 1925; NeospirorchisPrice, 1934) has no ventral sucker, a single cecum or cyclocoel, a coiled or looped testis extending into the anterior body half, a convoluted post-cecal and post-gonadal uterus, a uterine pouch, no metraterm, no Manter's organ, and non-filamented eggs. Plattidae n. fam. (Platt Roberts and Bullard, 2018 [type]; Hapalorhynchus Stunkard, 1922; Coeuritrema Mehra, 1933; Enterohaematotrema Mehra, 1940; CardiotremaDwivedi, 1967; Ruavermis Dutton and Bullard, 2020) has the anatomical sequence of a ventral sucker, external seminal vesicle, cirrus sac, anterior testis, ovary, transverse vitelline duct, and posterior testis (except Enterohaematotrema) plus a pars prostatica, dorsal genital pore, globular excretory vesicle, and no Manter's organ. Carettacolidae Yamaguti, 1958 (CarettacolaManter and Larson, 1950 [type]) has a spinose oral sucker, a spinose and contractile ventral sucker (transverse cavity present), numerous testes distributing in a post-ovarian, inter-cecal column, and an ovary and transverse vitelline duct that are pre-testicular. Hapalotrematidae (Stunkard, 1921) Poche, 1926 (Hapalotrema Loos, 1899 [type]; AmphiorchisPrice, 1934; LearediusPrice, 1934; Cheloneotrema Simha and Chattopadhyaya, 1980; NeocaballerotremaSimha, 1977; Satyanarayanotrema Simha and Chattopadhyaya, 1980; Shobanotrema Simha and Chattopadhyaya, 1980) has the anatomical sequence of a ventral sucker, anterior testis(es), external seminal vesicle and cirrus sac, ovary, and posterior testis(es). Atamatamidae n. fam. (Atamatam Bullard and Roberts, 2019 [type]; Paratamatam Bullard and Roberts, 2019; Pitiutrema Dutton and Bullard, 2019) has an oral sucker with anteroventral spines, vasa efferentia connecting to a pre-ovarian external seminal vesicle, a cirrus sac directed postero-laterad and having an internal seminal vesicle and glandular luminal wall, and a dendritic ovary flanked by the external seminal vesicle and oviducal seminal receptacle. A 28S phylogenetic analysis failed to reject the monophyly of these families.

Huffmanela cf. huffmani (Nematoda: Trichosomoididae) infecting swim bladder, peritoneum, and gonad of variable platyfish, Xiphophorus variatus (Cyprinodontiformes: Poeciliidae) and eastern mosquitofish, Gambusia holbrooki (Poeciliidae) in Florida; taxonomy, phylogenetic analysis, and pathological changes.

Variable platyfish, Xiphophorus variatus (Meek, 1904) (Cyprinodontiformes: Poeciliidae) and eastern mosquitofish, Gambusia holbrooki Girard, 1859 (Poeciliidae) from earthen ponds in west central Florida were examined for parasitic infections. At necropsy, we observed myriad nematodes (adults and eggs), which we identified as Huffmanela cf. huffmani, infecting the swim bladder, gonad, and visceral peritoneum. Nucleotide sequences (small subunit ribosomal DNA, 18S) of H. cf. huffmani from variable platyfish and eastern mosquitofish were identical; likewise for newly obtained 18S sequences of Huffmanela huffmani Moravec, 1987 from the swim bladder of red breast sunfish, Lepomis auritus (Linnaeus, 1758) (Centrarchiformes: Centrarchidae) and warmouth, Lepomis gulosus (Cuvier, 1829) from the San Marcos River (type locality for Huffmanela huffmani Moravec, 1987), Texas. The sequences of H. huffmani and H. cf. huffmani differed by 7 (1%) nucleotides. Pathological changes comprised proliferation of the tunica externa of the swim bladder in low-intensity infections in addition to inflammation, proliferation, and tissue necrosis of swim bladder, peritoneum, and gonad in high-intensity infections. The lesion was severe, affecting the cellular constituents of the swim bladder wall and reducing the size of the swim bladder lumen; potentially reducing swim bladder physiological efficiency. The present study is the first record of a freshwater species of Huffmanela Moravec, 1987 from beyond the San Marcos River, first record of a species of Huffmanela from a livebearer, first nucleotide sequences and phylogenetic analysis for Huffmanela, and first evidence that an infection by a species of Huffmanela causes pathological changes that could impact organ function.

