Journal of Great Lakes Research 39 (2013) 555–559 Contents lists available at ScienceDirect Journal of Great Lakes Research journal homepage: www.elsevier.com/locate/jglr What's in a name? Taxonomy and nomenclature of invasive gobies in the Great Lakes and beyond Carol A. Stepien ⁎, Matthew E. Neilson 1 The Great Lakes Genetics/Genomics Laboratory, Lake Erie Center and the Department of Environmental Sciences, The University of Toledo, 6200 Bayshore Road, Toledo, OH 43616, USA a r t i c l e i n f o Article history: Received 10 April 2013 Accepted 14 August 2013 Available online 6 October 2013 Communicated by Wendylee Stott Index words: Gobiidae Invasive species Neogobius Proterorhinus Round Goby Tubenose Goby a b s t r a c t The species identities, scientific names, and relationships of Eurasian gobies that invaded the Laurentian Great Lakes – and other species that are predicted to invade in the future – are evaluated here using recently resolved DNA characters. The Round Goby and the Freshwater Tubenose Goby entered the Great Lakes ca. 1990 via ballast water originating from Black Sea ports. The Round Goby spread extensively throughout the Great Lakes and adjacent rivers, whereas the Freshwater Tubenose Goby recently began to expand its range. Both species also are widely invasive in Eurasia, dispersing via canals and shipping. Several of their relatives – the Monkey, Racer, and Bighead gobies – also are invasive in Eurasia, and are predicted to invade the Great Lakes. We discuss results from phylogenetic analyses of DNA sequences from 4 mitochondrial and nuclear gene regions, and provide a revision of their scientific nomenclature. The Freshwater Tubenose Goby was redefined as Proterorhinus semilunaris, which is markedly different and distinctive from the Marine Tubenose Goby Proterorhinus marmoratus. The genus Neogobius, as formerly defined, contained multiple evolutionary lineages and incorrect scientific names. We thus restricted Neogobius to just 4 species—including the Round Goby Neogobius melanostomus and the Black Sea Monkey Goby Neogobius fluviatilis. Several previously recognized subgenera, which were incorrectly grouped in Neogobius, were elevated to the level of genera. Notably, the Racer Goby became Babka gymnocephalus and the Bighead Goby now is Ponticola kessleri. These changes made the names consistent with their true relationships and species characters, which are essential for identifying and characterizing these gobies in invasive and native habitats. © 2013 International Association for Great Lakes Research. Published by Elsevier B.V. All rights reserved. Introduction Identifying and distinguishing among taxa (i.e., species, populations, and their evolutionary lineages) underlie the essence of biology, being essential to understanding patterns of biogeographic variation, history on our planet, ecological adaptations, and physiological responses (Bortolus, 2008; Cracraft, 2002). Taxonomic or nomenclatural errors can have large, cascading effects on ecological studies and their derived applications, such as environmental remediation and management (see examples in Bortolus, 2008). Lack of taxonomic knowledge also can confound effective management of pests (Kirton, 2005), inhibit improvements of crops (Gupta and Baum, 1986), mask the distinction between invasive and native species (Geller, 1999), and/or preclude ability to make correct ecological comparisons across a range of habitats (Stepien et al., 2005). Understanding the phylogenetic placement of a taxon also provides a framework for its organization and interpreting its biogeographic patterns. In particular, the contributions of taxonomy, systematics, and ⁎ Corresponding author. Tel.: +1 419 530 8362 (Office). E-mail address: carol.stepien@utoledo.edu (C.A. Stepien). 1 Present address: Cherokee Nation Technology Solutions, contracted to USGS Nonindigenous Aquatic Species Program, Southeast Ecological Science Center, 7920 NW 71st Street, Gainesville, FL 32653, USA. biogeography to the conservation of native species and the growing impacts of invasive species have increased over time, especially in the advent of understanding genetic and genomic adaptations. For example, morphological and genetic identification was essential to the discovery that 2 Eurasian species were involved in the mid-1980s dreissenid mussel invasion of the Laurentian Great Lakes of North America — the Zebra Mussel Dreissena polymorpha and the Quagga Mussel Dreissena rostriformis (Jones and Ricciardi, 2005; Spidle et al., 1995; Stepien et al., 2013; Stoeckmann, 2003). These 2 species, although morphologically similar and belonging the same