expanding our understanding of the trade in marine aquarium … · 2017-01-26 · submitted 12 june...

Submitted 12 June 2015Accepted 30 December 2016Published 26 January 2017

Corresponding authorAndrew L Rhyne arhynerwuedu

Academic editorRobert Toonen

Additional Information andDeclarations can be found onpage 33

DOI 107717peerj2949

Copyright2017 Rhyne et al

Distributed underCreative Commons CC-BY 40

OPEN ACCESS

Expanding our understanding of thetrade in marine aquarium animalsAndrew L Rhyne12 Michael F Tlusty23 Joseph T Szczebak4 andRobert J Holmberg3

1Biology and Marine Biology Roger Williams University Bristol RI United States2Anderson Cabot Center for Ocean Life New England Aquarium Boston MA United States3 School for the Environment University of Massachusetts Boston Boston MA United States4Center for Economic and Environmental Development Roger Williams University Bristol RI United States

ABSTRACTThe trade of live marine animals for home and public aquaria has grown into amajor global industry Millions of marine fishes and invertebrates are removed fromcoral reefs and associated habitats each year The majority are imported into theUnited States with the remainder sent to Europe Japan and a handful of othercountries Despite the recent growth and diversification of the aquarium trade to datedata collection is not mandatory and hence comprehensive information on speciesvolume and diversity is lacking This lack of information makes it impossible to studytrade pathways Without species-specific volume and diversity data it is unclear howimporting and exporting governments can oversee this industry effectively or howsustainability should be encouraged To expand our knowledge and understandingof the trade and to effectively communicate this new understanding we introduce thepublically-available Marine Aquarium Biodiversity and Trade Flow online database(httpswwwaquariumtradedataorg) This tool was created to communicate thevolume and diversity of marine fishes andor invertebrates imported into the USover three complete years (2008 2009 and 2011) and three partial years (2000 20042005) To create this tool invoices pertaining to shipments of live marine fishes andinvertebrates were scanned and analyzed for species name species quantities country oforigin port of entry and city of import destination Here we focus on the analysis of thelater three years of data and also produce an estimate for the entirety of 2000 2004 and2005 The three-year aggregate totals (2008 2009 2011) indicate that just under 2300fish and 725 invertebrate species were imported into the US cumulatively althoughjust under 1800 fish and 550 invertebrate species were traded annually Overall thetotal number of live marine animals decreased between 2008 and 2011 In 2008 2009and 2011 the total number of individual fish (82 73 and 69 million individuals) andinvertebrates (42 37 and 36 million individuals) assessed by analyzing the invoicedata are roughly 60 of the total volumes recorded through the Law EnforcementManagement Information System (LEMIS) dataset Using these complete years weback-calculated the number of individuals of both fishes and invertebrates importedin 2000 2004 and 2005 These estimates (93 108 and 112 million individual fishper year) were consistent with the three years of complete data We also use thesedata to understand the global trade in two species (Banggai cardinalfish Pterapogonkauderni and orange clownfish Amphiprion ocellaris percula) recently considered forEndangered Species Act listing Aquariumtradedataorg can help create more effective

How to cite this article Rhyne et al (2017) Expanding our understanding of the trade in marine aquarium animals PeerJ 5e2949 DOI107717peerj2949

management plans for the traded species and ideally could be implemented at key tradeports to better assess the global trade of aquatic wildlife

Subjects Marine Biology Science PolicyKeywords Marine aquarium trade Wildlife trade Coral reef Data visualization

INTRODUCTIONThe live aquarium fish trade faces a multitude of potential threats including incidences ofreduced biodiversity from over-extraction habitat destruction in some source countries(Francis-Floyd amp Klinger 2003 Gopakumar amp Ignatius 2006) and negative impacts ofspecies invasions in the US and elsewhere (Chucholl 2013 Garciacutea-Berthou 2007Holmberget al 2015 Padilla amp Williams 2004) Despite these threats the aquarium trade can be aunique and significant positive force in reef-side communities (Rhyne Tlusty amp Kaufman2014) Benefits include but are not limited to saving threatened species from the brinkof extinction through the development of captive breeding programs (Tlusty 2002) andcatalyzing habitat preservation through sustainable supply-side practices (Firchau Dowd ampTlusty 2013) These sustainable practices include stewardship mechanisms for sustainablelivelihoods via poverty alleviation and the protection of threatened ecosystems (RhyneTlusty amp Kaufman 2014) Finally consumer education of aquarium trade sustainabilitycan promote widespread public appreciation for the worldrsquos aquatic ecosystems with theultimate goal of minimizing negative impacts of this trade (Tlusty et al 2013) While aproactive stance can transform a large consumer base into a powerful agent for biodiversityconservation increased sustainability and human well-being inaction will likely amplifythe deleterious threats currently faced by the trade Currently the lack of oversight leadingto a poor concept of trade volume and subsequent regulatory inefficiency has greatlyhampered the development of a sustainable industry Increasing the sustainability of themarine aquarium animal industry should be considered a primary initiative for the entireaquarium industry transport chain (Tlusty et al 2013) Increasing the sustainability ofthe transport chain of marine aquarium animals is achieved through a more thoroughunderstanding of the magnitude of the trade (Fujita et al 2014) which begins by assessingthe scale of imports into the US (the primary destination) (Rhyne et al 2012) Once theannual volume of US imports is gauged other relevant issues that lead to environmentaland economic benefits can then be tackled including animal quality and shipping survival

There is no clear picture of the number of live marine fish and invertebrate species orindividuals involved in the aquarium trade primarily due to insufficient global trackingof the import and export of these animals (Bruckner 2001 Fujita et al 2014 Green 2003Lunn amp Moreau 2004 Tissot et al 2010 Wabnitz et al 2003) Multiple sources of datahave been used to monitor this trade (Green 2003 Smith et al 2008 Wabnitz et al 2003Wood 2001) However not all of these data systems are sufficient for or were even intendedfor monitoring the aquarium trade For example compulsory data are maintained underfederal mandates for species listed by the Convention on International Trade in Endangered

Rhyne et al (2017) PeerJ DOI 107717peerj2949 236

Species of Wild Fauna and Flora (CITES) However previous studies found that CITESrecords were inaccurate incomplete or insufficient (Bickford et al 2011 Blundell ampMascia 2005 Rhyne et al 2012) Furthermore CITES-listed species (namely stony coralsgiant clams and seahorses) account for only a fraction of the total trade in live aquaticanimals Only a handful of studies (Rhyne et al 2012 Smith et al 2009 Smith et al 2008)have attempted to quantify the movement of non-CITES-listed aquarium species fromsource to market The Global Marine AquariumDatabase (GMAD) encouraged companiesto provide data on the marine aquarium trade (Green 2003) and until now GMAD wasthe only source for aquarium trade data recorded at a species-specific level While GMADcontains trade data from 1988 to 2003 some years and countries (eg Haiti) are missingand values reported in GMAD were found to be drastically fewer than other estimates oftrade data (Murray et al 2012) Furthermore the aquarium trade has been transformedby new technologies and husbandry breakthroughs since 2003 (Rhyne amp Tlusty 2012)which may dilute the relevance of this data set In addition to CITES and GMAD the LawEnforcement Management Information System (LEMIS) database has been used to betterunderstand the aquarium trade The United States Fish and Wildlife Service (USFWS)inspects wildlife shipments and through LEMIS maintains species-specific data in the caseof transporting CITES-listed species per CITES requirements However non-CITES-listedspecies are recorded with general codes (eg marine aquarium tropical fish regardless ofspecies are coded MATF) Recording data in this generalized manner eliminates specificinformation regarding the diversity and volumes of species traded (Smith et al 2009)While the need for accurate accounts of aquarium trade flow continually increases thecurrent monitoring methods remain static (Bickford et al 2011) The lack of specific datasystems for recording all species exported and imported for the marine aquarium traderaises two main concerns (1) because of the lack of trade data it is unclear how importingand exporting governments can monitor this industry effectively (2) it is also unclear howsustainability should be encouraged given the paucity of data

