Invasive Species Compendium

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Datasheet

Chama macerophylla
(leafy jewelbox)

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Datasheet

Chama macerophylla (leafy jewelbox)

Summary

  • Last modified
  • 24 March 2020
  • Datasheet Type(s)
  • Documented Species
  • Preferred Scientific Name
  • Chama macerophylla
  • Preferred Common Name
  • leafy jewelbox
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Mollusca
  •       Class: Bivalvia
  •         Subclass: Heterodonta
  • Summary of Invasiveness
  • Chama macerophylla is a common epifaunal chamid bivalve native to the western Atlantic (the Caribbean, the Gulf of Mexico, and the southern east coast of the United States). This species and several others of t...

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Pictures

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PictureTitleCaptionCopyright
Chama macerophylla (leafy jewelbox); habit, showing a dense aggregation on sub-tidal rocks within native range. Scout Key, Monroe County, Florida Keys, Florida, USA. April 2004. (Now in the collection of the Field Museum of Natural History, Chicago, FMNH 306132).
TitleHabit
CaptionChama macerophylla (leafy jewelbox); habit, showing a dense aggregation on sub-tidal rocks within native range. Scout Key, Monroe County, Florida Keys, Florida, USA. April 2004. (Now in the collection of the Field Museum of Natural History, Chicago, FMNH 306132).
Copyright©Rüdiger Bieler
Chama macerophylla (leafy jewelbox); habit, showing a dense aggregation on sub-tidal rocks within native range. Scout Key, Monroe County, Florida Keys, Florida, USA. April 2004. (Now in the collection of the Field Museum of Natural History, Chicago, FMNH 306132).
HabitChama macerophylla (leafy jewelbox); habit, showing a dense aggregation on sub-tidal rocks within native range. Scout Key, Monroe County, Florida Keys, Florida, USA. April 2004. (Now in the collection of the Field Museum of Natural History, Chicago, FMNH 306132).©Rüdiger Bieler
Chama macerophylla (leafy jewelbox); upper valve. Collected in 1987 from a floating polystyrene foam buoy in shallow water, within native range. nr Ohio Key, Monroe County, Florida Keys, Florida, USA. Largest specimen dimension, 57.6 mm. (Now in the collection of the Field Museum of Natural History, Chicago, FMNH 227408)
TitleUpper valve
CaptionChama macerophylla (leafy jewelbox); upper valve. Collected in 1987 from a floating polystyrene foam buoy in shallow water, within native range. nr Ohio Key, Monroe County, Florida Keys, Florida, USA. Largest specimen dimension, 57.6 mm. (Now in the collection of the Field Museum of Natural History, Chicago, FMNH 227408)
Copyright©Rüdiger Bieler
Chama macerophylla (leafy jewelbox); upper valve. Collected in 1987 from a floating polystyrene foam buoy in shallow water, within native range. nr Ohio Key, Monroe County, Florida Keys, Florida, USA. Largest specimen dimension, 57.6 mm. (Now in the collection of the Field Museum of Natural History, Chicago, FMNH 227408)
Upper valveChama macerophylla (leafy jewelbox); upper valve. Collected in 1987 from a floating polystyrene foam buoy in shallow water, within native range. nr Ohio Key, Monroe County, Florida Keys, Florida, USA. Largest specimen dimension, 57.6 mm. (Now in the collection of the Field Museum of Natural History, Chicago, FMNH 227408)©Rüdiger Bieler
Chama macerophylla (leafy jewelbox); lower (attached) valve. Collected in 1987 from a floating polystyrene foam buoy in shallow water, within native range. nr Ohio Key, Monroe County, Florida Keys, Florida, USA. Largest specimen dimension, 57.6 mm. (Now in the collection of the Field Museum of Natural History, Chicago, FMNH 227408)
TitleLower valve
CaptionChama macerophylla (leafy jewelbox); lower (attached) valve. Collected in 1987 from a floating polystyrene foam buoy in shallow water, within native range. nr Ohio Key, Monroe County, Florida Keys, Florida, USA. Largest specimen dimension, 57.6 mm. (Now in the collection of the Field Museum of Natural History, Chicago, FMNH 227408)
Copyright©Rüdiger Bieler
Chama macerophylla (leafy jewelbox); lower (attached) valve. Collected in 1987 from a floating polystyrene foam buoy in shallow water, within native range. nr Ohio Key, Monroe County, Florida Keys, Florida, USA. Largest specimen dimension, 57.6 mm. (Now in the collection of the Field Museum of Natural History, Chicago, FMNH 227408)
Lower valveChama macerophylla (leafy jewelbox); lower (attached) valve. Collected in 1987 from a floating polystyrene foam buoy in shallow water, within native range. nr Ohio Key, Monroe County, Florida Keys, Florida, USA. Largest specimen dimension, 57.6 mm. (Now in the collection of the Field Museum of Natural History, Chicago, FMNH 227408)©Rüdiger Bieler

