BioInvasions Records (2019) Volume 8, Issue 2: 400–409

CORRECTED PROOF

Rapid Communication First record of introduction of magister Dana, 1852 (Crustacea: : ) and range extension of sinensis Milne-Edwards, 1853 (Crustacea: Decapoda: ) in the Long Island Sound

David M. Hudson1,2,3,*, Sandi Schaefer-Padgett1, Barrett L. Christie1 and Richard Harris4 1The Maritime Aquarium at Norwalk, 10 N. Water St., Norwalk, CT 06854 USA 2Center for Environmental Sciences and Engineering, University of Connecticut, Storrs, CT 06269 USA 3Southern Connecticut State University, New Haven, CT 06515 USA 4Copps Island Oysters, 7 Edgewater Place, Norwalk CT 06855 USA Author e-mails: [email protected] (DMH), [email protected] (SS-P), [email protected] (BLC), [email protected] (RH) *Corresponding author

Citation: Hudson DM, Schaefer-Padgett S, Christie BL, Harris R (2019) First record Abstract of introduction of Metacarcinus magister Dana, 1852 (Crustacea: Decapoda: Invasive species have been present in the Long Island Sound, Cancridae) and range extension of northwestern Atlantic Ocean, for over two centuries. Three new records of Eriocheir sinensis Milne-Edwards, 1853 introduction are recorded here from collections by local fishermen. Two records are (Crustacea: Decapoda: Varunidae) in the for male Dungeness , Metacarcinus magister (Dana, 1852), collected in the Long Island Sound. BioInvasions Records 8(2): 400–409, https://doi.org/10.3391/bir. Western Long Island Sound (2017) and Cape Cod Bay (2018). The other record is 2019.8.2.21 that of a range extension documented by a single male Chinese mitten , Eriocheir sinensis (Milne-Edwards, 1853), found in New Haven Harbor, Connecticut. Received: 4 September 2018 Both species could potentially harbor nonnative epibionts and endoparasites. Accepted: 13 February 2019 Additionally, E. sinensis may be more likely to establish, as it has in numerous Published: 24 April 2019 locations in the region and worldwide. Handling editor: Darren Yeo Thematic editor: April Blakeslee Key words: introduced species, , Brachyura, , Copyright: © Hudson et al. , Connecticut, Massachusetts This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International - CC BY 4.0). Introduction OPEN ACCESS. Invasive are common in the Long Island Sound, northwestern Atlantic Ocean, with both the European green crab, Carcinus maenas (Linnaeus, 1758) and the Asian shore crab, sanguineus (De Haan, 1835), already established in the region and interacting with one another for decades (Lohrer and Whitlatch 2002). Invasive crabs have historically caused problems for ecological communities in intertidal and subtidal systems worldwide. Though not an exhaustive list, some of the most damaging crustacean invaders are the European green crab, Carcinus maenas, Chinese mitten crab, Eriocheir sinensis (Milne-Edwards, 1853), two members of the Hemigrapsus, H. sanguieneus, and H. takanoi (Asakura and Watanabe, 2005), the Harris mud crab, Rhithropanopeus harrisii (Gould, 1841), the blue crab, (Rathbun, 1896), and the Asian paddle crab, japonica (Milne-Edwards, 1861)

Hudson et al. (2019), BioInvasions Records 8(2): 400–409, https://doi.org/10.3391/bir.2019.8.2.21 400 Metacarcinus magister and Eriocheir sinensis in the Long Island Sound

(reviewed in Brockerhoff and McLay 2011; McLay 2015). This list of first records for invasions is growing ever larger with the addition of a first record of an eastern Pacific Ocean spider crab, Amphithrax armatus (de Saussure, 1853) to southern Taiwan (Ng et al. 2018). Two species that are regularly consumed by humans have recently appeared in the northwestern Atlantic region: Dungeness crab and Chinese Mitten crab. Dungeness crabs Metacarcinus magister (Dana, 1852) are native to the northeastern Pacific Ocean, and are a major fishery in that region, with the United States fishery landing over 29,000 metric tons worth over $222 million US dollars in the last year on record (National Marine Fisheries Service 2017). They are part of a male-only fishery for which some individuals are shipped live throughout the United States. While there are no reported established populations in the northwestern Atlantic Ocean, an individual was caught off Thatcher Island, Massachusetts in 2006 (Cohen 2006), and another in Cape Ann, Massachusetts in 2009 (Judith Pederson, Ph.D., MIT Sea Grant College Program, pers. comm.). Here, we report the first record of M. magister in the Long Island Sound and an additional record in Massachusetts. Chinese mitten crabs, E. sinensis, are already documented in the general vicinity of the Long Island Sound (Schmidt et al. 2009; reviewed in Dittel and Epifanio 2009). Here, we report an individual record of this species in the Long Island Sound.