To date Balboa (2003) Wabnitz et al (2003) and Rhyne et al (2012) have cataloguedspecies-specific information provided on trade invoices Rhyne et al (2012) furthercompared invoice information to associated shipment declarations It was this effortthat led to the development of the Marine Aquarium Biodiversity and Trade Flow onlinedatabase (httpswwwaquariumtradedataorg) a public portal offering anonymized livemarine animal trade data collected through trade invoices Here we describe three years(2008 2009 2011) of fish and invertebrate invoice-based data from US imports that wereanalyzed for country of origin port of entry and quantity of species and individualsassociated with each port We also relate the findings from these three years of completetrade data to previously existing invoice-based trade data from the LEMIS databaseSpecifically Rhyne et al (2012) described one contiguous year of import data based on a12-month period from June 2004 (sevenmonths) untilMay 2005 (fivemonths) and Balboa(2003) described one month (October) of data from 2000 To address the missing monthsof data from these years and to increase the scope of the dataset we modeled data for themissing months based on the monthly volume patterns analyzed from the three completeyears of data available (2008 2009 2011) This work provides an enlarged snapshot of the


volume biodiversity and trade pathways for live marine aquarium fish and invertebratespecies beyond the information provided by other reporting systems (Wabnitz et al2003) Further this work demonstrates that LEMIS while well-designed for importexportcompliance and management of USFWS staffing needs is not designed to monitor thespecies-level activity of the live marine aquarium trade Finally we present two casestudies (the Banggai cardinalfish Pterapogon kauderni and the commonorange clownfishAmphiprion ocellarispercula) to demonstrate the use of these data for better understandingthe trade in marine species a first step toward advancing industry sustainability

METHODSThe goal of this project was to evaluate the number of live marine aquarium speciesimported into the US and to create a trade path analysis of the diversity of animalsinvolved in the trade The methods used to analyze trade invoices were described byRhyne et al (2012) and are briefly summarized here We reviewed all USFWS shipmentdeclarations and the attached commercial invoices coded as Marine Aquarium TropicalFish (MATF) for 2008 2009 and 2011 as indicated in the LEMIS database Whileabout 22000 invoices were marked as containing MATF in the LEMIS database wereceived approximately 20000 shipment declarations and their attached invoices CITESdata are specially coded on import declarations and thus are not integrated into thisdatabase

Invoices were considered a true statement of shipping contents as we were not able toassess the veracity of the information contained on the invoice Shipment information (dateport of origin and destination port) was collected from the declaration page and speciesand quantity information was tabulated from the associated invoices and catalogued into adatabase Within the trade of live marine organisms re-export information is not recordedand thus could not be assessed here Both manual entry and automated optical characterrecognition (OCR) software (ABBYY FlexiCapture 90) customized for wildlife shipments(Fig 1) were utilized to retrieve the above information from these documents The inputmethod varied with invoice quality and length Manual entry was utilized when invoiceswere of poor quality (blurry speckled darkened fonts less than six point handwritten) orbrief (less than 12 page) whereas all others were read using the OCR software Once allnecessary data were captured species names were verified using World Register of MarineSpecies (WoRMS Editorial Board 2015) FishBase (Froese amp Pauly 2015) and the primaryliterature (Appeltans et al 2011 Froese amp Pauly 2011) We corrected species informationonly when species names were misspelled listed under a junior synonym or listed byonly a common name Species were identified to the greatest taxonomic detail availableOccasionally genera were listed without species (eg lsquoChrysiptera sprsquo) or the specieswas otherwise ambiguous (eg lsquohybrid Acanthurus tangrsquo) In these cases the genus (andall higher-level taxonomic information) would be recorded and the species would berecorded as unknown Genus and species were recorded as unknown when listed onlyby an ambiguous common name (eg lsquocolorful damselrsquo lsquounknown damselrsquo lsquoassorteddamselsrsquo)


Figure 1 FlexiCapture 90 verification screen used to capture shipping data for the wwwaquariumtradedataorg database (A) Shipping decla-ration (B) shipping invoice (C) invoice table produced from Optimal Character Recognition (OCR) software Note grey shaded cells indicate au-tocorrected fields and red flags within cells indicate errors for user to correct manually

To determine if the volume of captive-bred P kauderni imported into the US hasincreased in recent years we reviewed invoice data from Los Angeles-based importerQuality Marine for two additional recent years of imports Quality Marine sourced fishfrom Thailand and these were known to have been captive-bred given that Thailandlies outside the native range of P kauderni At our request all shipments of MATF fromThailand to Quality Marine over the period of March 2012 to July 2014 were supplied andreviewed

In accordance with Rhyne et al (2012) this report focused on major geographic tradeflows the frequency of invoice detail to species-level and how invoice data compared toLEMIS data Invoice data for both fishes and invertebrates were retrieved concurrentlyTo help organize and visualize the trade data a publically accessible representation of thetrade data was created the Marine Aquarium Biodiversity and Trade Flow online database(httpswwwaquariumtradedataorg) This web-based graphical user interface poweredby the open source JavaScript library D3 (httpd3jsorg) is data-rich visually appealingand allows users to query more than 28000 invoices containing over 27 million lines ofinvoice data from 2000 2004 2005 2008 2009 and 2011

To back-calculate the estimated total annual imports of marine animals during yearswith incomplete data sets (years 2000 2004 2005) we first determined the proportion ofindividuals imported during the time interval (one month for 2000 sevenmonths for 2004and five months for 2005) based on the three years for which we had a complete 12-monthdataset (2008 2009 and 2011) For these three years there was variation between monthsbut the intra-month variation was less than that of the inter-month variation suggesting


that monthly import volumes were proportionally consistent This proportion was thenused to calculate the number of individuals for the unknown months using the followingrelation

(nPr)minusn

where n is the known number of imports per year for 1 (2000) 5 (2004) or 7 (2005)months and Pr is the average proportion of known imports from corresponding monthsfrom 2008 2009 and 2011 This estimated number of animals was then allocated across theunknown months proportionally for 2000 2004 and 2005 This method appears robustgiven the consistency of the most voluminous species across years (see results) We alsogenerated estimates for the source countries and ports of entry The database user canselect the type of data to view and within the selection tool can select to view actual dataor whether estimates should be included Even though these numbers are estimates wepresent them as numbers so that they coalesce with the complete database

RESULTSAll data presented in this report are available at theMarineAquariumBiodiversity andTradeFlow online database (httpswwwaquariumtradedataorg) This site was developed toallow users to generate database queries using dropdownmenus On the lsquolsquoHomersquorsquo tab initialqueries can be filtered through large-scale source areas such as ocean basins or countries oforigin for a defined time period (Fig 2) Following user selections the software compilesdetailed information in the form of maps timeline charts and other data charts that allowusers to access data at a depth uncommon in user interfaces for the wildlife or seafoodtrades On further analysis it is possible using the lsquolsquoSpeciesrsquorsquo tab to query a single taxonomicfamily genus or species for one or more countries andor ports of entry The user-friendlydropdown menus are tree-based and progressive Figure 3 demonstrates successive screenswhere the user has successively selected the family Pomacentridae the genus Amphiprionand the species complex Amphiprion perculaocellaris The dashboard displays (A) adistribution map depicting the relative geographic importance using proportionally-sizedred dots and (B C) two graphs displaying export country- and port of entry-specificvolumes for the selected query To enhance the utility of the website and promote thedissemination of the data the user can download charts and graphs of data queries Userscan also share these charts directly to Facebook and Twitter (Fig 4) Further to ensure thedata within the invoice-based database is an accurate representation of the trade users canreport possible errors in data or features on the website If users find species that are likelyincorrect in distribution or taxa we can examine the invoice record verify its contents andupdate the database if needed This system also logs how users interact with the databasewhich provides feedback on the number and types of queries users generated

General trendsIn 2008 a total of 8299467 individual fishes (974 identified to species-level) representing1788 species were imported into the US The total number of fishes imported decreasedto 7102246 in 2009 and to 6892960 in 2011 However the number of species imported


A

B

Figure 2 Main dashboard page of wwwaquariumtradedataorg (A) Interactive trade flow map depict-ing exporting countries (blue circles) and ports of entry in the US (green circles) (B) Timeline chart offishes and invertebrates imported into the US based on user-selected dates

increased to 1798 by 2011 While no more than 1800 species were imported in a singleyear 2278 unique species were imported across the three-year span (Table 1)

A similar trend in the overall trade was observed for non-CITES-listed invertebratesduring this time period although the invertebrate data were less voluminous and speciouscompared to the fish data A total of 43 million invertebrates representing 545 species wereimported into the US in 2008 The total number of invertebrates imported decreased toabout 37 million in 2009 and 2011 (Table 2) A total of 724 species were imported overthe three-year span Compared to fishes relatively fewer invertebrates were identified tospecies-level (729)


A

B

C

Figure 3 Drop downmenus for user-generated queries in wwwaquariumtradedataorg (A) Main tabwith Exporting CountriesOcean and Ports of Entry inputs (B) Species tab with taxa selection displayinglsquolsquoTop 20 Speciesrsquorsquo chart (C) lsquolsquoCountries of Originrsquorsquo and lsquolsquoPorts of Entryrsquorsquo charts generated by the query