Paraharmotrema karinganiense n. gen., n. sp. (Digenea: Liolopidae) infecting the intestine of serrated hinged terrapin (Pelusios sinuatus), east African black mud turtle (Pelusios subniger), and South African helmeted turtle (Pelomedusa galeata) and a phylogenetic hypothesis for liolopid genera.

We herein describe Paraharmotrema karinganiense n. gen., n. sp. Dutton & Bullard (Liolopidae Dollfus, 1934) from specimens infecting the intestine of the serrated hinged terrapin (Pelusios sinuatus), east African black mud turtle (Pelusios subniger) (both Nwanedzi River, Mozambique), and South African helmeted terrapin (Pelomedusa galeata) (North-western Zululand, KwaZulu-Natal Province, South Africa). The new genus can be easily differentiated from the other accepted liolopid genera (Liolope Cohn, 1902; Helicotrema Odhner, 1912; Harmotrema Nicoll, 1914; Dracovermis Brooks & Overstreet, 1978) by the combination of having a linguliform body approximately 6-9 × longer than wide, tegumental spines/scales, a minute ventral sucker located in the anterior 1/7-1/8 of the body, deeply lobed testes that are transverse and abut the caeca (spanning the intercaecal space), a uterus that is lateral to the anterior testis (not ventral to the anterior testis), a lobed ovary that is dextral and nearest the posterior testis, and a vitellarium that does not extend anteriad to the level of the ventral sucker and that does not fill the intercaecal space. Nucleotide sequences of large subunit ribosomal DNA (28S) and internal transcribed space region (ITS2) from all analyzed specimens of the new species were identical, respectively; the 28S sequences differed from that of Liolope copulans Cohn, 1902 and from that of Harmotrema laticaudae Yamaguti, 1933 by 103 (8%) and 105 (8%) nucleotides, respectively. The 28S phylogenetic analysis recovered the new genus sister to a clade comprising L. copulans and H. laticaudae. A key to liolopid genera is provided herein. The present study comprises the first nucleotide-based phylogenetic placement of Harmotrema and first record of a liolopid from South Africa or Mozambique. It is the first proposal of a new liolopid genus in 43 yrs, and it documents a second liolopid genus from P. subniger while tripling the number of liolopid turtle hosts reported from the continent of Africa.

Resurrection and emendation of Pseudoparamacroderoides Gupta & Agrawal, 1968 (Digenea: Macroderoididae), description of a new species from Vietnam, and comments on the systematics of macroderoidid genera.

We herein resurrect and emend Pseudoparamacroderoides Gupta & Agrawal, 1968 (Digenea: Macroderoididae) and describe a new species, Pseudoparamacroderoides dongthapensis n. sp., from adult specimens infecting the intestine of a riverine catfish, Mystus mysticetus Roberts, (Siluriformes: Bagridae) in the Mekong River, Vietnam. Pseudoparamacroderoides (Pseudoparamacroderoides seenghali Gupta & Agrawal, 1968 [type species]; Pseudoparamacroderoides vittati Kakaji, 1969 [= Ps. vittatusi]; Pseudoparamacroderoides raychaudhurii Agarwal & Kumar, 1983; and Pseudoparamacroderoides keni Agarwal & Agarwal, 1984) differs from other macroderoidid genera by having the combination of a subspherical oral sucker that lacks distinctly-enlarged circumoral spines; caeca that extend posteriad beyond the testes without forming a cyclocoel; testes that are approximately ≤1/3 maximum body width in diameter; a cirrus sac that is claviform, slightly dorsal to and predominantly lateral to the ventral sucker (cirrus sac partially dorsolateral to dextral or sinistral margin of ventral sucker); symmetrical vitelline fields that extend posteriad to the middle of the post-testicular space (not restricted to the inter-gonadal space) and that remain separate (not confluent) anteriorly and posteriorly; and an excretory vesicle that is I-shaped (with or without anterior swelling) and wholly post-ovarian, inter-testicular, or median to the posterior testis. Pseudoparamacroderoides dongthapensis n. sp. differs from its congeners by having an elongate hindbody (>2× forebody length) and an excretory vesicle that is approximately half as long as the body and that extends anteriad beyond the anterior testis. This is the first record of a species of Pseudoparamacroderoides from beyond the Indian sub-continent, from M. mysticetus, and from the Mekong River or from Vietnam. A diagnostic key to macroderoidid genera and a key to Pseudoparamacroderoides spp. are provided.