Identity

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Preferred Scientific Name

  • Chama macerophylla Gmelin, 1791

Preferred Common Name

  • leafy jewelbox

Other Scientific Names

  • Chama citrea Gmelin, 1791
  • Chama imbricata Lamarck, 1801

Local Common Names

  • Bermuda: rock oyster

Summary of Invasiveness

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Chama macerophylla is a common epifaunal chamid bivalve native to the western Atlantic (the Caribbean, the Gulf of Mexico, and the southern east coast of the United States). This species and several others of the same genus readily settle on artificial substrata including ship hulls and have high invasive potential. C. macerophylla has been reported from attached specimens in Pacific island localities (Hawaii and Guam), from floating debris that had drifted across the Atlantic Ocean to the coasts of Ireland and the United Kingdom, and from barcoded larval samples in various Indo-Pacific locales. However, there are few reports of its becoming established outside its native range. Ecological impacts remain unstudied. The species is listed in the US NEMESIS database (Fofonoff et al., 2019) and has been placed on several watch lists in the Pacific Ocean (e.g. the Galapagos -- Keith et al., 2015) and the Atlantic Ocean (e.g. the genus is so listed in Wales -- Welsh Government, 2017).

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Mollusca
  •             Class: Bivalvia
  •                 Subclass: Heterodonta
  •                     Order: Veneroida
  •                         Unknown: Chamoidea
  •                             Family: Chamidae
  •                                 Genus: Chama
  •                                     Species: Chama macerophylla

Notes on Taxonomy and Nomenclature

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The small marine bivalve family Chamidae contains fewer than 70 living species, with the majority currently classified in the genus Chama Linnaeus, 1758 (MolluscaBase, 2019). The western Atlantic native Chama macerophylla Gmelin, 1791 has two accepted synonyms, C. citrea Gmelin, 1791 and C. imbricata Lamarck, 1801 (Campbell et al., 2004). Morphology-based species-level identification, especially of young specimens, is often difficult; the genus Chama and the closely related (and possibly synonymous) genus Pseudochama Odhner, 1917, are in need of comprehensive worldwide taxonomic review.

Description

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The circular to irregularly oval shell of Chama macerophylla is large for the genus (reaching more than 10 cm in greatest shell dimension) and has a protruding, weakly spiral umbonal region. The shell is consistently attached to the substratum by its left valve and usually has a very extensive attachment area. The attached valve is usually larger and more deeply cupped than the lid-like upper valve. The shell outside bears thin concentric fronds, lengthening into spine-like foliations toward the margin. Young specimens often resemble members of the bivalve genus Spondylus, with spine-like foliation that sometimes covers the entire shell; older specimens, especially in high-energy settings, are often eroded and may appear smooth. The distinctly visible radial grooves on most of the shell fronds are a key distinguishing feature of this species. The external shell coloration may be uniform or combinations of lemon-yellow, purplish or reddish-brown, orange, or white, with the umbones frequently differently coloured. The inner shell margins are finely crenulate. The interior colour is off-white or flushed with the colour of the exterior; in most cases with brownish purple stains at the ventral margin. For detailed morphological data and additional illustrations of this species see Campbell et al. (2004), Mikkelsen and Bieler (2007), Bieler et al. (2014), and Simone et al. (2015). Published morphological and molecular characterizations of this species are based near-exclusively on specimens from Florida. Specimens are often heavily encrusted with a diverse assemblage of fouling organisms (e.g. sponges, polychaetes and tunicates), and/or penetrated by borers (Mikkelsen and Bieler, 2007).