Figure 4 Exported chart from user-generated query countries of origin for A percula and A ocellarisin wwwaquariumtradedataorg Export header includes query details and export footer includes data at-tributes and date of last database update

Export countriesForty-five countries in total exported marine fishes to the US during the three years(Table 1) with 41 37 and 36 countries noted in 2008 2009 and 2011 respectively ThePhilippines exported 56 of the cumulative total volume (127 million fishes Fig 5)The overall volume of fishes traded decreased by 17 between 2008 and 2011 withcommensurate export decreases from the Philippines and Indonesia Third-ranked SriLanka exported consistently across the three years Exports from fourth-ranked Haitidecreased by nearly 50 between 2008 and 2011 likely due to earthquake activity in 2010

The US imported marine invertebrates from a total of 38 countries during the threeyears (Fig 5 Table 2) although only 27 (2008 2009) or 28 (2011) countries were noted peryear The number of individuals exported per year decreased 14 between 2008 and 2011a rate similar to that of fishes The countries exporting the greatest volume over the threeyears were the Philippines (36 million invertebrates) and Haiti (31 million invertebrates)The number of individual invertebrates exported from the Philippines increased by 24


Table 1 Countries that exported live aquarium fishes to the USData include the number of identifiable species number of species exported in quantities greater than1000 individuals (gt1000 (No)) number of individuals exported across species and the proportion of individuals identifiable to species-level (Known ()) by countryand year

ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)

Australia 115 3 12877 1000 162 2 8773 1000 199 1 9573 996 298 6 31224 999Belize 49 4 9472 999 45 3 8846 994 39 4 14976 996 63 6 33351 996Brazil 71 3 8742 994 61 0 3349 985 45 0 2005 995 87 2 14147 992Canada 2 0 52 1000 2 0 3 1000 1 0 26 423 5 0 81 815Cook Islands 23 2 4763 1000 27 2 3317 1000 ndash ndash ndash ndash 34 2 8080 100Costa Rica 22 2 7903 1000 46 0 3538 1000 28 2 6139 881 53 4 17580 958Curacao 15 0 529 1000 26 0 1383 1000 34 1 3367 997 47 1 5279 998DominicanRepublic

26 3 22121 863 19 4 28944 772 48 8 34272 967 52 8 93872 865

Egypt ndash ndash ndash ndash ndash ndash ndash ndash 20 0 953 921 20 0 953 921Eritrea 52 2 9506 996 44 0 3986 993 ndash ndash ndash ndash 62 2 13519 995Fed States ofMicronesia

ndash ndash ndash ndash ndash ndash ndash ndash 131 0 5550 974 131 0 5550 974

Fiji 187 28 115520 989 228 19 88289 978 311 25 156680 976 363 44 362444 981French Poly-nesia (Tahiti)

106 6 42846 999 101 3 30187 995 73 3 29011 99 144 7 102182 995

Ghana 19 0 509 998 19 0 686 961 22 0 708 955 33 0 1931 968Guatemala 3 0 1055 100 3 0 343 1000 ndash ndash ndash ndash 3 0 1398 1000Haiti 99 23 240552 977 114 23 215909 976 89 19 126799 99 133 30 588516 979Hong Kong 9 0 262 992 4 0 5 100 1 0 16510 03 14 0 16777 19Indonesia 973 214 2402733 972 1009 186 1998195 969 992 181 1867946 973 1284 234 6331781 972Israel ndash ndash ndash ndash 7 0 666 1000 10 2 21985 1000 10 0 22651 1000Japan 44 0 1133 1000 92 0 1137 1000 62 0 569 924 132 0 2839 985Kenya 173 27 144211 977 210 24 139129 978 186 21 101910 99 249 39 388376 981

(continued on next page)

Rhyne

etal(2017)PeerJDOI107717peerj2949

1036

Table 1 (continued)

ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)

Kiribati 67 6 122971 991 52 7 78812 982 72 6 105679 976 103 8 308889 984Malaysia 11 0 622 998 ndash ndash ndash ndash 1 0 13 1000 12 0 635 998Mauritius 63 0 823 934 ndash ndash ndash ndash 41 0 680 982 81 0 1503 956Mexico 90 1 5174 994 40 2 12688 962 62 3 15135 986 118 5 33504 978NetherlandsAntilles

9 0 319 1000 ndash ndash ndash ndash ndash ndash ndash ndash 9 0 319 1000

New Caledo-nia

2 0 84 1000 2 0 75 1000 17 0 387 997 17 0 546 998

Nicaragua 72 2 8986 980 ndash ndash ndash ndash 31 0 1847 932 83 1 10833 972Papua NewGuinea

132 2 6816 998 111 2 8313 986 ndash ndash ndash ndash 176 2 15243 991

Philippines 980 255 4694961 975 1053 248 4024693 973 1016 258 3901058 973 1320 315 12732212 974Rep of Mal-dives

141 5 24574 964 109 4 22093 989 67 11 34360 100 174 19 81275 986

Rep of theMarshall Isl

96 6 37972 940 138 9 115686 751 139 13 142068 787 227 19 334174 788

Saudi Arabia 16 0 326 1000 4 0 19 1000 ndash ndash ndash ndash 20 0 345 1000Singapore 36 1 2606 1000 14 1 2520 998 42 4 13949 995 71 3 19081 996Solomon Is-lands

134 8 47262 965 133 6 34773 949 138 10 41673 925 180 15 125588 947

Sri Lanka 419 30 202632 980 468 34 217116 971 461 28 212407 967 633 57 638606 972Taiwan 33 0 1511 981 29 0 897 853 26 1 2444 1000 63 0 5007 963Thailand 10 3 39887 1000 3 1 8310 1000 ndash ndash ndash ndash 10 0 48197 1000The Bahamas 85 0 951 1000 45 0 432 998 8 0 297 1000 98 0 1681 999Tonga 207 6 27857 895 92 2 8047 923 82 0 2676 91 227 8 39253 902United ArabEmirates


United King-dom



Rhyne


1136

Table 1 (continued)

ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Vanuatu 190 3 19704 972 123 1 12671 968 183 0 14405 941 240 11 47195 962Vietnam 146 1 14593 998 112 1 6545 991 102 0 4826 995 183 6 26022 996Yemen 10 3 10340 100 14 3 11871 1000 ndash ndash ndash ndash 16 3 22211 1000Total 1788 443 8299467 974 1780 411 7102246 967 1798 413 6892960 967 2278 518 22538637 970

Rhyne


1236

Table 2 Countries that exported live aquarium invertebrates to the USData include the number of identifiable species number of species exported in quantitiesgreater than 1000 individuals (gt1000 (No)) number of individuals exported across species and the proportion of individuals identifiable to the species-level (Known()) by country and year

Export Country 2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)

Australia 3 0 231 372 16 0 1881 998 34 0 1020 906 43 1 3132 922Belize 8 2 49515 561 7 3 83922 573 12 6 292176 582 17 7 425613 578Brazil ndash ndash ndash ndash ndash ndash 1 00 ndash ndash ndash ndash ndash ndash 1 00Canada 1 0 2 1000 ndash ndash ndash ndash ndash ndash 28 00 1 0 30 67China 3 0 1260 1000 ndash ndash ndash ndash ndash ndash ndash ndash 3 0 1260 100Costa Rica ndash ndash ndash ndash 3 0 64 1000 ndash ndash ndash ndash 3 0 64 100Curacao ndash ndash ndash ndash 1 0 15 1000 4 0 911 891 5 0 926 893Dominican Re-public

6 2 93781 999 5 2 133056 1000 18 4 107103 963 19 4 333940 988

Federated Statesof Micronesia

ndash ndash ndash ndash ndash ndash ndash ndash ndash ndash 1 00 ndash ndash 1 00

Fiji 6 2 52228 154 9 1 25502 205 23 3 28462 224 29 4 106192 185French Polyne-sia (Tahiti)

ndash ndash 766 00 3 00 48 479 ndash ndash ndash ndash 3 0 814 28

Ghana 3 0 2395 122 3 0 135 1000 3 0 768 535 5 0 3298 254Guatemala ndash ndash 3000 00 ndash ndash ndash ndash ndash ndash ndash ndash ndash ndash 3000 00Haiti 55 24 1409841 925 63 24 1011683 933 47 26 676134 944 79 34 3097658 932Hong Kong 1 1 1520 1000 ndash ndash ndash ndash 1 1 23255 1000 1 1 24775 1000Indonesia 323 57 709736 612 317 51 610264 649 301 48 575657 686 413 90 1895657 646Japan 8 0 425 765 17 0 556 64 12 0 168 911 25 0 1149 726Kenya 18 2 13955 570 17 2 44426 261 22 1 14750 537 30 6 73131 376Kiribati ndash ndash 6 00 ndash ndash 18 00 1 0 80 15 1 0 104 115Mauritius 1 0 198 1000 ndash ndash ndash ndash ndash ndash ndash ndash 1 0 198 1000Mexico 24 1 1429 994 8 2 4035 976 17 2 17678 533 38 5 23142 639New Caledonia ndash ndash ndash ndash ndash ndash ndash ndash ndash 4 00 ndash ndash 4 00Nicaragua 30 8 58918 838 ndash ndash ndash ndash 19 3 31052 745 41 11 89970 806