REDESCRIPTION OF CATHARIOTREMA SELACHII (MACCALLUM, 1916) JOHNSTON AND TIEGS, 1922 (MONOGENOIDEA: MONOCOTYLIDAE), EMENDATION OF MONOTYPIC CATHARIOTREMA JOHNSTON AND TIEGS, 1922, AND PROPOSAL OF CATHARIOTREMATINAE N. SUBFAM. BASED ON MORPHOLOGICAL AND NUCLEOTIDE EVIDENCE.

We herein redescribe the enigmatic Cathariotrema selachii (MacCallum, 1916) Johnston and Tiegs, 1922 based on the holotype, paratypes, and newly collected specimens infecting the olfactory organ of 5 shark species from the Gulf of Mexico (all new host records): scalloped hammerhead shark, Sphyrna lewini (Griffith and Smith, 1834) (Carcharhiniformes: Sphyrnidae); great hammerhead shark, Sphyrna mokarran (Rüppell, 1837); blacktip shark, Carcharhinus limbatus (Müller and Henle, 1839) (Carcharhiniformes: Carcharhinidae); spinner shark, Carcharhinus brevipinna (Müller and Henle, 1839); and Atlantic sharpnose shark, Rhizoprionodon terraenovae (Richardson, 1836) (Carcharhinidae). These specimens were morphologically indistinguishable from each other and from MacCallum's holotype and paratypes. Those sequenced had identical first internal transcribed spacer (ITS1) and large subunit ribosomal DNA (28S) nucleotide sequences. As such, C. selachii infects sharks of 2 orders (Carcharhiniformes, Lamniformes) and 3 families (Carcharhinidae, Sphyrnidae, Lamnidae) in the Northwestern Atlantic Ocean (type locality) and Gulf of Mexico (new records herein). This report is the first of new specimens of C. selachii in the Atlantic Ocean Basin in 95 yr and corrects long-standing error cascades and ambiguities concerning the morphology and systematic placement of C. selachii. Considering morphology and nucleotide-based phylogenetic evidence (28S, Bayesian analysis), we herein emend monotypic CathariotremaJohnston and Tiegs, 1922 and propose Cathariotrematinae Bullard n. subfam. for it and 4 other genera (all formerly assigned to Merizocotylinae Johnston and Tiegs, 1922). These genera comprise species infecting only the nose of sharks (monotypic Cathariotrema, SqualotremaKearn and Green, 1983 and SeptitremaKheddam, Chisholm, and Tazerouti, 2020 plus 3 species of TriloculotremaKearn, 1993) and nose of a chimaera (monotypic HolocephalocotyleDerouiche, Neifar, Gey, Justine, and Tazerouti, 2019). Cathariotrematinae differs from Merizocotylinae by having a 3-part attachment organ and by lacking open loculi that symmetrically encircle a cluster of >2 loculi in the center of the haptor. Monophyletic Cathariotrematinae (with sequences representing species of Cathariotrema, Triloculotrema, and Holocephalocotyle only) was sister to monophyletic Merizocotylinae, which together were sister to monophyletic Calicotylinae Monticelli, 1903. These subfamilies comprise a monophyletic group of monocotylids that have a double vagina and infect extrabranchial, enclosed niches (urogenital system, body cavity, olfactory chamber/nose) on their shark, ray, and chimaera hosts (all other monocotylids have a single vagina and infect the gill or body surfaces of rays only). Monocotylinae Taschenberg, 1879 and Decacotylinae Chisholm, Wheeler, and Beverley-Burton, 1995 were recovered as monophyletic. Heterocotylinae Chisholm, Wheeler, and Beverley-Burton, 1995 remained paraphyletic. We accept ParacalicotyleSzidat, 1970.

Two new species of Cephalogonimidae Looss, 1899 (Digenea: Plagiorchioidea) from Africa (Mozambique and Guinea), including a new phylogenetic hypothesis for related plagiorchioids.