Distribution

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Native range: Chama macerophylla is widely distributed in the warm-temperate to tropical western Atlantic, where it also has an extensive fossil record since the Miocene (Campbell et al., 2004). Its extant range includes Bermuda, the U.S. coast from North Carolina south to the Gulf of Mexico, and the Caribbean. (Note that the range includes some states for which there are not specific records in the Distribution table). It has been stated to occur in Brazil (e.g. Rios, 2009), but Simone et al. (2015) found no specimens from that region with similar morphological features and excluded Brazilian records from the native range pending further studies.

Introduced range: Chama macerophylla has been introduced to Pearl Harbor, Hawaii (Coles et al., 1999a, b), reportedly from the Philippines (Coles et al., 1999b), although there do not appear to be records of its presence in the Philippines. It was introduced to Guam in 1999, but most individuals soon died and it was not expected to become established (Paulay et al., 2002; Paulay, 2003). There have been occasional reports of the species on floating debris in the north-east Atlantic (Fenwick, 2019), and DNA barcoding projects have inferred the presence of C. macerophylla larvae in various Indo-Pacific locales such as Saudi Arabia and French Polynesia (MGnify, 2018). Indo-Pacific records of this species in collection databases need individual verification, especially as this species name and its synonyms have been misapplied or confused with other species in this genus in the literature (see Campbell et al., 2004).

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Last updated: 06 Apr 2020
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Asia

PhilippinesPresentColes et al. (1999a)Described as source region for introduction in Hawaii

Europe

IrelandPresent, Few occurrencesIntroducedFenwick (2019)On floating debris

North America

Antigua and BarbudaPresentNativeZhang (2011)
BahamasPresentNativeRedfern (2013)
BelizePresentNativeMacintyre et al. (1982)
BermudaPresentNativeJensen and Pearce (2009)
Bonaire, Saint Eustatius and Saba PresentCABI (2020a)Present, based on regional distribution
-BonairePresentNativeHewitt and van Leeuwen (2017)
Costa RicaPresentNativeRobinson and Montoya (1987)
CubaPresentNativeTurgeon et al. (2009)
CuraçaoPresentNativevan Benthem Jutting (1927)
Dominican RepublicPresentNativeMassemin et al. (2009)
GuadeloupePresentNativeMassemin et al. (2009)
HaitiPresentNativeMassemin et al. (2009)
HondurasPresentNativeRosenberg G (2009)
JamaicaPresentNativeHumfrey M (1975)
MartiniquePresentNativeMassemin et al. (2009)
MexicoPresentNativeWiley et al. (1982)
PanamaPresentNativeBirkeland et al. (1976)
Puerto RicoPresentNativeWarmke and Abbott (1961)
United StatesPresent, LocalizedNative and IntroducedRay (2005); Turgeon et al. (2009); CABI (2020a)Native to Gulf of Mexico and east coast from North Carolina southwards; introduced to Hawaii.
-FloridaPresentNativeCampbell et al. (2004)
-GeorgiaPresentNativeWolfe (2008)
-HawaiiPresent, LocalizedIntroduced1992Coles et al. (1999); Coles et al. (1999a); Ray (2005); Fofonoff et al. (2019)Pearl Harbor, Oahu
-LouisianaPresentNativeGarcía and Lee (2002)
-North CarolinaPresentNativePorter (1989)
-TexasPresentNativeGarcía and Lee (2002)

Oceania

GuamAbsent, Formerly present20001999Paulay (2003)Arrived on a floating dock from Hawaii; most individuals died soon after arrival

Sea Areas

Atlantic - NortheastPresent, Few occurrencesIntroducedFenwick (2019)
Atlantic - Western CentralPresent, WidespreadNativeCampbell et al. (2004)
Pacific - Eastern CentralPresent, LocalizedFofonoff et al. (2019)Hawaii
Pacific - Western CentralPresent, LocalizedIntroduced1999Paulay (2003)

South America

BrazilAbsent, Invalid presence record(s)Rios (2009); Simone et al. (2015)
ColombiaPresentNativeDaccarett and Bossio (2011)
French GuianaPresentNativeMassemin et al. (2009)
GuyanaPresentNativeDiaz Merlano and Puyana Hegedus (1994)
VenezuelaPresentNativeMassemin et al. (2009)