genus, are very phylogenetically distinct, separated by ca. 11–13 million years of evolutionary diversification (Stepien et al., 1999, 2001, 2013). The Quagga Mussel's classification has been controversial: it was previously defined as “D. bugensis”2 (Spidle et al., 1995), was relegated to the subspecies “D. rostriformis bugensis” (Therriault et al., 2004), and recent data have shown that it lacks genetic and morphological distinctiveness from other D. rostriformis, refuting its designation as a separate subspecies (Stepien et al., 2013). Over the past decade the Quagga Mussel has greatly outcompeted the Zebra Mussel in nonindigenous habitats where the 2 species are sympatric, with the latter declining in numbers 2 Note: Quotation marks denote formerly used scientific names that have been changed and corrected. 0380-1330/$ – see front matter © 2013 International Association for Great Lakes Research. Published by Elsevier B.V. All rights reserved. http://dx.doi.org/10.1016/j.jglr.2013.09.020 556 C.A. Stepien, M.E. Neilson / Journal of Great Lakes Research 39 (2013) 555–559 (Nalepa et al., 2009; Ricciardi and Whoriskey, 2004; Wilson et al., 2006), rendering this identification critically important. Two Ponto-Caspian Goby species, the Round and the Tubenose Gobies (identified as Neogobius melanostomus and Proterorhinus “marmoratus”, respectively), first were found in the St. Clair River ca. 1990, where they had been accidentally introduced into the Laurentian Great Lakes basin via ballast water discharge from ocean-going vessels (Jude et al., 1992). Since these introductions, the Round Goby rapidly spread throughout the entire Great Lakes basin and beyond, following the spread of one of its preferred native food sources, Dreissena mussels. The Tubenose Goby has been slower to spread from its original location in the Lake St. Clair region south into Lake Erie (Grant et al., 2012; Kocovsky et al., 2011), although it was found in Duluth Harbor in western Lake Superior in 2001, likely transported via ballast water discharge from vessels traveling this highly-trafficked shipping route (http://www.seagrant.umn.edu/ newsletter/2002/03/tubenose_goby_leaps_to_duluthsuperior.html). The goby and dreissenid mussel invasions, along with introductions of many other Ponto-Caspian species, have stimulated much research on the ecological role and impact of introduced species in the Great Lakes. Correct taxonomic identifications are essential to make accurate ecological comparisons and understand their adaptations. Stepien and Tumeo (2006) and Neilson and Stepien (2009a, 2009b, 2011) analyzed the molecular systematic relationships of PontoCaspian Gobiidae, including the Round and Tubenose Gobies, and their relatives (Table 1). They addressed classification issues and evolutionary relationships within the group, and a revised taxonomy and nomenclature was presented by Neilson and Stepien (2009a) that reflects current understanding. However, recent publications from a variety of authors, including several from this journal (Journal of Great Lakes Research), have continued to use outdated or incorrect names for these gobies in the Great Lakes and Europe. The goal of the present paper thus is to provide clarification of the current nomenclature for these species and their close relatives, from results of recent genetic analyses in relation to morphological characters. We base our analyses on DNA sequence data from the mitochondrial (mt) DNA gene regions cytochrome oxidase I (COI) and cytochrome b (cyt b), and the nuclear DNA S7 ribosomal protein intron 1 (S7) and the recombinationactivating gene 1 (RAG1) previously presented in Neilson and Stepien (2009a, 2009b, 2011), and whose results are summarized in Fig. 1 and Table 1. brackish waters at Sevastopol along the Crimean Peninsula of Ukraine. Thus, these are the designated “type” specimens that form the morphological and biogeographic reference for the genus Proterorhinus and the species P. marmoratus. Additionally, 3 freshwater species of tubenose gobies were described by early ichthyologists (Heckel, 1837; Kessler, 1877; Kriesch, 1873), including “Gobius” semilunaris from specimens collected in the Maritza River from the Balkan region of Bulgaria (Heckel, 1837). All freshwater forms of tubenose gobies later were synonymized with the marine/brackish P. marmoratus by Berg (1949). A full synonymy and taxonomic history of Proterorhinus was presented in Neilson and Stepien (2009a, 2009b). Stepien et al. (2005) and Stepien and Tumeo (2006) discerned that freshwater specimens of the invasive Tubenose Goby from the Great Lakes and freshwater areas of Europe significantly