Materials and methods The Long Island Sound is an estuary in the northwestern Atlantic Ocean located between the glacial moraine of Long Island to the south and the state of Connecticut to the north. Rhode Island Sound and Cape Cod are to the east, and the City metropolitan area is on the Long Island Sound’s western end. It is a highly seasonal estuary, with southern migratory species moving in during summer and northern species in winter. The Long Island Sound and surrounding estuaries are historically commercial fishing areas and have seen an impact from European development since the 1600s. On 26 July 2017, a local fisherman captured a Dungeness crab, M. magister, in a trap due south of Norwalk Harbor, Connecticut, on Copp’s Island Oysters’ Lot 006 in over 60 meters of water (41.014833°N; 73.394250°W). The 1.13 kg (male, 180.59 mm carapace width) individual was transported to The Maritime Aquarium at Norwalk (TMA) for identification. The staff at TMA identified it as M. magister, with some spots of shell disease, and reported the finding to the Connecticut Department of Energy and Environmental Protection (CT DEEP). In November 2018, another individual was caught by fisherman Kevin Scola and submitted to Massachusetts Division of Marine Fisheries. That individual was another male, measuring 217.68 mm carapace width, caught in Cape Cod Bay

Hudson et al. (2019), BioInvasions Records 8(2): 400–409, https://doi.org/10.3391/bir.2019.8.2.21 401 Metacarcinus magister and Eriocheir sinensis in the Long Island Sound

(42.03833°N; 70.38500°W). With concerns of confounding the specimens with the Atlantic rock crab irroratus, for which the authors have had a number of false alarms, the identification of these individuals was checked against Mary Wicksten’s (2011) most recent key for the Pacific Ocean. In April of 2018, a captain from Copps Island Oysters collected a Chinese mitten crab, E. sinensis, in an oyster dredge on the southern side of the Lot 12 lease in New Haven Harbor, Connecticut (41.248389°N; 72.934833°W). That individual, a male (56 g after freezing, 58.91 mm carapace width), was transported to Copps Island Oysters, in Norwalk, Connecticut and identified by Richard Harris as E. sinensis, who reported the finding to the CT DEEP. This specimen was submitted to the Yale Peabody Museum of Natural History in New Haven, CT (Specimen number: YPM IZ 103488); the M. magister specimen was accessioned in the Maritime Aquarium’s living collection (accession no. Norwalk-00348), but upon death was preserved and was also deposited at Yale Peabody Museum of Natural History (Specimen number: YPM IZ 103916).

Results and discussion The Long Island Sound were captured using commercial fishing gear in areas of high commercial shipping traffic (mid-Long Island Sound off Norwalk and New Haven, Figure 1C), and in an area of Cape Cod Bay that is an approved shellfishing area due west of Provincetown, Massachusetts (Figure 1B). The most likely vector of introduction of both species is the live trade, since port seizures of E. sinensis imports continue to happen every few years and M. magister can be found at markets both locally and throughout the United States. Humans have long been considered a vector for accidental introduction of invertebrates in the western North Atlantic via shipping (Carlton 2003; Chapman et al. 2003), but import of live animals for seafood and fishing bait is also an important documented vector for introductions in marine systems (Weigle et al. 2005; Fowler et al. 2016). Purchase of live animals at seafood markets and online, and their release in the wild, is probably how both of these crabs ended up in the Long Island Sound. Crustaceans that are involved in fisheries are easily the subject of illegal transportation, selling and potential release (Carlton 2003). The M. magister fishery is all male, making the likelihood of establishment of this species by the live seafood vector, barring a mistake, near zero. All the respective regulatory agencies of the states and provinces in the range of this (Alaska, British Columbia, Washington, Oregon, and California) do not permit anything but mature male fishery. Reflective of this vector, both the Long Island Sound (2017) and Cape Cod Bay (2018) specimens are male (Figure 2) and both of the individuals collected in Massachusetts in 2006 and 2009 were also male (Judith Pederson, Ph.D., MIT

Hudson et al. (2019), BioInvasions Records 8(2): 400–409, https://doi.org/10.3391/bir.2019.8.2.21 402 Metacarcinus magister and Eriocheir sinensis in the Long Island Sound

Figure 1. (A) Collection locations of introduced crabs. The two Dungeness crabs, M. magister, were collected in (B) Cape Cod Bay (triangle, 42.03833°N; 70.38500°W) and (C) in deeper water off of Norwalk (triangle, 41.014833°N; 73.394250°W). The non-native Chinese mitten crab, E. sinensis, was collected in (C) the Long Island Sound in New Haven Harbor (circle, 41.248389 °N, 72.934833°W).