History of Introduction and Spread

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The earliest published Hawaiian record dates from 1992, provided by Coles at al. (1999a, 1999b) who reported a Chama species introduced from the fouling community of a floating drydock towed to Hawaii from the Philippines in 1992; by 1996 the species was “abundant on the hull of the floating drydock […] and single specimens were found at three other stations” in Pearl Harbor (Coles et al., 1999b; see also Godwin, 2003; O’Connor et al., 2008; Clarke Murray, 2012). The species was initially tentatively identified as the Red Sea species Chama elatensis Delsaerdt, 1986, and then re-identified as the Caribbean C. macerophylla (G. Paulay in Eldredge and Smith, 2001). The stated Philippine source region in this context is of interest as C. macerophylla had not been reported as present in the Philippines. Subsequently, the species was transported from Hawaii to Guam on a floating dry dock in July 1999 and reported by Paulay et al. (2002) and Paulay (2003), who deemed it unlikely that it would become established in Guam as most introduced animals had died shortly after arrival.  There have been occasional reports of the species on floating debris in the north-east Atlantic (Fenwick, 2019).

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Hawaii Philippines 1992 Hitchhiker (pathway cause) Yes No Coles et al. (1999b); Fofonoff et al. (2019)
Guam Hawaii 1999 Hitchhiker (pathway cause) No No Paulay et al. (2002) Considered unlikely to become established

Risk of Introduction

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Chama macerophylla settles on ship hulls and other floating structures, and so can be readily transported.

Habitat

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Like all chamids, Chama macerophylla is a suspension feeder, living epifaunally and often in gregarious clumps on hard substrata such as rocks, dead coral, larger shells, and mangrove roots, as well as on artificial surfaces including metal (e.g. navigational markers, ship hulls, underwater platforms) and concrete (e.g. seawalls, pilings).  It has been reported from the lower intertidal zone to a depth of 525 m (Woodring, 1925), but verified deep records are based on empty shells, with confirmed live occurrences restricted to waters shallower than 50 m. It is primarily euhaline.

Chama macerophylla readily settles on and cements its shells to artificial substrata and has been reported from both fixed and floating surfaces. In its native range, Merrill (1963) reported it growing on metal buoys off Cumberland Island, Georgia, and it has been found on a floating polystyrene foam buoy in the Florida Keys (R. Bieler, Field Museum of Natural History, Chicago, Illinois, USA, unpublished observation). Anderson (2009) described it from oil and gas platforms on the Louisiana continental shelf, where it was the fourth most abundant species identified, and although individuals were not numerically abundant, the valves of both live and dead individuals of this species formed an important substratum for other members of the encrusting community. Bieler et al. (2017) found it to be a common member of the fouling community that develops on previously cleaned vessels that have been deliberately scuttled to serve as artificial reefs off the south Florida coast. In both states it is recorded on such substrates to a depth of about 30m.

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Marine
Inshore marine Principal habitat Natural
Coral reefs Principal habitat Natural

Biology and Ecology

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Reproductive Biology

The reproductive biology of Chama macerophylla remains largely unstudied, but chamids are generally considered gonochoristic (Yonge, 1967). Gametes and early fertilization were studied by Hylander and Summers (1977).

Nutrition

Chama macerophylla feeds by filtering a wide range of particles, such as algae, zooplankton and organic matter.

Associations

Chama macerophylla serves as a substratum for other members of the fouling community, including sponges, polychaetes, and vermetid gastropods.

Means of Movement and Dispersal

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Natural Dispersal

Natural dispersal of Chama macerophylla is passive, with veliger larvae transported from colonized areas through water currents.

Vector Transmission (Biotic)

Long distance dispersal by hitchhiking on other organisms is also possible: Frazier et al. (1985) reported specimens of Chama macerophylla on the carapace of Loggerhead Sea Turtles (Caretta caretta Linnaeus, 1758) off Georgia, USA.

Accidental Introduction

The most likely route of introduction of Chama macerophylla is by vessel fouling, as was established for the introductions in Hawaii and Guam. Within ocean basins, it can also be moved over long distances by floating debris. For instance, floating stone crab traps from Florida bearing several species of Chama (including, based on comparative shell size, attached valves of C. macerophylla) have been reported from the coast of Cornwall (United Kingdom). Another specimen of this species was reported from Ireland on floating plastic debris that likewise probably originated in the southern USA (Fenwick, 2019).

Intentional Introduction

This has not been reported, but the aquarium trade is an additional source for potential introductions. Chama macerophylla is a species that is sold (as an attached organism) with “wild”-collected or locally cultured 'live rock' for the aquarium trade (Mikkelsen and Bieler, 2007), and has become available for online purchases.