diverged from the Marine Tubenose Goby P. marmoratus. They analyzed mtDNA cyt b gene sequence data variation of native marine/brackish water and freshwater tubenose gobies from the Black Sea basin (including marine/brackish water specimens collected by Stepien at the type location in Sevastopol, Ukraine) versus introduced locations within the North American Great Lakes and other areas in Europe. The conclusion was that the freshwater versus marine/brackish water tubenose gobies comprised 2 separate species due to pronounced genetic differences. Stepien and Tumeo (2006) recommended that the name P. semilunaris be resurrected for the species of Freshwater Tubenose Goby native to the Black Sea rivers and invasive in the North American Great Lakes, the Baltic Sea, and the Danube River, as that goby markedly differed from the marine/brackish water P. marmoratus. In subsequent work on these and an expanded series of specimens, Neilson and Stepien (2009a, 2009b) found that other gene regions, including the mtDNA COI gene, and the nuclear DNA RAG1 gene and S7 intron also revealed pronounced divergences, and combined with morphological variation demonstrated that the freshwater P. semilunaris and the marine P. marmoratus are indeed separate species. Thus, all tubenose gobies we have analyzed throughout the Great Lakes are P. semilunaris. No P. marmoratus have been identified in North America, and that species appears to be exclusively found in marine and brackish waters. In addition, other species of tubenose gobies from other areas of their ranges have been identified by us (Neilson and Stepien, 2009a, 2009b) and other researchers (Freyhof and Naseka, 2007; Naseka et al., 2005). Taxonomic history of Neogobius Taxonomic history of the tubenose gobies Proterorhinus Tubenose gobies were first described in the early 1800s by Pallas (1814); (as “Gobius marmoratus”) from specimens collected in marine/ Similar to P. marmoratus, the Round Goby originally was described by Pallas (1814) from specimens collected in brackish waters at Sevastopol in the Crimean Peninsula of Ukraine. Berg (1949) provided the Table 1 Taxonomy of select Ponto-Caspian Gobies, including taxa invasive in North America and/or Europe. Former genus namea Correct genus Species Common name Invasive? Freshwater (F), marine (M), and/or brackish (B) waters COI barcode GenBank accession number Neogobius Neogobius Neogobius Proterorhinus Proterorhinus (Neogobius) (Neogobius) (Neogobius) (Neogobius) (Neogobius) (Neogobius) (Neogobius) (Neogobius) (Neogobius) (Neogobius) (Neogobius) Neogobius Neogobius Neogobius Proterorhinus Proterorhinus Ponticola Ponticola Ponticola Ponticola Ponticola Ponticola Ponticola Ponticola Ponticola Ponticola Babka melanostomus fluviatilis caspius marmoratus semilunaris cephalargoides constructor cyrius eurycephalus gorlap kessleri platyrostis ratan rhodioni syrman gymnotrachelus Round Gobyb Monkey Goby Caspian Goby Black Sea Marine Tubenose Goby Freshwater Tubenose Gobyb Pinchuk's Goby Caucasian Goby Kura River Goby Mushroom Goby Caspian Bighead Goby Bighead Goby Flatsnout Goby Ratan Goby Riverine Goby Syrman Goby Racer Goby Yes, North America & Eurasia Yes, Eurasia No No Yes, North America and Eurasia No No No No Yes, Europe Yes, Europe No No No No Eurasia F, B, M F, B, M. F, M, B M, B F F, B F F M, B, F M, B, F F, B F, B F, B F F, B F, B FJ526800 FJ526804 FJ526811 EU444689 EU444674 FJ526828 FJ526846 FJ526844 FJ526815 FJ526817 FJ526823 FJ526826 FJ526848 FJ526842 FJ526829 FJ526820 a b In parentheses if recently changed by Neilson and Stepien (2009a). Official common name per Page et al. (2013). C.A. Stepien, M.E. Neilson / Journal of Great Lakes Research 39 (2013) 555–559 557 Fig. 1. Phylogeny of Ponto-Caspian gobies based on combined analysis of 4 gene regions using maximum likelihood, summarized and adapted from Neilson and Stepien (2009a, 2009b, 2011). Numbers at nodes indicate bootstrap support (2000 pseudoreplications), with * = 100%; ◊ indicates species introduced into the Laurentian Great Lakes, ♦ indicates species introduced or expanding in Europe, ‡ indicates species on the GLANSIS (2012) watchlist for potential Great Lakes' introductions. The genera Ponticola (Po.) and Babka formerly were included in the genus Neogobius (yellow shading). The genus Neogobius (blue shading) now is restricted to N. caspius, N. fluviatilis, N. melanostomus, and N. pallasi. Several tubenose gobies in the genus Proterorhinus were removed from synonymy with P. marmoratus; only one of these – P. semilunaris – has been introduced to the Laurentian Great Lakes (purple shading). first modern usage of the genus Neogobius, including