Two new species of Cephalogonimidae Looss, 1899 (from Emoleptalea Looss, 1900 and Masenia Chatterji, 1933) are described from African freshwater fishes. Emoleptalea mozambiquensis n. sp. infected the turquoise killifish, Nothobranchius furzeri Jubb, in Mozambique and differs from its nine congeners by the combination of differences in body shape and size, oral sucker shape, sucker width ratio, configuration of the digestive tract and gonads, vitelline follicle shape and vitellarium configuration. Emoleptalea dollfusi Srivastava, 1960 is a synonym of Emoleptalea loossi Srivastava, 1960, thus there are still nine accepted species. Masenia baroensis n. sp. infected the globe fish, Tetraodon lineatus L., in the Republic of Guinea and differs from its five African congeners and 15 Asian congeners by the combination of circumoral spine count, oral sucker shape, caecal extent, ovary shape, genital pore position, and configuration of the vitellarium. Masenia dayali (Gupta & Puri, 1984) Chandra & Saxena, 2016 and Masenia pushpanjalii are nomina dubia. We propose Masenia ritai (Agrawal, 1964) n. comb., with M. ritai Sircar & Sinha, 1970 its junior synonym. Heterorchis cf. crumenifer (identified tentatively due to egg size) is reported from the West African lungfish, Protopterus annectens (Owen), in Mozambique (new geographical record). Heterorchis protopteri Thomas, 1958 and Heterorchis ghanensis Thomas, 1968 are species inquirendae. Sequences (28S rDNA) from these parasites were included in a Bayesian phylogenetic analysis with 37 other ingroup taxa. Both new species formed a clade with Masenia nkomatiensis Dumbo, Dos Santos & Avenant-Oldewage, 2019 from Africa. These three species formed a sister relationship with the other available cephalogonimids: Cephalogonimus americanus Stafford, 1902 and Cephalogonimus retusus (Dujardin, 1845), both frog parasites from North America and Europe, respectively. Heterorchis cf. crumenifer represented a distinct lineage within the Plagiorchioidea but formed a polytomy with species from 10 plagiorchioid families.

Redescription of the Type Species of Cardicola Short, 1953 (Digenea: Aporocotylidae) and Description of a New Congener Infecting Yellowedge Grouper, Hyporthodus flavolimbatus (Perciformes: Serranidae), from the Gulf of Mexico.

Cardicola Short, 1953 is the most speciose aporocotylid genus (35 species) and includes marine and estuarine species of fish blood flukes that infect "higher ray-finned fishes" (Euteleostei). Several clades within Cardicola are recovered in phylogenetic analyses of the large subunit ribosomal DNA (28S), but morphological synapomorphies for those nucleotide-based clades remain elusive. The type species, Cardicola cardiocola (Manter, 1947) Short, 1953, has not been recollected in 73 yr and the original description was incomplete; making a genus revision challenging because of the ambiguous systematic position of its type species. Herein, we redescribe C. cardiocola by using the holotype (USNM 1337732) and new specimens collected from the type host, jolthead porgy, Calamus bajonado (Sparidae), from nearby the type locality. It differs from its congeners by the combination of having a body that is 5 times longer than wide, an anterior sucker with concentric rows of spines, 2-6 tegumental body spines per row, an esophageal gland that is 22-43% of the esophageal length, a testis that is 3-5 times longer than wide and that fills the intercecal space, a vitelline duct connecting to the anterior aspect of the oötype, an ascending uterus that lacks any coil, a descending uterus yielding a single coil, an obvious cirrus sac separated by constriction from the seminal vesicle, a tegumental protrusion surrounding the terminal end of cirrus sac, and a male genital pore that is posterior to the remainder of the genitalia. We also describe a new congener infecting the heart of yellowedge grouper, Hyporthodus flavolimbatus (Serranidae), from the Gulf of Mexico. It differs from its congeners by the combination of having an anterior sucker that does not extend beyond the anterior body margin, 2-5 tegumental body spines per row, posterior ceca that are 9 times length of the anterior ceca and that lack any coil, a testis that is 3 times longer than wide and that does not fill the intercecal space, an ovary that is >60% of the body width, a vitelline duct that connects to the anterior aspect of the oötype, a uterus that is >10% of the body width and that extends posterior to all genitalia, and a rounded posterior body margin. It is the first species of Cardicola to be described from a grouper (Serranidae). The 28S and internal transcribed spacer 2 phylogenetic analyses recovered the new species as a distinct lineage within the clade of Cardicola spp.

First Record of a Polystome from Alligator Snapping Turtle, Macrochelys temminckii (Cryptodira: Chelydridae) or Mississippi; with Comments on "Neopolystoma orbiculare (Stunkard, 1916)" and Its Junior Subjective Synonyms.