Rhyne


1336

Table 2 (continued)


Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(N

o)

Kno

wn(

)

Papua NewGuinea

23 0 2323 905 21 2 6336 839 ndash ndash ndash 34 3 8659 856


3 0 95 211 2 0 686 26 ndash 890 00 4 0 1671 23

Rep of the Mar-shall Islands

3 2 47362 1000 7 5 200088 421 24 3 39588 688 29 6 287038 554

Saudi Arabia ndash ndash ndash ndash ndash ndash 5 00 ndash ndash ndash ndash ndash 5 00Singapore 15 0 2654 459 11 0 2063 647 11 1 7017 567 21 2 11734 557Solomon Is-lands

17 2 12521 514 10 1 4084 674 8 2 16753 437 21 3 33358 495

Sri Lanka 63 11 251373 902 60 9 309053 919 54 11 261004 881 87 17 821430 902Thailand 1 0 250 1000 ndash ndash ndash ndash ndash ndash ndash ndash 1 0 250 1000The Bahamas 9 0 92 978 6 0 28 786 ndash ndash ndash ndash 13 0 120 933Tonga 18 3 135089 656 8 2 31214 610 8 1 81918 526 23 3 248221 607United ArabEmirates


United King-dom


Vanuatu 8 0 672 999 4 0 96 979 5 0 132 795 11 0 900 967Vietnam 25 6 293733 81 23 5 108699 272 24 4 106411 122 38 8 508843 131Total 545 137 4256372 734 537 126 3732213 731 535 138 3662980 722 724 220 11651565 729

Rhyne


1436

Figure 5 Trade flow of marine aquarium fishes and invertebrates from source nations into the USduring 2008 2009 and 2011Numbers within circles denote percent contribution to total imports Piechart within US represents Ports of Entry (with the Midwest starting at 0 degrees and clockwise NE SESW and NW)

between 2008 and 2011 This was likely a response to the decrease in volume from Haiti(52 decline from 2008 to 2011) Third-ranked Indonesia (18 million invertebrates)exported a consistent volume across the three years Even though Indonesia ranked thirdin volume it exported the most species (413) during the three years The Philippines (395)and Sri Lanka (87) were second and third respectively in terms of the number of speciesexported to the US

SpeciesMore than half (52) of the total fishes imported into the US (identified to species-levelTable 3) were represented by 20 species There was a great deal of consistency within thesetop 20 species among the years of this study The species ranking was identical between


Table 3 Top 20 live aquarium fish and invertebrate species imported into the US by yearData include proportion of import volume (individuals as Total) andnumber of export countries (Countries (No)) for each

Rank 2008 2009 2011

Species Total Countries (No) Species Total Countries (No) Species Total Countries (No)

Fishes1 Chromis viridis 102 13 Chromis viridis 105 16 Chromis viridis 116 132 Chrysiptera

cyanea56 8 Chrysiptera


parasema47 6

3 Dascyllustrimaculatus

50 11 Dascyllustrimaculatus

44 12 Chrysipteracyanea

44 7

4 Dascyllusaruanus

37 9 Chrysipteraparasema


37 10

5 Chrysipteraparasema

35 3 Dascyllusaruanus


36 8

6 Amphiprionocellaris

32 10 Amphiprionocellaris

30 10 Nemateleotrismagnifica

30 8

7 Nemateleotrismagnifica



30 10

8 Chrysipterahemicyanea

26 2 Chrysipterahemicyanea

24 3 Pterapogonkauderni

19 5

9 Dascyllusmelanurus

18 3 Dascyllusmelanurus

20 6 Centropygeloricula

19 9

10 Pterapogonkauderni

18 3 Paracanthurushepatus

17 14 Pseudocheilinushexataenia

16 9

11 Pseudocheilinushexataenia



16 3

12 Paracanthurushepatus


16 7 Sphaeramianematoptera

15 7

13 Synchiropussplendidus



15 3

14 Centropygeloricula

13 8 Valencienneapuellaris

14 10 Synchiropussplendidus

15 5

15 Labroidesdimidiatus



14 7

16 Salarias fasciatus 13 8 Gramma loreto 13 6 Paracanthurushepatus

13 15

17 Gramma loreto 12 6 Salarias fasciatus 13 10 Salarias fasciatus 12 1118 Valenciennea

puellaris11 9 Sphaeramia

nematoptera13 6 Centropyge

bispinosa12 14


Rhyne


1636

Table 3 (continued)

Rank 2008 2009 2011


19 Centropygebispinosa

11 12 Labroidesdimidiatus


11 11

20 Sphaeramianematoptera

10 7 Centropygebispinosa

11 12 Valencienneastrigata

09 10

Invertebrates1 Paguristes

cadenati220 3 Paguristes


cadenati140 4

2 Lysmataamboinensis

102 6 Lysmataamboinensis


123 8

3 Clibanariustricolor

80 1 Clibanariustricolor

57 2 Mithraculussculptus

73 3

4 Mithraculussculptus


42 2 Trochusmaculatus

46 3

5 Stenopus hispidus 29 11 Lysmata debelius 30 5 Condylactisgigantea

33 4

6 Trochusmaculatus

27 1 Calcinus elegans 28 4 Clibanariustricolor

27 2

7 Nassariusvenustus


28 2 Stenopus hispidus 26 10

8 Tectus fenestratus 25 2 Stenopus hispidus 27 12 Lysmata debelius 25 3

9 Dardanusmegistos

24 5 Tectus fenestratus 25 3 Entacmaeaquadricolor

24 12

10 Percnon gibbesi 23 5 Tectus pyramis 24 4 Dardanusmegistos

24 6

11 Tectus pyramis 21 2 Percnon gibbesi 23 3 Protoreasternodosus

23 5

12 Lysmata ankeri 20 1 Condylactisgigantea

21 5 Nassariusdorsatus

22 1

13 Lysmata debelius 20 5 Sabellastartespectabilis

17 5 Percnon gibbesi 18 5

14 Condylactisgigantea

19 5 Heteractis malu 15 6 Nassariusvenustus

16 1

15 Sabellastartespectabilis

18 4 Lysmata ankeri 15 1 Sabellastartespectabilis

16 4


Rhyne


1736

Table 3 (continued)

Rank 2008 2009 2011


16 Heteractis malu 14 6 Protoreasternodosus

13 4 Nassariusdistortus

16 1

17 Calcinus elegans 13 3 Enginamendicaria

12 2 Heteractis malu 15 4

18 Archaster typicus 13 6 Entacmaeaquadricolor

11 12 Lysmata ankeri 15 1

19 Enginamendicaria


11 1 Pusiostomamendicaria

14 2

20 Stenorhynchusseticornis

11 5 Archaster typicus 11 4 Archaster typicus 14 6

Rhyne


1836

2008 and 2009 and only the 20th ranked fish was different in 2011 (the blueband gobyValenciennea strigata replaced the royal gramma Gramma loreto) The order of the topseven fish species was consistent across the years and represented nearly 33 of total fishimports The green chromis Chromis viridis was the most popular fish species across allthree years (gt10 of total fish imports) and was exported by 13ndash16 different countriesdepending on the year This Chromis species was unique in being collected from a largenumber of countries The only other fish that was equally sourced from a large number ofcountries (an average of 15 per year) was the blue tang Paracanthurus hepatus (Table 3)

Invertebrates demonstrated a similar but more extreme trend The top 20 species ofinvertebrates imported into theUSwere responsible for approximately 75of total imports(identified to species-level Table 3) yet there was more variability in the invertebrate top20 species list compared to the fish list Only the top two species (the scarlet hermitcrab Paguristes cadenati and the scarlet skunk cleaner shrimp Lysmata amboinensis) wereconsistently ranked across the three years Overall 25 invertebrate species were representedon the three yearly top 20 lists (Table 3)

Each country could be represented by a single most exported species Overall the singlemost imported species averaged 37 (fishes) or 63 (invertebrates) of total species volumeexported from that country (Tables 4 and 5) In general countries that exported greaterquantities of marine animals relied less on the contribution of the single most importantspecies to export volume (Fig 6)