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Aquarium tradeComponent of “live rock” aquarium trade Yes Mikkelsen and Bieler, 2007
Hitchhiker Yes Yes

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Ship hull fouling Yes Yes

Impact: Economic

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None known except for costs of monitoring and vessel cleaning.

Impact: Environmental

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Impact on Habitat

Chama macerophylla is an early settler on artificial materials (e.g. Bieler et al., 2017) and provides calcareous substratum for other fouling species.

Impact on Biodiversity

Chama macerophylla is not known to have had any impact on biodiversity, although the related species C. pacifica Broderip, 1835 has become invasive in the Western Mediterranean via the Suez Canal and now outnumbers the native C. gryphoides Linnaeus, 1758 in that region (Zurel et al., 2012; Crocetta and Russo, 2013). The same invasive species and additional members of Chama have been reported as non-native in Hawaii (Carlton and Eldredge, 2009, 2015).

Impact: Social

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None known.

Risk and Impact Factors

Top of page Invasiveness
  • Has a broad native range
  • Abundant in its native range
  • Pioneering in disturbed areas
  • Gregarious
Impact mechanisms
  • Fouling
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Highly likely to be transported internationally deliberately
  • Difficult to identify/detect as a commodity contaminant
  • Difficult to identify/detect in the field

Uses

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Chama macerophylla is a species that is sold (as an attached organism) with “wild”-collected or locally cultured 'live rock' for the aquarium trade (Mikkelsen and Bieler, 2007).

Local fishermen on some of the Caribbean islands use it as line bait or chum (R. Bieler, Field Museum of Natural History, Chicago, Illinois, USA, unpublished observation).

Uses List

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Animal feed, fodder, forage

  • Bait/attractant

General

  • Pet/aquarium trade

Detection and Inspection

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Adult specimens of Chama macerophylla can be identified, but might need input by taxonomic specialists. Molecular barcoding data are available for this species, although not for many other Chama species.

Similarities to Other Species/Conditions

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The taxonomy of Chama is challenging and shell-based identifications, especially of juvenile specimens, often remain tentative. Chama macerophylla is now well characterized in its native range, with detailed anatomical data (Simone et al., 2015) and an extensive range of molecular markers from Florida populations on GenBank (e.g., as published by Sharma et al., 2012), the latter providing the basis for molecular barcoding of suspected invasive specimens. Relatively few of the approximately 55 other worldwide species of Chama and Pseudochama currently have molecular barcoding information available, and species-level morphology-based determination needs relevant taxonomic expertise.

Prevention and Control

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Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

No specific measures for prevention of the spread of Chama macerophylla have been documented.

Physical/Mechanical Control

According to Molnar et al. (2008) and Eldredge and Smith (2001), Chama macerophylla is “a notoriously stubborn fouling species; adults are very difficult to remove from ships’ hulls.”

Gaps in Knowledge/Research Needs

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Comprehensive taxonomic revision and additional molecular barcoding data are needed to reliably identify members of this group. Likewise, information on salinity tolerances, duration of planktonic veliger stages, and individual life spans are needed for all chamid species.

References

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Anderson LC, 2009. Molluscs: Mollusca from Oil and Gas Platforms on the Louisiana Continental Shelf: A Biotechnology Survey. In: Evaluation of oil and gas platforms on the Louisiana continental shelf for organisms with biotechnology potential [ed. by Rouse L]. New Orleans, Louisiana, USA: US Department of the Interior, Minerals Management Service, Gulf of Mexico OCS Region.37-42. https://espis.boem.gov/final%20reports/4881.pdf OCS Study MMS 2009-059

Bieler R, Granados-Cifuentes C, Rawlings TA, Sierwald P, Collins TM, 2017. Non-native molluscan colonizers on deliberately placed shipwrecks in the Florida Keys, with description of a new species of potentially invasive worm-snail (Gastropoda: Vermetidae). PeerJ, 5(3158), e3158. https://peerj.com/articles/3158/

Bieler R, Mikkelsen PM, Collins TM, Glover EA, González VL, Graf DL, Harper EM, Healy J, Kawauchi GY, Sharma PP, Staubach S, Strong EE, Taylor JD, Tëmkin I, Zardus JD, Clark S, Guzmán A, McIntyre E, Sharp P, Giribet G, 2014. Investigating the Bivalve Tree of Life -- an exemplar-based approach combining molecular and novel morphological characters. Invertebrate Systematics, 28(1), 32-115. http://www.publish.csiro.au/IS/IS13010