Herein, we describe several newly-collected specimens of Neopolystoma cf. orbiculare from the urinary bladder of 2 alligator snapping turtles, Macrochelys temminckii (Troost in Harland, 1835) (Cryptodira: Chelydridae Gray, 1831) from Comet Lake (30°35'46.94″N, 88°36'3.12″W), Pascagoula River, Mississippi. Our specimens differed from all previous descriptions of N. orbiculare and its junior subjective synonyms by the combination of having intestinal ceca adorned with triangular pockets and that terminate dorsal to the haptor, distinctive hooklets each having a handle and guard of approximately equal length and having a much longer and curved blade, 16 genital coronet spines that each possess 1-2 flanges per spine, pre-testicular vaginal pores, and vaginal ducts that are anterior to the junction of the oviduct and genito-intestinal canal. Some of our specimens were enantiomorphic (4 and 3 had a dextral and sinistral ovary, respectively). Nucleotide sequences (large subunit ribosomal DNA [28S], small subunit ribosomal DNA [18S], and cytochrome oxidase subunit 1 mitochondrial gene [COI]) for our specimens were most similar to GenBank sequences ascribed to N. orbiculare. Single-gene and concatenated phylogenetic analyses confirmed that NeopolystomaPrice, 1939 is polyphyletic and that our isolates share a recent common ancestor with those ascribed to N. orbiculare. This is the first record of a polystomatid from Mississippi, from the Pascagoula River, and from the alligator snapping turtle (and only the second species of Neopolystoma reported from any snapping turtle).

A new species and emendation of the seldom reported Enterohaematotrema Mehra, 1940 (Digenea: Schistosomatoidea), including a revised phylogenetic hypothesis for turtle blood flukes.

Enterohaematotrema Mehra, 1940 is emended herein based upon a review of the literature and a description of a new species (Enterohaematotrema triettruongi n. sp.) infecting yellow-headed temple turtles, Heosemys annandalii (Boulenger) (Cryptodira: Geoemydidae), in the Mekong River, Vietnam. The new species differs from the published descriptions of its congeners Enterohaematotrema palaeorticum Mehra, 1940 and Enterohaematotrema hepaticum (Simha, 1958) Simha & Chattopadhyaya, 1980 by having two distinctive oesophageal glands, a short and eversible cirrus (vs protrusive with 3 distinct processes), a dorsal common genital pore that is sinistral (vs ventral and medial), a transverse (vs longitudinal) external seminal vesicle, an oviducal seminal receptacle that is sinistral (vs dextral), and a vitellarium distributing from the caecal bifurcation (anterior to the ventral sucker) to the caecal tips (vs vitellarium not extending anteriad beyond ventral sucker in E. palaeorticum or vitellarium wholly posterior to the terminal genitalia in E. hepaticum). A phylogenetic analysis of the D1-D3 domains of the nuclear large subunit ribosomal DNA (28S) recovered Enterohaematotrema and Platt Roberts & Bullard, 2016 as sister taxa that share a recent common ancestor with the clade comprising Ruavermis Dutton & Bullard, 2020 and Coeuritrema Mehra, 1933. These flukes collectively comprise a monophyletic group of southeast Asian turtle blood flukes. This analysis also indicated that the massive, longitudinal metraterm of species of Enterohaematotrema and Uterotrema Platt & Pichelin, 1994 represents homoplasy (convergent evolution). The present study comprises the first morphological study of original specimens of any species of Enterohematotrema in more than 50 years and is the first molecular phylogenetic placement of the genus among the various turtle blood fluke lineages.

An updated geographic distribution of Myxobolus cerebralis (Hofer, 1903) (Bivalvulida: Myxobolidae) and the first diagnosed case of whirling disease in wild-caught trout in the south-eastern United States.