Sea Grant College Program, pers. comm.). However, an additional concern with introduced species is not only what trouble they can cause in an adventive community, but also the diseases they may bring to the new area’s related species. Instances include the spread of bacterial gaffkaemia in , fungal “ plague” in Europe (Alderman 1996), and the rhizocephalan Loxothylacus panopaei (Gissler, 1884) in the western Atlantic (O’Shaughnessy et al. 2014). The parasites associated with marine bioinvasions are reviewed in Goedknegt et al. (2016), and have been identified as a critically understudied area deserving of greater attention (Lagrue 2017). The parasites and brood predators known from M. magister are summarized in Table 1, though the host specificity of many of these species is not fully

Hudson et al. (2019), BioInvasions Records 8(2): 400–409, https://doi.org/10.3391/bir.2019.8.2.21 403 Metacarcinus magister and Eriocheir sinensis in the Long Island Sound

Figure 2. (A) Dorsal and (B) ventral view of the M. magister male captured in the Long Island Sound. Photo by David Sigworth and The Maritime Aquarium at Norwalk; (C) dorsal and (D) ventral view of the M. magister male captured in Cape Cod Bay further north. Photos by David Hudson.

Table 1. Published parasite fauna of Metacarcinus magister. Species Type Reference Carcinonemertes errans Nemertea Wickham 1978 Microphallus similis Trematoda Ching 1991 Mesanophrys pugettensis Ciliophora Morado and Small 1994 Paranophrys sp. Ciliophora Armstrong et al. 1981 Ameson metacarcini Microsporida Small et al. 2014 Nadelspora canceri Microsporida Olson et al. 1994

known, complicating assessments of risk to native crustaceans in Long Island Sound and Cape Cod Bay. The Long Island Sound specimen was noted to have shell disease upon collection, which progressed to the state seen in Figure 2A and 2B until its death and preservation, and the live Cape Cod Bay specimen notably had early lesions as well. Spread from a localized invasion of E. sinensis in the Hudson River of New York (Schmidt et al. 2009) could be the origin of the specimen in New Haven Harbor (Figure 3) and of the individual found in 2012 in the Mianus River of Greenwich, Connecticut (CT DEEP 2012). However, with yet another instance of smuggling of this species recorded at a Cincinnati, Ohio (USA) express confinement facility in October 2018 (US CBP 2018), the possibility that these might alternatively be independent introductions from other smuggling events cannot be ruled out either. While the risk posed by parasites and brood predators of M. magister is relatively understudied, the potential dangers posed by the parasite and epibiont flora and fauna of E. sinensis are more evident. Chinese mitten crabs are known to harbor an

Hudson et al. (2019), BioInvasions Records 8(2): 400–409, https://doi.org/10.3391/bir.2019.8.2.21 404 Metacarcinus magister and Eriocheir sinensis in the Long Island Sound

Figure 3. (A) Dorsal and (B) ventral view of the E. sinensis male captured near New Haven. Photo by David Hudson.

array of potential pathogens, including a virus, two , five fungi/microsporidians, two protozoans, three worms, nine mite species, and a rhizocephalan, which are given a cursory overview in Table 2. Among these species there are two organisms of special concern, namely the zoonotic Paragonimus westermani, a lung fluke (Kuntz 1969), and Aphanomyces astaci, causative agent of “” (Benisch 1940; Schrimpf et al. 2014), the latter of which is well established as the etiological agent of numerous crustacean disease epidemics worldwide, including within the United States. The potential of E. sinensis to act as a

Hudson et al. (2019), BioInvasions Records 8(2): 400–409, https://doi.org/10.3391/bir.2019.8.2.21 405 Metacarcinus magister and Eriocheir sinensis in the Long Island Sound