Comparison to LEMIS dataThe LEMIS database contains information regarding non-CITES-listed species recordedon declarations under general codes LEMIS data is produced by US-based importersfrom shipment declarations where importers input shipment data into the required 3-177declaration form and present the completed shipment declaration with correspondinginvoice to USFWS prior to shipment clearance We have demonstrated elsewhere (Rhyne etal 2012) that this method of gathering import data is fraught with errors first importerscommonly mislabel shipments as containing marine aquarium species when they onlycontain freshwater fish non-marine species or non-aquarium fish (all increasing the totalnumber of fish reported in the LEMIS database) second the data do not appear to beupdated if shipments are canceled or modified (there is sometimes a significant mismatchbetween the number of individuals on the declaration and the corresponding values onthe invoices) third importers commonly misrepresent the country of origin and source(wild-caughtcaptive-bred) of species in shipments As previously discussed (Rhyne ampTlusty 2012) LEMIS is a tool designed for internal use by USFWS primarily relating tovolume of boxes arriving at ports and CITES compliance Shipments of non-CITES-listedspecies andor unregulated species are not held to any data integrity standards We proposethat the invoice-based method of data collection presented here can rectify many of thedata deficiency issues that currently exist within the marine aquarium trade Through thiswork it was observed that the numbers of fishes imported into the US were routinely60ndash72 fewer than the import volumes reported by the LEMIS database (Fig 7) A largeproportion of the declaration form overestimate was a result of importers misclassifying


Table 4 Top live aquarium fish species exported to the US from each exporting country (1) with proportion of countryrsquos export volume (individuals) ( Total) by year

Export Country 2008 2009 2011

1 Species Total 1 Species Total 1 Species Total

Australia Amphiprion ocellaris 423 Choerodon fasciatus 147 Choerodon fasciatus 169Belize Gramma loreto 224 Gramma loreto 270 Holacanthus ciliaris 258Brazil Holacanthus ciliaris 230 Holacanthus ciliaris 286 Holacanthus ciliaris 445Canada Eumicrotremus orbis 769 Rhinoptera jayakari 667 Eptatretus stoutii 423Cook Islands Pseudanthias ventralis 450 Pseudanthias ventralis 460 ndash ndashCosta Rica Thalassoma lucasanum 311 Thalassoma lucasanum 280 Elacatinus puncticulatus 281Curacao Elacatinus genie 280 Liopropoma carmabi 186 Gramma loreto 312Dominican Republic Gramma loreto 598 Gramma loreto 604 Gramma loreto 360Egypt ndash ndash ndash ndash Zebrasoma xanthurum 317Eritrea Zebrasoma xanthurum 232 Zebrasoma xanthurum 246 ndash ndashFed States of Micronesia ndash ndash ndash Pseudanthias bartlettorum 123Fiji Pseudanthias squamipinnis 96 Pseudanthias squamipinnis 122 Chromis viridis 203French Polynesia (Tahiti) Neocirrhites armatus 439 Neocirrhites armatus 646 Neocirrhites armatus 680Ghana Balistes punctatus 202 Balistes punctatus 363 Holacanthus africanus 414Guatemala Selene brevoortii 588 Selene brevoortii 647 ndash ndashHaiti Gramma loreto 338 Gramma loreto 312 Gramma loreto 329Hong Kong Zebrasoma flavescens 763 Dascyllus trimaculatus 400 Chordata 997Indonesia Chromis viridis 105 Chromis viridis 89 Chromis viridis 108Israel ndash ndash Premnas biaculeatus 347 Amphiprion ocellaris 887Japan Parapriacanthus ransonneti 662 Parapriacanthus ransonneti 135 Paracentropogon rubripinnis 142Kenya Labroides dimidiatus 155 Labroides dimidiatus 142 Labroides dimidiatus 120Kiribati Centropyge loricula 701 Centropyge loricula 632 Centropyge loricula 651Malaysia Amphiprion ocellaris 357 ndash ndash Paracanthurus hepatus 1000Mauritius Amphiprion chrysogaster 120 ndash ndash Macropharyngodon bipartitus 132Mexico Holacanthus passer 495 Holacanthus passer 358 Holacanthus passer 390Netherlands Antilles Elacatinus genie 589 ndash ndash ndash ndashNew Caledonia Chaetodontoplus conspicillatus 845 Chaetodontoplus conspicillatus 853 Cirrhilabrus laboutei 403Nicaragua Apogon retrosella 224 ndash ndash Acanthemblemaria hancocki 395Papua New Guinea Amphiprion percula 297 Paracanthurus hepatus 234 ndash ndashPhilippines Chromis viridis 117 Chromis viridis 133 Chromis viridis 136Rep of Maldives Acanthurus leucosternon 129 Acanthurus leucosternon 127 Acanthurus leucosternon 147Rep of the Marshall Islands Centropyge loricula 537 Centropyge loricula 419 Centropyge loricula 401Saudi Arabia Dascyllus marginatus 307 Anampses caeruleopunctatus 368 ndash ndash


Rhyne


2036

Table 4 (continued)



Singapore Amphiprion ocellaris 423 Amphiprion ocellaris 730 Amphiprion percula 316Solomon Islands Paracanthurus hepatus 194 Paracanthurus hepatus 337 Paracanthurus hepatus 204Sri Lanka Valenciennea puellaris 225 Valenciennea puellaris 211 Valenciennea puellaris 268Taiwan Pomacanthus maculosus 248 Pomacanthus maculosus 194 Amphiprion ocellaris 552Thailand Amphiprion ocellaris 878 Amphiprion ocellaris 905 ndash ndashThe Bahamas Haemulon sciurus 175 Chromis cyanea 115 Haemulon flavolineatum 889Tonga Centropyge bispinosa 148 Centropyge bispinosa 127 Meiacanthus atrodorsalis 96United Arab Emirates ndash ndash ndash ndash Zebrasoma xanthurum 636United Kingdom Amphiprion ocellaris 765 ndash ndash ndash ndashVanuatu Centropyge loricula 94 Chrysiptera rollandi 128 Chromis viridis 69Vietnam Nemateleotris magnifica 83 Nemateleotris magnifica 167 Chaetodontoplus septentrionalis 123Yemen Zebrasoma xanthurum 482 Zebrasoma xanthurum 476 ndash ndashTotal Chromis viridis 100 Chromis viridis 102 Chromis viridis 112

Rhyne


2136

Table 5 Top live aquarium invertebrate species exported to the US from each exporting country (1) with proportion of countryrsquos export volume (individuals) ( Total)by year

Export country 2008 2009 2011


Australia Pseudocolochirus violaceus 756 Actinia tenebrosa 461 Actinia tenebrosa 372Belize Mithraculus sculptus 725 Mithraculus sculptus 519 Mithraculus sculptus 653Canada Enteroctopus dofleini 1000 ndash ndash ndash ndashChina Actinia equina 705 ndash ndash ndash ndashCosta Rica ndash ndash Lysmata wurdemanni 781 ndash ndashCuracao ndash ndash Lysmata wurdemanni 1000 Paguristes cadenati 947Dominican Republic Paguristes cadenati 922 Paguristes cadenati 951 Paguristes cadenati 880Fiji Linckia laevigata 780 Linckia laevigata 870 Linckia laevigata 365French Polynesia (Tahiti) ndash ndash Holothuria (Halodeima) atra 522 ndash ndashGhana Actinia tenebrosa 853 Actinia equina 948 Lysmata grabhami 995Haiti Paguristes cadenati 463 Paguristes cadenati 471 Paguristes cadenati 436Hong Kong Entacmaea quadricolor 1000 ndash ndash Entacmaea quadricolor 1000Indonesia Trochus maculatus 193 Trochus maculatus 191 Trochus maculatus 303Japan Aurelia aurita 529 Aurelia aurita 587 Catostylus mosaicus 294Kenya Lysmata amboinensis 560 Lysmata amboinensis 690 Lysmata amboinensis 489Kiribati ndash ndash ndash ndash Panulirus versicolor 1000Mauritius Heteractis magnifica 1000 ndash ndash ndash ndashMexico Pentaceraster cumingi 749 Turbo fluctuosus 401 Dardanus megistos 597Nicaragua Turbo fluctuosus 492 ndash ndash Coenobita clypeatus 523Papua New Guinea Archaster typicus 457 Archaster typicus 365 ndash ndashPhilippines Lysmata amboinensis 159 Lysmata amboinensis 188 Lysmata amboinensis 163Rep of Maldives Echinaster (Echinaster) sepositus 500 Echinaster (Echinaster) sepositus 833 Pusiostoma mendicaria 459Rep of the Marshall Islands Engina mendicaria 708 Calcinus elegans 452 ndash ndashSingapore Tectus niloticus 411 Entacmaea quadricolor 347 Sabellastarte spectabilis 540Solomon Islands Archaster typicus 464 Archaster typicus 658 Archaster typicus 667Sri Lanka Lysmata amboinensis 619 Lysmata amboinensis 637 Lysmata amboinensis 607Thailand Gecarcoidea natalis 1000 ndash ndash ndash ndashThe Bahamas Ophiocoma alexandri 467 Ancylomenes pedersoni 227 ndash ndashTonga Nassarius venustus 920 Nassarius venustus 776 Nassarius venustus 992United Kingdom ndash ndash ndash ndash Chrysaora quinquecirrha 500Vanuatu Linckia multifora 282 Entacmaea quadricolor 564 Entacmaea quadricolor 543Vietnam Macrodactyla doreensis 341 Macrodactyla doreensis 365 Entacmaea quadricolor 401Total Paguristes cadenati 221 Paguristes cadenati 209 Paguristes cadenati 143

