Clarifying the cryptogenic species Polydora neocaeca Williams & Radashevsky, 1999 (Annelida: Spionidae): a shell-bor
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ORIGINAL PAPER
Clarifying the cryptogenic species Polydora neocaeca Williams & Radashevsky, 1999 (Annelida: Spionidae): a shell-boring invasive pest of molluscs from locations worldwide Amore Malan 1 & Jason D. Williams 2 & Hirokazu Abe 3 & Waka Sato-Okoshi 4 & Conrad A. Matthee 1 & Carol A. Simon 1 Received: 26 January 2020 / Revised: 16 March 2020 / Accepted: 27 March 2020 # Senckenberg Gesellschaft für Naturforschung 2020
Abstract Several studies noted similarities between Polydora neocaeca Williams & Radashevsky, 1999, described from the northeast coast of the USA, and shell-boring Polydora haswelli Blake & Kudenov, 1978, described from southeast Australia, and emphasised the need for molecular data to resolve the taxonomy of these species. This study investigates the potential conspecificity of these two species. We provide morphological evidence based on the shape of the modified spines of chaetiger 5 and methyl green staining patterns to support the hypothesis that sand tube-dwelling (habitat of the holotype, i.e. P. haswelli sensu stricto) and shell-boring species identified as P. haswelli are different species. We also provide molecular (nuclear 18S rRNA and mitochondrial COI) and morphological evidence that shows that a previously unidentified shell-boring Polydora species from South Africa and shell-boring species previously recorded as P. haswelli from Japan and China are conspecific with P. neocaeca from its type locality in Rhode Island. We therefore recommend that shell-boring species previously identified as P. haswelli be referred to P. neocaeca. Furthermore, population structure analyses based on COI show distinct geographical groupings, with one shared haplotype among samples for Japan and South Africa. Although limited sampling makes it difficult to draw conclusions regarding the native range for P. neocaeca, we argue that this species represents a cryptic invasion and may have moved with infested molluscs for aquaculture or by shipping. Although described from the east coast of the USA, P. neocaeca may be indigenous to Australia, Asia or an unknown region. Keywords COI . 18S rRNA . Mollusc aquaculture . Cryptic invasion . Tube dwelling
Introduction An important component of non-indigenous species (NIS) associated with molluscs translocated for aquaculture is shell-boring or shell-fouling polydorid polychaetes (Simon
and Sato-Okoshi 2015). Such non-indigenous polydorids pose a considerable biosecurity risk in recipient regions, since they can spread among culture facilities (Simon et al. 2009; Haupt et al. 2012; Williams et al. 2016; Martinelli et al. 2020) and can become pests of native wild molluscs (see Sato-Okoshi
Communicated by D. Fiege * Carol A. Simon [email protected] Amore Malan [email protected]
Conrad A. Matthee [email protected] 1
Department of Botany and Zoology, Stellenbosch University, Matieland, Private Bag X1, Stellenbosch, South Africa
2
Department of Biology, Hofstra University, Hempstead, NY 11549-1140, USA
Hirokazu Abe [email protected]
3
Department of Biology,