Birkeland, C, Reimer, AR, Young, JR, 1976. Survey of marine communities in Panama and experiments with oil (Ecological research series: EPA-600/3-76-028). Naragansett, Rhode Island, USA: Office of Research and Development, U.S. Environmental Protection Agency.xv+177 pp. https://play.google.com/books/reader?id=r4oMclKXXgsC&hl=en&pg=GBS.PP1

Campbell, MR, Steiner, G, Campbell, LD, Dreyer, H, 2004. Recent Chamidae (Bivalvia) from the western Atlantic Ocean. Malacologia, 46(2), 381-415. https://biodiversitylibrary.org/page/28112961

Carlton JT, Eldredge L, 2009. Marine bioinvasions of Hawaii: The introduced and cryptogenic marine and estuarine animals and plants of the Hawaiian archipelago, Honolulu, Hawaii, USA: Bishop Museum Press , 202 pp. Bishop Museum Bulletin in Cultural and Environmental Studies, 4

Carlton JT, Eldredge LG, 2015. Update and revisions of the marine bioinvasions of Hawai‘i: the Introduced and cryptogenic marine and estuarine animals and plants of the Hawaiian Archipelago. In: Bishop Museum Bulletin in Zoology,9 [ed. by Evenhuis NL, Carlton JT]. 25-47. http://www.malamamaunalua.org/wp-content/uploads/Carlton-and-Eldredge-2015.pdf [Lucius G. Eldredge Memorial Volume]

Clarke Murray CL, 2012. The role of recreational boating in the introduction and spread of marine invasive species. Vancouver, Canada: University of British Columbia. xiv + 177 pp. https://open.library.ubc.ca/cIRcle/collections/ubctheses/24/items/1.0053492 [PhD thesis]

Coles, SL, DeFelice, RC, Eldredge, LG, 1999. Nonindigenous marine species introductions in the harbors of the south and west shores of Oahu, Hawaii. Bishop Museum Technical Report No. 15. Honolulu, Hawaii, USA: Bernice Pauahi Bishop Museum and Hawaii Biological Survey.x + 212 pp. http://hbs.bishopmuseum.org/pdf/southshore.pdf

Coles, SL, DeFelice, RC, Eldredge, LG, Carlton, JT, 1999. Historical and recent introductions of nonindigenous marine species into Pearl Harbor, Oahu, Hawaiian Islands. Marine Biology, 135(1), 147-158. https://doi.org/10.1007/s002270050612 doi: 10.1007/s002270050612

Crocetta, F., Russo, P., 2013. The alien spreading of Chama pacifica Broderip, 1835 (Mollusca: Bivalvia: Chamidae) in the Mediterranean Sea. Turkish Journal of Zoology, 37(1), 92-96. http://journals.tubitak.gov.tr/zoology/

Daccarett, EY, Bossio, VS, 2011. Colombian Seashells from the Caribbean Sea, L'Informatore Piceno.384 + 16 pp.

Diaz Merlano, JM, Puyana Hegedus, M, 1994. Moluscos marinos del Caribe Colombiano: un catálogo ilustrado, Colombia: Invemar, Fundación Natura, Colciencias.291 pp. + 78 plates.

Eldredge LG, Smith CM, 2001. A guidebook of introduced marine species in Hawaii. vi + 60 pp. http://www2.bishopmuseum.org/HBS/invertguide/species_pdf/guide.pdf [Bishop Museum Technical Report, 21. Contribution No. 2001-005 to the Hawai'i Biological Survey]

Fenwick, D, 2019. Aphotomarine. http://www.aphotomarine.com/

Fofonoff, PW, Ruiz, GM, Steves, B, Simkanin, C, Carlton, JT, 2019. National Exotic Marine and Estuarine Species Information System. Edgewater, Maryland, USA: Smithsonian Environmental Research Center.http://invasions.si.edu/nemesis/

Frazier, J., Margaritoulis, D., Muldoon, K., Potter, C. W., Rosewater, J., Ruckdeschel, C., Salas, S., 1985. Epizoan communities on marine turtles I. Bivalve and gastropod mollusks. Marine Ecology, 6(2), 127-140.