Myxobolus cerebralis (Bivalvulida: Myxobolidae), the aetiological agent of salmonid whirling disease, was detected in 2 river basins of North Carolina during 2015, which initiated the largest spatial-temporal monitoring project for the disease ever conducted within the south-eastern United States (focused mainly in eastern Tennessee and western North Carolina). A total of 2072 rainbow trout Oncorhynchus mykiss, 1,004 brown trout Salmo trutta and 468 brook trout Salvelinus fontinalis were screened from 113 localities within 7 river basins during June 2017 through October 2019. Infections were detected by pepsin-trypsin digest, microscopy and the species-specific nested polymerase chain reaction (PCR) in 19 localities across 6 river basins. Myxospore morphology was indistinguishable from the published literature. In 2019, five rainbow trout that symptomatic for whirling disease (sloping neurocranium and lordosis) were captured and processed for histopathology. Myxospores were detected in the calvarial cartilage of two deformed trout with associated erosion of the cartilage consistent with reported whirling disease lesions. This is the first report of M. cerebralis in Tennessee and the first histologically confirmed cases of whirling disease in southern Appalachian (south-eastern United States) rivers and streams and expands the distribution of M. cerebralis throughout western North Carolina and eastern Tennessee.

Ruavermis mikebargeri gen. et sp. n. (Digenea: Schistosomatoidea) infecting the yellow-headed temple turtle, Heosemys annandalii (Cryptodira: Geoemydidae), in Vietnam, including an updated phylogeny for the turtle blood flukes.

Ruavermis mikebargeri gen. et sp. n. infects the yellow-headed temple turtles Heosemys annandalii (Boulenger et Robinson) in the Mekong River Basin. It resembles Platt Roberts et Bullard, 2018 and Coeuritrema Mehra, 1933 by having the anterior to posterior anatomical sequence of a ventral sucker, external seminal vesicle, cirrus sac, anterior testis, ovary, transverse vitelline duct, and posterior testis. These genera are further similar by having the combination of an elongate/ovoid aspinous body, a ventral sucker at the level of the body constriction, an oesophagus that terminates in the anterior 1/5 of the body and that is ventral to the anterior nerve commissure, intestinal caeca that bifurcate in the anterior 1/3 of the body (not immediately anterior to the ventral sucker), a sinistral caecum that bends toward the midline at level of the cirrus and common genital pore, an external seminal vesicle that abuts the anterodextral margin of the cirrus sac, an oviduct that emerges from the dextral margin of the ovary, and an oviducal seminal receptacle that comprises the middle portion of the oviduct. These genera lack lateral oesophageal diverticulae and a median oesophageal diverticulum. The new genus is unique by having a papillate ventral body surface, an external seminal vesicle lateral to the cirrus sac, vasa efferentia that are ventral to the gonads, an oviduct that is convoluted, a Laurer's canal pore that is preovarian, a Laurer's canal that extends anterolaterad, and an excretory vesicle that is Y-shaped. The 28S rDNA phylogenetic analysis recovered the new species sister to Coeuritrema platti Roberts et Bullard, 2016, with that clade sister to Hapalorhynchus spp. and Platt spp. The new turtle blood fluke is the fourth from Vietnam, second from a Vietnam geomydid, and first from Heosemys Stejneger as well as the first endohelminth from the yellow-headed temple turtle.

New Genus and Species of Turtle Blood Fluke (Platyhelminthes: Digenea: Schistosomatoidea) Infecting Six-Tubercled Amazon River Turtles, Podocnemis sextuberculata (Pleurodira: Podocnemididae) from the Amazon River Basin (Peru).

Herein we describe a new species of turtle blood fluke (TBF) and propose a new genus to accommodate it, Pitiutrema revelae n. gen., n. sp. This blood fluke infects the heart of six-tubercled Amazon River turtles (Podocnemis sextuberculata [Cornalia, 1849] [Pleurodira: Podocnemididae]) in the headwaters of the Amazon River near Iquitos, Peru. It resembles the other 2 described species of South American freshwater TBFs (Atamatam amazoniensis Bullard and Roberts, 2019, and Paratamatam iquitosiensis Bullard and Roberts, 2019) by having a dorsoventrally flattened and ovoid body, an oral sucker with anteroventral spines, 2 inter-cecal testes arranged in a column, inter-gonadal terminal genitalia, an inter-cecal and post-ovarian Laurer's canal pore, and a Y-shaped excretory bladder. It differs from all other nominal TBFs by having the combination of an aspinose body that lacks mammillae, a tapered (not broadly rounded) posterior body end, a ventral sucker, slightly M-shaped or inverse U-shaped ceca, a deeply-lobed (dendritic) ovary, a transverse uterus, and a dispersed vitellarium. The new genus is further unique among TBF genera by having an anterior to posterior sequence of ventral sucker, anterior testis, ovary, cirrus sac (lateral to posterior half of ovary), and posterior testis. The phylogenetic results and placement of the new taxon (1) were both predicted by our morphological diagnosis and comparisons with related taxa, (2) further indicated monophyly of the nominal South American freshwater TBFs, (3) reaffirmed the marine derived lineage identity of the nominal South American freshwater TBFs, and (4) highlighted that the single cercarial sequence (TBF sp. W-810) from an ampullariid in Brazil does not share a recent common ancestor with any of the nominal South American freshwater TBFs. The new species is the eighth TBF reported from a side-necked turtle (Pleurodira), the first TBF from a member of Podocnemididae, and the third freshwater TBF from South America.

