Proteocephalus perplexus La Rue, 1911

Toma, Scholz, s, Choudhury, Anindo & McAllister, Chris T., 2022, A young parasite in an old fish host: A new genus for proteocephalid tapeworms (Cestoda) of bowfin (Amia calva) (Holostei: Amiiformes), and a revised list of its cestodes, International Journal for Parasitology: Parasites and Wildlife 18, pp. 101-111 : 102-103

3.2.1. Synonym: Proteocephalus perplexus La Rue, 1911

Material studied: holotype ( USNM 1347286) and paratypes ( USNM 1348673–1348680) from Amia calva , Illinois River, Illinois, USA, collected by H.B. Ward in June and July 1910; vouchers: one specimen from A. calva (host code US 029/701), Pascagoula River, Mississippi, USA, collected by V.V. Tkach in March 2009 ( MHNG-PLAT- 0063348); numerous specimens, including two scoleces for SEM, from A. calva (host codes US 610a–i, 611b, 612a–g, 620a, 621c, 622a,b), Green Bay, Lake Michigan, Wisconsin, USA, T. Scholz and A. Choudhury, 15 and 16 May 2017; numerous specimens from A. calva (629d–g, 635a, 636a), Lake Poygan, Wisconsin, T. Scholz and A. Choudhury, 17 May 2017; 14 specimens, their cross and longitudinal (frontal) sections and one scolex for SEM from A. calva ( US 841a, g, 854a), Pascagoula River, Mississippi, USA, T. Scholz, R. Kuchta and M. Oros, 20 and 22 June 2019 ( IPCAS C-774/1); two mounted specimens and one scolex for SEM from A. calva ( US 925a), Turner Pond, McCurtain County, Oklahoma, USA, C.T. McAllister, 1 June 2019 ( IPCAS C-774/1); 1 specimen from A. calva

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(FR19_173.2), Lake Erie , Presque Isle, Pennsylvania, USA, F. Reyda, 15 May 2019 .

Type host: Bowfin, Amia calva Linnaeus ( Amiiformes : Amiidae ).

Additional reported hosts (unconfirmed – see Remarks): Ambloplites rupestris (Rafinesque) (Centrarchiformes: Centrarchidae ); Ameiurus melas (Rafinesque) , A. nebulosus (Lasueur) , Ictalurus punctatus (Rafinesque) ( Siluriformes : Ictaluridae ); Esox sp. ( Esociformes : Esocidae ); Lepisosteus osseus (Linnaeus) , L. platostomus Günther ( Lepisosteiformes : Lepisosteidae ).

Type locality: Illinois River in Havana (40.297067, – 90.060004), Mason County, Illinois, USA (Mississippi River basin) .

Type specimens: holotype – a contracted, overstained immature specimen ( USNM 1347286 ) , paratypes – three slides with fragments of the strobila and ten slides with cross, longitudinal and sagittal sections of scoleces and gravid proglottids ( USNM 1348673–1348680 ) .

Distribution (new geographical records marked with asterisk): Canada (Ontario – Erie, Huron and Ontario Lakes), USA (Illinois, Kentucky, Michigan, Minnesota, Mississippi *, New York, Oklahoma * (southwesternmost distribution in North America), Pennsylvania *, South Carolina, Tennessee, Wisconsin) – La Rue (1911, 1914), Pearse (1924), Hunter (1929), Van Cleave and Mueller (1934), Bangham and Venard (1942), Bangham (1944, 1955), Fischthal (1950), Dechtiar (1972), Bauer and Harley (1973), Anthony (1984), Dechtiar and Christie (1988), Dechtiar et al. (1988), Amin (1990), Aho et al. (1991), Olson and Caira (1999), de Chambrier et al. (2009).

McDonald and Margolis (1995) cited Jarecka et al. (1990) for the record of this tapeworm from New Brunswick, Canada, but that study does not provide any information about source of the bowfin hosts. Amia calva does not occur in New Brunswick (Scott and Crossman, 1973), so the fish hosts must have been collected elsewhere. Contemporaneous studies on another bowfin tapeworm, Haplobothrium globuliforme Cooper, 1914 by two of Jarecka’ s coauthors (see MacKinnon and Burt, 1985 a –c, MacKinnon et al., 1985) state Lake Ontario as the source of bowfin in those studies.

Life cycle: Not elucidated completely. In an abstract from a conference, Jarecka et al. (1990) reported calanoid copepods as intermediate hosts of L. perplexa , but did not provide any details. Plerocercoids presumably conspecific with L. perplexa were found in Ictalurus sp. ( Siluriformes : Ictaluridae ), Morone chrysops (Rafinesque) ( Perciformes : Moronidae ), Pimephales notatus (Rafinesque) ( Cypriniformes : Leuciscidae ) ( Hoffman, 1999); these fish may serve as second intermediate or paratenic hosts.

Representative DNA sequences: (1) 28S rRNA gene (D1–D3): adult from A. calva, Hay Bay, Lake Ontario, Ontario, Canada (1,314 bp; AF286940 – Olson and Caira, 1999); adult from A. calva, Hay Bay, Lake Ontario, Ontario, Canada, 17 July 1995, MHNG-INVE-0025658 (1,009 bp; AJ275228 – de Chambrier et al., 2004); two adults from A. calva, Reelfoot Lake , Tennessee, USA, 30 June 2002, MHNG-INVE-0035366, 0035321 (1,010 bp; FM956089 – de Chambrier et al., 2009); adult from A. calva, Reelfoot Lake , Tennessee, USA, 30 June 2002, MHNG-INVE-36139 (1,005 bp; FM956090 – de Chambrier et al., 2009); adult from A. calva, Hay Bay, Lake Ontario, Ontario, Canada, LRP 8299 (1,396 bp; KF685873 – Caira et al., 2004); adult from Ictalurus punctatus, Reelfoot Lake , Tennessee, USA, 20 June 2002, MHNG-INVE-0036277, sequence identical to MHNG-INVE-0035366 (FM956089). (2) 18S rRNA gene: adult from A. calva, Hay Bay, Lake Ontario, Ontario, Canada (2,037 bp; AF124472 – Olson and Caira, 1999); adult from A. calva, Hay Bay, Lake Ontario, Ontario, Canada, LRP 8299 (1,978 bp; KF685833 – Caira et al., 2004). (3) Mitochondrial 16S rRNA gene: adult from A. calva, Lake Ontario, Ontario, Canada (427 bp; AJ275216 – M. Zehnder, unpublished). (4) Elongation factor-1α: adult from A. calva, Hay Bay, Lake Ontario, Ontario, Canada (987 bp; AF124805 – Olson and Caira, 1999).

Phylogenetic relationships: Laruella perplexa is a member of the “Neotropical fish” superclade of de Chambrier et al. (2015), but its relationships to members of this species-rich clade are not resolved ( de

Chambrier et al., 2004, 2015).