García, EF, Lee, HG, 2002. Report on molluscan species found in the offshore waters of Louisiana, including many extensions of known range and un-named species. American Conchologist, 30(4), 10-13.

Godwin, L. S., 2003. Hull fouling of maritime vessels as a pathway for marine species invasions to the Hawaiian Islands. Biofouling, 19(Suppl), 123-131. https://doi.org/10.1080/0892701031000061750

Hewitt, SJ, van Leeuwen, S, 2017. Marine molluscs from the island of St. Eustatius, Eastern Caribbean: results of the Statia Marine Biodiversity Expedition 2015. Vita Malacologica, 16(annual issue), 44-80.

Humfrey M, 1975. Sea shells of the West Indies: a guide to the marine molluscs of the Caribbean, Taplinger Publishing Company.351 pp.

Hylander BL, Summers RG, 1977. An ultrastructure analysis of the gametes and early fertilization in two bivalve mollusks, Chama macerophylla and Spisula solidissima with special reference to gamete binding. Cell and Tissue Research, 182(4), 469-489. https://link.springer.com/article/10.1007%2FBF00219830

Jensen, RH, Pearce, TA, 2009. Marine mollusks of Bermuda: checklist and biography, Wilmington, Delaware, USA: Delaware Museum of Natural History.x + 473 pp.

Keith I, Dawson T, Collins K, Banks S, 2015. Marine invasive species in the Galapagos Marine Reserve: A case for additional research, improved management, and policy review. In: Galapagos Report 2013-2014 . Puerto Ayora, Galapagos, Ecuador: GNPD, GCREG, CDF and GC.83-88. https://www.galapagos.org/wp-content/uploads/2015/09/GalapagosReport_2013-2014-12-Keith-83-88.pdf

Macintyre, IG, Rützler, K, Norris, JN, Fauchald, K, 1982. A submarine cave near Columbus Cay, Belize: A bizarre cryptic habitat. In: The Atlantic barrier reef ecosystem at Carrie Bow Cay, Belize, I. Structure and communities. (Smithsonian Contributions to Marine Science, 12), [ed. by Rützler, K, Macintyre, IG]. Washington, DC, USA: Smithsonian Institution. 127-141. https://repository.si.edu/handle/10088/1116

Massemin, D, Lamy, D, Pointier, JP, Gargominy, O, 2009. Coquillages et escargots de Guyane, Mèze, France: Biotope.456 pp. http://www.biotope-editions.com/index.php?article98/coquillages-et-escargots-de-guyane-seashells-and-snails-from-french-guiana

Merrill AS, 1963. Mollusks from a buoy off Georgia. Nautilus, 77(2), 68-70.

MGnify, 2018. Amplicon sequencing of Tara Oceans DNA samples corresponding to size fractions for protists. https://doi.org/10.15468/2hv1be

Mikkelsen PM, Bieler R, 2007. Seashells of Southern Florida - Living Marine Mollusks of the Florida Keys and Adjacent Regions: Bivalves, Princeton, New Jersey, USA: Princeton University Press.x + 503 pp. .

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van Benthem Jutting, T, 1927. Marine molluscs of the island of Curaçao. Bijdragen tot de Dierkunde, 25(1), 1-36. https://www.repository.naturalis.nl/document/548200

Warmke, GL, Abbott, RT, 1961. Caribbean seashells, Narberth, Pennsylvania, USA: Livingston Publishing Co.xx + 348 pp.

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Wiley, GN, Circé, RC, Tunnell, Jr JW, 1982. Mollusca of the rocky shores of east central Veracruz State, Mexico. The Nautilus, 96(2), 55-61.

Wolfe, DA, 2008. Mollusks taken by beam trawl in the vicinity of Gray’s Reef National Marine Sanctuary on the continental shelf off Georgia, southeastern U.S. (NOAA Technical Memorandum NOS NCCOS, 88). USA: National Oceanic and Atmospheric Administration.iv + 42 pp. http://aquaticcommons.org/2083/1/wolfeGRNMS.pdf

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Distribution References

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Macintyre IG, Rützler K, Norris JN, Fauchald K, 1982. A submarine cave near Columbus Cay, Belize: A bizarre cryptic habitat. In: The Atlantic barrier reef ecosystem at Carrie Bow Cay, Belize, I. Structure and communities. (Smithsonian Contributions to Marine Science, 12). [ed. by Rützler K, Macintyre IG]. Washington, DC, USA: Smithsonian Institution. 127-141. https://repository.si.edu/handle/10088/1116

Massemin D, Lamy D, Pointier JP, Gargominy O, 2009. Coquillages et escargots de Guyane. Mèze, France: Biotope. 456 pp. http://www.biotope-editions.com/index.php?article98/coquillages-et-escargots-de-guyane-seashells-and-snails-from-french-guiana

Paulay G, 2003. Marine Bivalvia (Mollusca) of Guam. Micronesica. 35-36 (no issue number), 218-243.