the Round Goby's name as N. melanostomus and designating the Monkey Goby N. fluviatilis as the type species for the genus. Berg (1949) also grouped many of the Ponto-Caspian Gobiidae into the genus Neogobius (see Table 1). In any future revisions, the rules of the International Code for Zoological Nomenclature (henceforth termed “The Code”) mandates that the name Neogobius must remain with the taxon containing N. fluviatilis. Stepien et al. (2005) and Stepien and Tumeo (2006) analyzed mtDNA sequence data for several species of Neogobius and other Ponto-Caspian gobies, including invasive and native taxa. They discerned that the Round Goby (N. melanostomus) and the Monkey Goby (N. fluviatilis) are separate related species, but found that the genus Neogobius was not monophyletic, but instead comprised multiple lineages. Thus the Round and Monkey Gobies did not rightly belong to the same lineage as either the Racer or the Bighead Gobies (formerly classified as “N. gymnotrachelus” and “N. kessleri”, respectively; Table 1. The species of “Neogobius” analyzed by Stepien et al. (2005) and Stepien and Tumeo (2006) comprised several lineages. This further was confirmed with phylogenetic analysis of the other mtDNA gene and 2 nuclear DNA regions by Neilson and Stepien (2009a). DNA evidence (and morphological characters) thus showed that the Monkey and Round Gobies were closely related, and belonged to the same genus, but the other gobies were not their nearest relatives. In evolutionary analysis, each lineage (and name) must include all members of a single lineage descended from a common ancestor. Grouping together of multiple lineages or excluding lineages is discouraged, as it leads to incorrect classification through paraphyly (exclusion of a subset of a monophyletic group) or polyphyly (inclusion of multiple unrelated lineages; Funk and Omland, 2003). Berg's (1949) classification of the PontoCaspian gobies thus was incorrect and paraphyletic. Moreover, Iljin (1927) classified the Round Goby and the Monkey Goby in 2 separate subgenera, putting the Round Goby into the monotypic subgenus “Apollonia” and the Monkey Goby into the subgenus Neogobius. That incorrect classification also was followed more recently by Miller and Vasil'eva (2003). Morphological work by Dobrovolov et al. (1995) had recommended that the Round and Monkey Gobies both be reclassified as “Apollonia”, and should not be included in the same genus as the Racer and Bighead Gobies. That change would have been consistent with the DNA sequence data findings by Stepien and Tumeo (2006). However, that change did not follow “The Code” since N. fluviatilis is the type species (i.e., the name bearing taxon) for both the genus and subgenus Neogobius. We thus conducted extensive background research in consultation with the American Fisheries Society Fish Names Committee (Dr. William 558 C.A. Stepien, M.E. Neilson / Journal of Great Lakes Research 39 (2013) 555–559 Eschmeyer of the California Academy of Sciences and the late Dr. Joseph Nelson of the University of Alberta) and colleague Dr. Nina Bogutskaya from the Russian Academy of Sciences in St. Petersburg, Russia, in order to clarify the scientific names. This nomenclature research revealed that the genus name for the Monkey Goby and the Round Goby needed to remain as Neogobius, and the other gobies needed to be excluded from Neogobius due to paraphyly revealed in our DNA work (Neilson and Stepien, 2009a; Stepien and Tumeo, 2006). This can be seen on our phylogenetic tree in Fig. 1. These nomenclature clarifications and changes are detailed in Neilson and Stepien (2009a, 2009b, 2011), who analyzed the relationships of many other gobiid taxa and their relatives. Our nomenclatural changes for the Round Goby and the Tubenose Goby were adopted in the recently published ‘Common and Scientific Names of Fishes from the United States, Canada, and Mexico’ (Page et al., 2013). Other former inconsistencies were that Miller and Vasil'eva (2003) recommended classifying both the Monkey and Round Gobies as belonging to the subgenus “Apollonia”, and the Caspian Goby (Neogobius caspius) in a monotypic subgenus Neogobius. That also would not be allowed by “The Code”, as the type species for the genus and subgenus name Neogobius is the Monkey Goby N. fluviatilis. DNA sequence analyses of the Caspian Goby by us revealed that it belongs with the Round and Monkey Gobies in a restricted genus Neogobius (Neilson and Stepien, 2009a). The Caspian Goby thus now is N. caspius (Table 1). We moreover found that the Caspian Sea Monkey Goby and the Black Sea Monkey Goby are different species, thus we resurrected an old