Neotropical Turtle Blood Flukes: Two New Genera and Species from the Amazon River Basin with a Key to Genera and Comments on a Marine-Derived Parasite Lineage in South America.

Two new genera and species of freshwater turtle blood flukes (TBFs) are described herein based on specimens infecting the nephritic and mesenteric blood vessels of "matamatas" (a side-necked turtle, Chelus fimbriata [Schneider, 1783] [Pleurodira: Chelidae]) from the Amazon River Basin, Peru. These taxa comprise the first-named species and the first-proposed genera of freshwater TBFs from the continent of South America. A new comparison of all TBF genera produced 6 morphologically diagnosed groups that are discussed in light of previous TBF classification schemes and a novel phylogenetic hypothesis based on the nuclear large subunit ribosomal DNA (28S). Considering external and internal anatomical features, species of the new genera (Atamatam Bullard and Roberts n. gen., Paratamatam Bullard and Roberts n. gen.) are most similar to each other and are together most similar to those of several marine TBF genera. The 28S phylogenetic analysis supported the monophyly of all 6 morphologically diagnosed groups of genera. Most notably, the freshwater TBFs of South America comprise a derived group nested within the clade that includes the paraphyletic marine TBFs. Not surprisingly in light of morphology, another marine TBF lineage (Neospirorchis Price, 1934) clustered with the freshwater TBFs of Baracktrema Roberts, Platt, and Bullard, 2016 and Unicaecum Stunkard, 1925. Our results, including an ancestral state reconstruction, indicated that (1) freshwater TBFs have colonized marine turtles twice independently and that (2) the South American freshwater TBFs comprise a marine-derived lineage. This is the first evidence that TBFs have twice independently transitioned from a marine to freshwater definitive host. Marine incursion is considered as a possible mechanism affecting the natural history of marine-derived freshwater TBFs in South America. A dichotomous key to accepted TBF genera is provided.

First Record of A Species of Mermithidae Braun, 1883 Infecting a Decapod, Palaemon paludosus (Palaemonidae).

The hemocoel of 26 of 30 (86%) eastern grass shrimps, Palaemon paludosus (Gibbes, 1850) (Decapoda: Palaemonidae), captured during June 2017 from several freshwater lakes near Leesburg and Lake Kissimmee, Florida, was infected by juveniles of a nematode species (Mermithidae sp.). Some infected eastern grass shrimps were preserved whole for histopathology, whereas others were dissected to excise parasitic juvenile nematodes, and still others were kept alive in glass aquaria such that post-parasitic (emerged) juvenile nematodes could be opportunistically observed alive and ultimately preserved. Parasitic and post-parasitic juvenile nematodes had cuticle cross-fibers, cephalic papillae, cup-shaped amphids, a horn-shaped vagina, a trophosome, and a caudal appendage, which collectively diagnosed them as Mermithidae sp. They differed from those of nematomorphs (Nematomorpha) by lacking 4 giant cells anteriorly, tegumental bristles, scale or plate-like areoles, a bifurcate or trifurcate posterior end, and an anus. A phylogenetic analysis of the small subunit rDNA (18S) that included all of the available mermithid sequences corresponding to morphologically diagnosed specimens recovered Mermithidae sp. within the clade of mermithids and sister to Ovomermis sinensis Chen, Jian, and Ren, 1991 . This is the first record of a mermithid infection in a decapod and first report of a mermithid infection in an aquatic crustacean from North America (another mermithid infects a terrestrial isopod there). The high prevalence of infection and the multiple geographic localities harboring infected eastern grass shrimps indicated that these infections were not spurious. Because no other decapod is confirmed as a mermithid host, we suspect that these specimens likely represent a new species with a life cycle worth studying, since none for a mermithid involving a decapod nor a crustacean has been elucidated to date. We also provide a table of all mermithid and nematomorph infections in crustaceans.