Porter HJ, 1989. Continental Shelf molluscan zoogeography of North and South Carolina. In: North Carolina Coastal Oceanography Symposium, Wilmington, North Carolina, 30 September - 2 October 1987 (NOAA-National Undersea Research Program Research Report, 89-2) [North Carolina Coastal Oceanography Symposium, Wilmington, North Carolina, 30 September - 2 October 1987 (NOAA-National Undersea Research Program Research Report, 89-2)], [ed. by George RY, Hulbert AW]. Rockville, Maryland, USA: National Oceanographic and Atmospheric Administration. 231-278.

Ray GL, 2005. Invasive marine and estuarine animals of Hawai’i and other Pacific islands (ERDC/TN ANSRP-05-3)., 19 pp. https://apps.dtic.mil/dtic/tr/fulltext/u2/a437988.pdf

Redfern C, 2013. Bahamian Seashells: 1161 Species from Abaco, Bahamas. Boca Raton, Florida, USA: Bahamianseashells.com, Inc. 501 pp. http://www.bahamianseashells.com/

Rios EC, 2009. Compendium of Brazilian sea shells. Rio Grande, Brazil: Universidade Federal do Rio Grande, Museu Oceanográfico Prof. Eliézer de Carvalho Rios. viii + 668 pp.

Robinson DG, Montoya M, 1987. The marine molluscs of the Atlantic coast of Costa Rica. (Los moluscos marinos de la Costa Atlántica de Costa Rica). Revista de Biologia Tropical. 35 (2), 375-400. https://www.researchgate.net/publication/267212718_Los_moluscos_marinos_de_la_Costa_Atlantica_de_Costa_Rica

Rosenberg G, 2009. Malacolog 4.1.1: A Database of Western Atlantic Marine Mollusca., Philadelphia, Pennsylvania, USA: The Academy of Natural Sciences of Drexel University. http://www.malacolog.org/

Simone LRL, Mikkelsen PM, Bieler R, 2015. Comparative anatomy of selected marine bivalves from the Florida Keys, with notes on Brazilian congeners (Mollusca: Bivalvia). Malacologia. 58 (1-2), 1-127. http://dx.doi.org/10.4002/040.058.0201 DOI:10.4002/040.058.0201

Turgeon DD, Lyons WG, Mikkelsen P, Rosenberg G, Moretzsohn F, 2009. Bivalvia (Mollusca) of the Gulf of Mexico. In: Gulf of Mexico origin, waters, and biota: Volume 1, Biodiversity, [ed. by Felder DL, Camp DK]. Texas, USA: Texas A&M University Press. 711-744.

van Benthem Jutting T, 1927. Marine molluscs of the island of Curaçao. Bijdragen tot de Dierkunde. 25 (1), 1-36. https://www.repository.naturalis.nl/document/548200

Warmke GL, Abbott RT, 1961. Caribbean seashells. Narberth, Pennsylvania, USA: Livingston Publishing Co. xx + 348 pp.

Wiley GN, Circé RC, Tunnell Jr JW, 1982. Mollusca of the rocky shores of east central Veracruz State, Mexico. The Nautilus. 96 (2), 55-61.

Wolfe DA, 2008. Mollusks taken by beam trawl in the vicinity of Gray’s Reef National Marine Sanctuary on the continental shelf off Georgia, southeastern U.S. (NOAA Technical Memorandum NOS NCCOS, 88)., USA: National Oceanic and Atmospheric Administration. iv + 42 pp. http://aquaticcommons.org/2083/1/wolfeGRNMS.pdf

Zhang DY, 2011. Antiguan shallow-water seashells. Wellington, Florida, USA: MdM Publishing. 210 pp.

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05/08/19: Original text by:

Rüdiger Bieler, Field Museum of Natural History, Chicago, Illinois, USA

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