Rhyne


2236

10 100 1000 1000000

01

02

03

04

05

06

07

08

09

10

Cum

ulat

ive

Sum

mat

ion

( in

divi

dual

s)

AustraliaBelizeBrazilCosta RicaCuraccedilaoDominican RepublicEgyptFederated States of MicronesiaFijiFrench Polynesia (Tahiti)GhanaHaitiIndonesiaIsraelJapanKenyaKiribatiMauritiusMexicoNew CaledoniaNicaraguaPhilippinesRepublic of MaldivesRepublic of the Marshall IslandsSingaporeSolomon IslandsSri LankaTaiwanTongaVanuatuVietnam

A

10 100 100000

01

02

03

04

05

06

07

08

09

10

AustraliaBelizeDominican RepublicFijiHaitiIndonesiaJapanKenyaMexicoNicaraguaPhilippinesRepublic of the Marshall IslandsSingaporeSolomon IslandsSri LankaTongaVietnam

B

Number of species exported per country (log10)

Figure 6 The cumulative summation of the number of (A) fishes and (B) invertebrates exported percountry by rank order of species The most-exported species represents a significant proportion of the to-tal individuals exported per country and this importance decreases as a country exports a greater numberof species

shipments as MATF when they only contained freshwater species Occasionally entirefreshwater shipments were erroneously listed as MATF A second unknown portion ofthis error was missing invoices Not all invoices were recovered from the LEMIS systemSeveral hundred records were either missing the invoice or exhibited invoicedeclarationmismatch making the data impossible to verify Similarly invoice-based data reporteda total of 45 countries exporting MATF which was only 60 of the 76 export countriesreported by the LEMIS database (Table 6) These countries represented 5 6 and 11of the total volume of MATF imported into the US according to the LEMIS databaseduring 2008 2009 and 2011 respectively (Table 6) Third is that the declaration is typicallycompleted days before the order is packed which invites variation between estimated andactual order volume Finally there was a lack of adherence to differentiating lsquolsquowild-caughtrsquorsquoand lsquolsquocaptive-bredrsquorsquo animals (Rhyne Tlusty amp Kaufman 2012) The case studies presentedbelow use the invoice-based dataset to shed light on this discrepancy

Estimated fishIn October of 2000 810705 fishes and 124308 invertebrates were imported Duringthe years 2008 2009 and 2011 October represented on average 87 of fish and 86of invertebrate imports into the US Thus it can be estimated that 9327754 fish and1442859 invertebrates were imported into the US during calendar year 2000 Followingthis example 10766706 and 11229443 fish were imported into the US in 2004 and 2005respectively no invertebrate data was available for 2004 and 2005 data


Table 6 Countries reported in LEMIS database that export live aquarium fishes to the USData include number of individuals exported (LEMIS (No)) and propor-tion of LEMIS invoices recovered in the present study (Invoice ()) Shaded countries are those represented in the invoice-based assessment of fish imports to the US

Country 2008 2009 2011 2008ndash2011LEMIS (No) Invoice () LEMIS (No) Invoice () LEMIS (No) Invoice () LEMIS (No) Invoice ()

Antarctica ndash ndash ndash ndash 49 ndash 49 ndashAustralia 16908 762 13973 628 10545 908 41426 754Barbados ndash ndash ndash ndash 82 ndash 82Belgium 266 ndash ndash ndash ndash ndash 266 ndashBelize 19957 475 18214 486 27840 538 66011 505Brazil 61247 143 15342 218 3179 631 79768 177Canada 53 981 119 25 56 464 228 355Cayman Islands 14 ndash ndash ndash ndash ndash 14 ndashChina 354093 ndash 126218 ndash 44151 ndash 524462 ndashChristmas Island ndash ndash ndash ndash 1712 ndash 1712 ndashColombia 24386 ndash ndash ndash 12594 ndash 36980 ndashDem Rep of the Gongo ndash ndash 185 ndash ndash ndash 185 ndashCook Islands 4793 994 3330 996 ndash ndash 8123 995Costa Rica 38904 203 15770 224 6220 987 60894 289Cuba 20 ndash ndash ndash ndash ndash 20 ndashCuracao ndash ndash ndash ndash 2992 1125 2992 1764Czech Republic 1154 ndash 428022 ndash 80500 ndash 509676 ndashDominican Republic 58497 378 64254 45 39687 864 162438 578Ecuador ndash ndash 5990 ndash ndash ndash 5990 ndashEgypt ndash ndash ndash ndash 755 1262 755 1262Eritrea 2796 3400 3046 1309 ndash ndash 5842 2314Fed States of Micronesia ndash ndash ndash ndash 5515 1006 5515 1006Fiji 138801 832 119535 739 171364 914 429700 843French Polynesia (Tahiti) 50261 852 30789 980 30914 938 111964 913Ghana 1119 455 982 699 808 876 2909 664Guatemala 7755 136 343 1000 ndash ndash 8098 173Guinea 357 ndash ndash ndash ndash ndash 357 ndashHaiti 306084 786 280196 771 135890 933 722170 815Hong Kong 205408 01 25806 0 141888 116 373102 45India 5374 ndash 4932 ndash ndash ndash 10306 ndashIndonesia 3044574 789 2743170 728 2228446 838 8016190 790Israel 22 ndash 818 814 25990 846 26830 844Japan 1911 593 1790 635 820 694 4521 628Kenya 175607 821 178715 778 123248 827 477570 813


Rhyne


2436

Table 6 (continued)


Kiribati 143615 856 93586 842 118164 894 355365 869Republic of Korea ndash ndash ndash ndash 6 ndash 6 ndashMalaysia 6516 95 11937 ndash 15255 01 33708 19Mauritania 184 ndash ndash ndash ndash ndash 184 ndashMauritius 6465 127 ndash ndash 681 999 7146 210Mexico 5147 1005 15805 803 22331 678 43283 774Morocco 141 ndash ndash ndash ndash ndash 141 ndashNetherlands ndash ndash 87 ndash 175 ndash 262 ndashNetherlands Antilles 2178 146 1558 ndash 628 ndash 4364 73New Caledonia 317 265 86 872 617 627 1020 535New Zealand ndash ndash ndash ndash 1 ndash 1 ndashNicaragua 15647 574 ndash ndash 2120 871 17767 610Nigeria 4005 ndash 4249 ndash 9446 ndash 17700 ndashNorway 196 ndash 2853 ndash 2903 ndash 5952 ndashPapua New Guinea 11899 573 13167 631 ndash ndash 25066 608Peru 80963 ndash 67609 ndash 10395 ndash 158967 ndashPhilippines 5504928 853 4815066 836 4545933 858 14865927 856Rep of Maldives 27215 903 23572 937 37423 918 88210 921Rep of the Marshall Isl 50143 757 163433 708 152000 935 365576 914Saudi Arabia 7304 45 2131 09 ndash ndash 9435 37Sierra Leone 244 ndash ndash ndash ndash ndash 244 ndashSingapore 310090 08 279794 09 325715 43 915599 21Slovakia 119 ndash ndash ndash ndash ndash 119 ndashSolomon Islands 66057 715 58397 595 42562 979 167016 752Sri Lanka 337521 600 1807583 12 1981399 107 4126503 155Suriname ndash ndash 214 ndash ndash ndash 214 ndashTaiwan 54623 28 47430 19 6950 352 109003 46United Republic of Tanzania 253 ndash ndash ndash ndash ndash 253 ndashThailand 207582 192 115952 72 1117626 ndash 1441160 33The Bahamas 1557 611 1524 283 1397 213 4478 375Tonga 57120 488 13787 584 2474 1082 73381 535Trinidad and Tobago ndash ndash 107805 ndash 46360 ndash 154165 ndashTunisia 558 ndash ndash ndash ndash ndash 558 ndashUkraine 93 ndash ndash ndash ndash ndash 93 ndashUnited Arab Emirates ndash ndash ndash ndash 79 975 79 975United Kingdom 3721 997 583 ndash 4 ndash 4308 861