Table 2. Published parasite fauna of Eriocheir sinensis. Species Type Reference sp. unk. Virus Wang and Gu 2002 Spiroplasma eriocherius* Bacteria Wang et al. 2005 sp. unk.** Bacteria Wang and Gu 2002 Septocylindrium eriocheir Fungus Alderman and Polgase 1985 Aphanomyces astaci Fungus Benisch 1940 Hepatospora eriocheir Microsporidia Wang and Chen 2007; Stentiford et al. 2011 Steinhausia mytilovum Microsporidia Peterson et al. 2011 Ichthyophonus hoferi Mesomycetozoea Sobecka et al. 2011 Epistylis sp. Ciliophora Sobecka et al. 2011 Branchiobdella sp. Annelida Sobecka et al. 2011 Trematoda sp. unk.*** Trematoda Sobecka et al. 2011 Paragonimus westermani Trematoda Kuntz 1969 Caspihalacarus hyrcanus Arachnida Normant et al. 2013 Copidognathus spp. Arachnida Normant et al. 2013 Halacarellus petiti Arachnida Normant et al. 2013 Porohalacarus alpinus Arachnida Normant et al. 2013 Soldanellonyx chappuisi Arachnida Normant et al. 2013 Soldanellox monardi Arachnida Normant et al. 2013 Hydrozetes lacustris Arachnida Normant et al. 2013 Trypocthoniellus longisetus Arachnida Normant et al. 2013 Piona pusilla Arachnida Normant et al. 2013 Polyascus gregarius Rhizocephala Li et al. 2011 * Etiological agent of “tremor disease” ** Rickettsia-like organism *** Encysted metacercariae

vector for A. astaci has global implications given the spread of the invasive crab to the Eastern Pacific, Europe, Canada, and the Mediterranean (Cohen and Carlton 1997; Streftaris and Zenetos 2006), and the demonstrated transmission of the disease to native crustaceans in Poland (Normant et al. 2007). The bacterium, Spiroplasma eriocherius, etiological agent of “tremor disease” is also a pathogen of concern, given that it is known to cause mass mortality in E. sinensis with characteristic neurological symptoms (Wang et al. 2005), and that species of Spiroplasma have also appeared in the crayfish, , causing systemic infections (Wang et al. 2005). The epifauna associated with crustaceans further compounds any assessment of ecological disturbance, as the diversity of epifaunal communities on crustaceans has been found to be surprisingly diverse (Dvoretsky and Dvoretsky 2010; Dražina et al. 2018), which portends an equally wide range of potential ecological impacts. Based on past records of disease, this potential range extension for E. sinensis into Long Island Sound would seem to have more immediate and alarming implications for the health of native crustaceans than the records of M. magister (which may not represent a viable population) in the Long Island Sound and Cape Cod Bay. However, the assessment of any non-native species requires consideration not just of the displaced organism, but rather the holobiont, i.e., the sum of all the associated microbial, commensal, epibiotic, and parasitic flora and fauna in our calculus of the risk of ecological disturbance.

Hudson et al. (2019), BioInvasions Records 8(2): 400–409, https://doi.org/10.3391/bir.2019.8.2.21 406 Metacarcinus magister and Eriocheir sinensis in the Long Island Sound

Both species have the potential to have broader ecosystem effects as predators and as ecosystem engineers. For instance, mesocosm work in Europe on E. sinensis revealed a broad range of impacts on freshwater macroinvertebrate communities (Rosewarne et al. 2016). Also, recent work on inducible defenses in prey show that M. magister can be quite effective in causing changes to invasive prey in its native range (Turner et al. 2017). Both of these examples show that these animals can be effective in altering food webs through their predatory behavior. A further concern with E. sinensis is its already well-documented physical cost, as in the example of the over 80 million Euros of damage since 1912 (Gollasch 2011) to the Danube River estuary by destabilizing the riverbanks. Continued invasion of these and other crustacean species clearly carry continued risk and costs to ecological systems in the areas where they occur.

Acknowledgements

The authors would like to thank partners and vessel crews at Copps Island Oysters for finding the two Connecticut crabs and bringing it to their attention, and to Ann Marie Lisi of The Maritime Aquarium’s education department for retrieving the animal for collection. Special thanks to fisherman Kevin Scola and Derek Perry, Marine Fisheries Biologist at the Massachusetts Division of Marine Fisheries for information about the Dungeness crab caught in Massachusetts. This work was supported through The Maritime Aquarium at Norwalk’s conservation ask program and the George S. and Pamela M. Humphrey Fund. Additionally, the authors would like to thank the anonymous reviewers and editor for their valuable comments on the manuscript.

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