name to describe the Caspian Monkey Goby as a separate species — N. pallasi (Neilson and Stepien, 2011). The Black Sea Monkey Goby remains as N. fluviatilis, as that is the type locality for that species. There thus now are 4 species in the genus Neogobius (see Fig. 1 and Table 1). Changes to names of other Ponto-Caspian gobies The error of paraphyly in the Ponto-Caspian Goby traditional nomenclature was corrected in Neilson and Stepien's (2009a) revised classification, by elevating 2 of Iljin's (1927) subgenera – Babka and Ponticola – for the remainder of the 'neogobiin' species. The new classification thus recognizes 5 genera: Proterorhinus, Mesogobius, Neogobius, Babka, and Ponticola. The scientific names of these gobies are important, as many are invasive in Eurasia, and several species have been cited as likely to invade the Great Lakes in the future, given the vectors, transport and ecological opportunity. Potential invaders to the Great Lakes on the GLANSIS (2012) High-Risk Invasive Fish Species list include the Black Sea Monkey Goby N. fluviatilis and the Racer Goby Babka gymnotrachelus. The tadpole gobies (or benthophilines; Anatirostrum + Benthophilus + Benthophiloides + Caspiosoma) now are known to be closely related to “negobiins” (the term traditionally used for Mesogobius + Proterorhinus + historic Neogobius; Neilson and Stepien, 2009a). The tadpole goby lineage can be discerned on the Fig. 1 tree. One of these, the Starry Tadpole Goby Benthophilus stellatus from the Caspian Sea, also has been identified as a possible future invader of the Great Lakes (GLANSIS, 2012). Conclusions The present work offers an easy-to-use guide to the Ponto-Caspian gobies for ecologists, to facilitate correct comparisons across habitats and among populations. Table 1 summarizes revised taxonomic names for the common Ponto-Caspian gobies, emphasizing those that have invaded the Great Lakes and those that have been predicted to invade in the future. Fig. 1 presents the phylogeny of these common Ponto-Caspian gobies, and highlights taxa that have undergone recent taxonomic revision. Systematics, taxonomy and nomenclature are inherently broad-based and foundational disciplines that, paradoxically, have the potential for rapid change and upheaval: additional data allow for the refinement of phylogenetic and taxonomic hypotheses for species and higher taxa, and the adjustment of the associated nomenclature for these groups. This fluctuation presents difficulties for those not familiar with the taxonomic literature. Yet, many other disciplines – such as community ecology and environmental biology – rely on accurate taxonomy (Godfray et al., 2007; Gotelli, 2004). Although newer citation databases are growing more effective at keeping track of nomenclatural and taxonomic changes (e.g., Web of Science (http:// thomsonreuters.com/products_services/science/science_products/az/web_of_science/) includes both Neogobius melanostomus and “Apollonia melanostoma” as part of their ‘KeyWords Plus’ for articles about Round Goby), it is important for ecologists and others working in the field to understand and know current correct taxonomy for their focal organisms, through working with taxonomists and museum personnel (Gotelli, 2004) or through the use of emerging online taxonomic resources such as the Integrated Taxonomic Information System (http:// www.itis.gov), the Global Biodiversity Information Facility (http:// www.gbif.org), and FishBase (http://www.fishbase.org). Acknowledgments DNA sequence data collection and phylogenetic analyses were funded by NSF Grant #DEB-0456972 to CAS. We are grateful to M. Evans, former Editor-in-Chief of the Journal of Great Lakes Research for encouraging us to write and submit this paper. We thank N. Bogutskaya, W. Eschmeyer, and the late J. Nelson for numerous conversations and valuable insights for resolving the nomenclature of this enigmatic group. We also thank M. Gray for scanning the pictures for the tree, and P. Maitland and K. Linsell for use of the drawings to illustrate the tree. Specimen collections for our original DNA sequence analyses were aided by N. Bogutskaya, V. Boldyrev, L. Corkum, I. Grigorovich, J. Herler, S. Ibrahimov, H. Jenner, J. Kornichuk, V. Kovac, Y. Kvach, A. Naseka, J. Ram, S. Rudnicka, M. Sapoto, P. Simonovic, Y. Slynko, A. Smirnov, and C. Wiesner. Some specimen identifications were resolved by V. Boldyrev and E. Vasil'eva. This is publication #2013-20 from the University of Toledo's Lake Erie Research Center. References Berg, L.S., 1949. Freshwater Fishes of the U.S.S.R. and Adjacent Countries. 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