Rhyne


2536

Table 6 (continued)


United States 828 ndash 87 ndash 45 ndash 960 ndashVanuatu 22850 862 15538 815 15924 905 54312 869Various States 31771 ndash 13369 ndash 23350 ndash 68490 ndashVietnam 26621 548 10832 604 9597 503 47050 553Yemen 20230 511 13114 905 ndash ndash 33344 666Zambia ndash ndash 1286 ndash ndash ndash 1286 ndashTotal (No Invoice Data) 505041 ndash 776984 ndash 1350027 ndash 1499694 ndashTotal (All Countries) 11529062 720 11783973 603 11586805 595 34899840 646

Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF

Figure 7 Comparison of total number of marine aquarium fish imports into the US Comparison oftotal number of marine aquarium fish imports into the US according to the Law Enforcement Manage-ment Information System (LEMIS) and The Marine Aquarium Biodiversity and Trade Flow (MABTF) on-line database across 4 years Data from 2008 2009 and 2011 is from the dataset presented here and datafrom 2005 was presented in Rhyne et al (2012)

Indonesia Sri Lanka Thailand0

300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011

Figure 8 Annual volume of Bangaii cardinalfish (Pterapogon kauderni) into the US by top exportcountries Exports from Sri Lanka (2009 2011) and Thailand (2013) illustrate the likely prevalence ofpreviously unrecognized captive-bred P kauderni in the trade

CONFUSION BETWEEN ldquoWILDrdquo AND ldquoCULTUREDrdquoPRODUCTIONThe Banggai cardinalfish Pterapogon kauderni is a popular marine fish in the aquariumtrade (ranked the 10th 11th and 8th most imported fish into the US during 2008 2009and 2011 Table 5) It was one of the original marine aquarium aquaculture success stories(Talbot et al 2013 Tlusty 2002) which was supposed to reduce the need for wild-caughtfishes While aquaculture should dominate the source of the fish all P kauderni importedduring this three-year span were reported as wild-caught fish Yet import records from SriLanka in 2009 and 2011 and Thailand in 2013 (both outside the natural geographic rangeof P kauderni) suggest this is not the case (Fig 8)


Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012

Figure 9 Temporal variability of the volume of captive-bred Bangaii cardinalfish (Pterapogon kaud-erni) imported into the US Temporal variability of the volume of assumed captive-bred Bangaii cardi-nalfish (Pterapogon kauderni) imported into Los Angeles California US This seasonal variability is con-sistent with the import of wild fish

The export volume of P kauderni from Thailand followed the typical aquarium tradepattern of lower volumes exported in the summer months (JunendashAugust) and in December(Fig 9) Interestingly in 2013 the only year with a 12-month data set starting in Januaryand ending in December the volume of P kauderni (sim120000 individualsyear) wasapproximately 75 of the average total import volume of this species recorded per yearfor 2008 2009 or 2011 Further these fish were listed on import declarations as rangingin size from 1 to 15 inches A 1-inch fish is smaller than the average wild-caught fish (ARhyne pers obs 2013) and instead represents the typical shipping size of a captive-bredfish Shipment manifests also list the number of Dead On Arrival (DOA) from previousshipments which were extremely low (lt05) for wild-caught P kauderni

The shipment manifests have common errors that can be observed on the 3ndash177 USFWSdeclarations On several occasions the importer incorrectly indicated that shipments werewild-caught animals (lsquolsquoWrsquorsquo) After examining the invoices and associated documents (iehealth and aquaculture certificates) we determined that all shipments of P kauderni fromThailand to Quality Marine during the period examined were captive-bred (lsquolsquoCrsquorsquo) and theimporter erroneously selected lsquolsquoWrsquorsquo in the Source box (Box 18B 3ndash177 form) Given thecurrent Endangered Species Act (ESA) listing for P kauderni (NOAA 2014) accurate andtimely trade data are crucial to the sustainable management of this species

MISIDENTIFICATION AND UNKNOWN TRADESimilar to the Banggai cardinalfish clownfishes exported from Southeast Asia arecommonly labeled as wild-caught while many are in fact captive-bred This inaccuracy iscompounded by the misidentification of clownfishes on export invoices especially amongspecies with similar morphological appearances (eg the orange clownfish Amphiprionpercula and the common clownfishA ocellaris) Use of httpwwwaquariumtradedataorgnot only sheds light on source-errors (as seen in the Banggai cardinalfish case study) butalso on potential species misidentifications


A ocellaris A percula0

100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native

Figure 10 Imports of orange clownfish (Amphiprion ocellarisA percula) into the US aggregated over2008 2009 and 2011 Export countries were grouped based on the documented geographic range of eachspecies All non-native individuals are either actually native (but of an unknown distribution) captive-bred or misidentified as to origin on the shipping invoice

Recently the orange clownfish (A percula) was proposed to be listed as threatened orendangered under the ESA mainly due to its small geographic distribution and obligaterelationship with giant sea anemones prone to bleaching events in the Coral TriangleHowever the proposition was also based on the assumption that out of the 400000individuals from the perculaocellaris complex imported into the US in 2005 (Rhyne etal 2012) (a) all specimens were wild-caught and (b) A percula and A ocellaris wereequally traded with 200000 individuals of each species being harvested Utilization ofhttpwwwaquariumtradedataorg can help clarify issues regarding origination of thesespeciesWhile in 2008 2009 and 2011 831398 individual clownfishes of the perculaocellariscomplex were imported into the US (Fig 10) only 163547 individuals were A percula(245 Fig 10) These data suggest that the original assumptions of trade volume used topetition for ESA listing were strongly overestimated

Further the Countries of Origin feature of the database revealed that of the ten exportcountries of A percula seven countries (41 of all individuals) fall outside the naturalgeographic range of this species (Fautin amp Allen 1997 Froese amp Pauly 2015) Furthermorefive of the seven non-native locations are established producers of captive-bred A perculaBased on this 7 of the non-native individuals are likely captive-bred specimens Theremaining two non-native countries (Singapore and the Philippines) account for theresidual 93 of the non-native individuals and are likely misidentifications Interestinglyboth Singapore and the Philippines fall within the natural geographic range of A ocellariswhich is commonly confused withA percula While these individuals may bemisidentifiedit is also important to note that Singapore is a known trans-shipping country and couldhave imported their specimens from another country making the true origin of thesespecimens unattainable Furthermore Singapore is a leader in captive-bred production ofaquarium fish and thus many clownfish could be of captive-bred origin

In summary 41 of A percula imported into the US over the three-year span (a)were misidentified as to species or export country (b) were misidentified as to source(wild-caught versus captive-bred) or (c) represent a recently expanded home range not yetnoted within the scientific literature (unlikely) Regardless of the reason the contribution


of A percula imports to the perculaocellaris complex is not only substantially less thanassumed but also is likely even lower based on the high percent of geographic anomaliesreported here Ultimately this case study confirms the need for more accurate and detailedtrade data such as that provided via httpwwwaquariumtradedataorg for any potentialESA listing activity

The orange clownfish case study demonstrates the effect that species-level nomenclatureconfusion can have on trade data Users of httpwwwaquariumtradedataorg haveilluminated other examples of species misidentification such as Centropyge argi anAtlantic species (Froese amp Pauly 2015) with a significant volume of reported exports fromIndonesia While species-level confusion may be common for specious genera of fishessuch as Centropyge and Amphiprion it is important to recognize instances of more blatantmisidentification For example the iconic yellow tang Zebrasoma flavescens althougha Hawaiian endemic appears in this database as being exported from 13 countries it isdifficult to imagine this fish being confused with anything else Continuous biogeographicalfiltering of invoice data by website users will help to minimize misidentification errorsOne of the egregious cases of identification error is invoice items listed as lsquolsquounknownrsquorsquoThese must find a home in the database and at this point in time are ultimately listedas lsquolsquoChordatarsquorsquo This lack of identification of fishes raises the issue that different countrieshave various reporting requirements that can create data deficiencies within the databaseIf one queries exports from Hong Kong there are no reporting requirements and thusall fish are entered into the database as the generic category lsquolsquoChordatarsquorsquo A deficiency ofreporting requirements for this database is the lack of between-country agreement

