Chama macerophylla (leafy jewelbox)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Biology and Ecology
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Impact: Economic
- Impact: Environmental
- Impact: Social
- Risk and Impact Factors
- Uses List
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Distribution Maps
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PicturesTop of page
IdentityTop of page
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 InvasivenessTop of page
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 TreeTop of page
- 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 NomenclatureTop of page
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.
DescriptionTop of page
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).
DistributionTop of page
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 TableTop of page
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|
|Philippines||Present||Coles et al. (1999a)||Described as source region for introduction in Hawaii|
|Ireland||Present, Few occurrences||Introduced||Fenwick (2019)||On floating debris|
|Antigua and Barbuda||Present||Native||Zhang (2011)|
|Belize||Present||Native||Macintyre et al. (1982)|
|Bermuda||Present||Native||Jensen and Pearce (2009)|
|Bonaire, Saint Eustatius and Saba||Present||CABI (2020a)||Present, based on regional distribution|
|-Bonaire||Present||Native||Hewitt and van Leeuwen (2017)|
|Costa Rica||Present||Native||Robinson and Montoya (1987)|
|Cuba||Present||Native||Turgeon et al. (2009)|
|Curaçao||Present||Native||van Benthem Jutting (1927)|
|Dominican Republic||Present||Native||Massemin et al. (2009)|
|Guadeloupe||Present||Native||Massemin et al. (2009)|
|Haiti||Present||Native||Massemin et al. (2009)|
|Honduras||Present||Native||Rosenberg G (2009)|
|Jamaica||Present||Native||Humfrey M (1975)|
|Martinique||Present||Native||Massemin et al. (2009)|
|Mexico||Present||Native||Wiley et al. (1982)|
|Panama||Present||Native||Birkeland et al. (1976)|
|Puerto Rico||Present||Native||Warmke and Abbott (1961)|
|United States||Present, Localized||Native and Introduced||Ray (2005); Turgeon et al. (2009); CABI (2020a)||Native to Gulf of Mexico and east coast from North Carolina southwards; introduced to Hawaii.|
|-Florida||Present||Native||Campbell et al. (2004)|
|-Hawaii||Present, Localized||Introduced||1992||Coles et al. (1999); Coles et al. (1999a); Ray (2005); Fofonoff et al. (2019)||Pearl Harbor, Oahu|
|-Louisiana||Present||Native||García and Lee (2002)|
|-North Carolina||Present||Native||Porter (1989)|
|-Texas||Present||Native||García and Lee (2002)|
|Guam||Absent, Formerly present||2000||1999||Paulay (2003)||Arrived on a floating dock from Hawaii; most individuals died soon after arrival|
|Atlantic - Northeast||Present, Few occurrences||Introduced||Fenwick (2019)|
|Atlantic - Western Central||Present, Widespread||Native||Campbell et al. (2004)|
|Pacific - Eastern Central||Present, Localized||Fofonoff et al. (2019)||Hawaii|
|Pacific - Western Central||Present, Localized||Introduced||1999||Paulay (2003)|
|Brazil||Absent, Invalid presence record(s)||Rios (2009); Simone et al. (2015)|
|Colombia||Present||Native||Daccarett and Bossio (2011)|
|French Guiana||Present||Native||Massemin et al. (2009)|
|Guyana||Present||Native||Diaz Merlano and Puyana Hegedus (1994)|
|Venezuela||Present||Native||Massemin et al. (2009)|
History of Introduction and SpreadTop of page
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).
IntroductionsTop of page
|Introduced to||Introduced from||Year||Reason||Introduced by||Established in wild through||References||Notes|
|Natural reproduction||Continuous 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 IntroductionTop of page
Chama macerophylla settles on ship hulls and other floating structures, and so can be readily transported.
HabitatTop of page
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 ListTop of page
|Inshore marine||Principal habitat||Natural|
|Coral reefs||Principal habitat||Natural|
Biology and EcologyTop of page
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).
Chama macerophylla feeds by filtering a wide range of particles, such as algae, zooplankton and organic matter.
Chama macerophylla serves as a substratum for other members of the fouling community, including sponges, polychaetes, and vermetid gastropods.
Means of Movement and DispersalTop of page
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.
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).
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 CausesTop of page
Impact: EconomicTop of page
None known except for costs of monitoring and vessel cleaning.
Impact: EnvironmentalTop of page
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: SocialTop of page
Risk and Impact FactorsTop of page Invasiveness
- Has a broad native range
- Abundant in its native range
- Pioneering in disturbed areas
- 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
UsesTop of page
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 ListTop of page
Animal feed, fodder, forage
- Pet/aquarium trade
Detection and InspectionTop of page
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/ConditionsTop of page
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 ControlTop of page
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.
Gaps in Knowledge/Research NeedsTop of page
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.
ReferencesTop of page
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05/08/19: Original text by:
Rüdiger Bieler, Field Museum of Natural History, Chicago, Illinois, USA
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