DISCUSSIONThe deficiency of comprehensive and overarching data relating to the global marineaquarium trade hinders progress toward its effective management (Foster Wiswedel ampVincent 2016 Fujita et al 2014Rhyne et al 2012)We believe that access tomore accuratedata will allow for increased public engagement in trade sustainability and guide responsibletrade management These data can stimulate action toward consumer education addresschallenges such as misidentification and better manage species Ideally these will resultin greater sustainability (Tlusty et al 2013) Currently there is no overarching systemfor tracking species-level importexport data for the marine aquarium trade This isexacerbated by the lack of standard recordkeeping between different countries (Green2003) Coupled with this is the fact that present data systems are either overly generalbased on declaration forms (LEMIS) or specific to the trade of rare and threatened species(CITES Foster Wiswedel amp Vincent 2016) The Global Marine AquariumDatabase (Green2003) attempted to make sense of some of these discrepancies but can be difficult to usedue to its data structures and relational databases These complications and data limitationspromote misinterpretation as evidenced by the trade volume under- and over-estimatesdiscussed here Changes to and standardization of the way live animal trade data arerecorded are necessary to accurately assess trade pathways and data inaccuracies This willavoid misinterpretations with potentially costly consequences of social economic and


ecological proportions (eg the use of such data to affect ESA listing status) Such costswill only be exacerbated as the aquarium industry continues to grow

Capturing invoice-based data can waylay many of the deficiencies of the extant databasesand should prove useful to both conservation organizations and government agencies byhelping to address the aquarium trade data deficiency that currently exists The benefit ofthe more detailed invoice data we focused on here is that it allowed for a truer estimate ofaquatic wildlife trade The recent ESA petitions for both the Banggai cardinalfish (NOAA2014) and the orange clownfish (Maison amp Graham 2015) were proposed based onincorrect trade data and in each of these cases we demonstrated that increased knowledgeof production areas and modalities do not support the underlining trade assumptions ofthese petitions The assumptions of these ESA listings were demonstrated to be erroneousin part due to reported source (wild-caught captive-bred farmed) inaccuracies of shippedanimals For example exporters will often mark farmed corals as wild corals even whenthey have proper CITES permits for the export of farmed corals (Rhyne Tlusty amp Kaufman2014) Many exporters do not have the appropriate paperwork or government supportneeded to accurately mark corals as captive-bred or farmed on CITES documents oftenbecause of the onerous process required to certify that corals are of farmed origin Whileimproved analysis of invoices will help limit some of this misreporting it will not be totallyunabated until a full fisheryfarm to retail traceability program is initiated

Further issues with the clownfish ESA listing occurred because of demonstratedgeographic misidentification Seven of the ten export countries of the orange clownfishfell outside the natural geographic range of this species Even though these can beindicated as erroneous data correcting these anomalies is outside the purview ofhttpwwwaquariumtradedataorg as it was deemed important to record data as indicatedon the invoice Further discussion of discrepancy can occur through in-depth analysis ofthese data and such efforts are welcomed As this database develops it will be important toidentify the commonlymisidentified species and to help the entire trade improve its speciesdocumentation Ideally lists of lsquolook-alikersquo species can be created and machine learningcan be used to derive biogeographically edited species lists One of the critical assumptionsof this work is that the invoice represents the true contents of the shipment There are anumber of reasons this may not be the case including but not limited to miscountingmisidentification and intentional substitution The question of invoice veracity has beenhighlighted by others (Jones 2008 Wabnitz et al 2003) and without a study directlycomparing invoice to box contents the exact correlation will remain unknown If it isassumed that the trade is based on above-board honest business practices then the invoiceshould be an accurate representation of the trade However the greater the dishonesty inthe trade (eg invoicing a shipment of corals as MATF) the greater the disconnect betweeninvoice and box contents This project was recently named a Grand Prize Winner of theWildlife Crime Tech Challenge (wwwwildlifecrimetechorg) which will spur continueddevelopment of this project with an emphasis on increasing honest business practices bymore easily identifying and enforcing illegal and inaccurate trade activity

The development of httpwwwaquariumtradedataorg is a first step toward improvingthe data which will allow for better management and oversight of the trade in marine


aquatic animals However the invoice analysis was developed from a post-importstandpoint The shipments were accepted at import the paperwork processed and theinvoices stored only to be recovered from storage and delivered for analysis within thisprogram However the OCR data processing has the potential to be utilized in real timeThis would allow for shipment diagnostics to be conducted which could potentiallyidentify misidentified or even illegal shipments Such an import risk-based screen toolexists under the FDArsquos Predictive Risk-based Evaluation for Dynamic Import ComplianceTargeting program (httpwwwfdagovForIndustryImportProgramucm172743htm)and we propose that a similar model would be effective for the aquatic wildlife tradeUltimately such an analysis would provide support to port agents to help them moreeffectively monitor the trade

Aquaculture is a significant means of fish production (Tlusty 2002) This will presenta challenge because the lack of reporting for domestic sales will obscure patterns in thetradeMurray amp Watson (2014) observed clownfish to be the most popular species kept byaquarists although they were not the most popular species imported in our database TheUS domestic production will obscure the relationship between fishes imported and thoseactually being kept by hobbyists However we need to step towards greater recordkeepingon species in the trade and while we focus on international trade it would be ideal ifrecords of domestic production could additionally be kept

While it was not implicitly necessary to estimate the number of individuals importedfor years of incomplete data (2000 2004 and 2005) incomplete data in 2004 and 2005compared to complete data in the later years could give the impression the trade wasincreasing A common query without the estimated number of fish would result in a figurewhere the total number of indivudals in 2000 was 8 while 2004 and 2005 data wouldbe approximately half that of 2008 2009 and 2011 Therefore the estimated fish numberswere calculated to create a more cohesive visual presentation of data and to avoid theincorrect analysis that numbers of US imports of marine aquarium fishes and invertebratesare increasing Prior analysis indicated the trade has decreased from its peak in 2005following the economic recession and a shift to smaller tank sizes (Rhyne amp Tlusty 2012Rhyne Tlusty amp Kaufman 2012) Estimated data should be treated with caution given their(by definition) degree of error We encourage users of this database to determine if moresufficient methods to estimate unknown data can be validated

In summary wildlife data tracking systems require improvement (Chan et al 2015Foster Wiswedel amp Vincent 2016) we are beyond the age of tracking animal shipmentvolume solely for the purpose of assessing port agent staffing needs The systems currentlyin place for tracking non-CITES-listed aquarium animals have proven ineffective inproducing meaningful data that can move the trade toward sustainability and conservation(Vincent et al 2014) The invoice-based dataset presented here while set up as a post-import assessment tool could be easily modified into a real-time aquarium trade datamonitoring system Ideally this can have a positive impact on increasing the veracity of theLEMIS system It behooves the largest aquatic wildlife importer in the world to showcasereal constructive intent to foster sustainability in the trade and to create positive changein an important industry This proof that the trade can be passivily monitored (Wallace


et al 2014) is a huge step toward driving positive change in the trade The next stepwill be to monitor aquarium trade pathways in real-time as this is crucial to effectivelyassist the management of the trade of marine aquarium wildlife for the home aquariumindustry A goal for real-time simultaenous monitoring of exports and imports is nowwithin reach

ACKNOWLEDGEMENTSWe thank the United States Fish and Wildlife Service (USWFS) for providing access toLEMIS data and associated invoices The authors are grateful to the dozens of RogerWilliams University undergraduates that spent thousands of hours cataloguing importdata C Buerner of Quality Marine provided invoices to better understand the trade ofP kauderni A draft of this paper was greatly improved through the efforts of K English SFerse J Murray and an anonymous reviewer

ADDITIONAL INFORMATION AND DECLARATIONS

FundingThe National Oceanic and Atmospheric Administration (NOAA) Coral Reef ConservationProgram and the National Fish and Wildlife Foundation provided funding for this workNOAA provided the data in the form of invoices acquired from USFWS

Grant DisclosuresThe following grant information was disclosed by the authorsNational Oceanic and Atmospheric Administration (NOAA) Coral Reef ConservationProgramNational Fish and Wildlife Foundation

Competing InterestsThe authors declare there are no competing interests

Author Contributionsbull Andrew L Rhyne conceived and designed the experiments performed the experimentsanalyzed the data contributed reagentsmaterialsanalysis tools wrote the paperprepared figures andor tables reviewed drafts of the paper project PI

bull Michael F Tlusty conceived and designed the experiments analyzed the data contributedreagentsmaterialsanalysis tools wrote the paper prepared figures andor tablesreviewed drafts of the paper project Co-PI

bull Joseph T Szczebak and Robert J Holmberg performed the experiments analyzed thedata contributed reagentsmaterialsanalysis tools wrote the paper prepared figuresandor tables reviewed drafts of the paper


Data AvailabilityThe following information was supplied regarding data availability

Marine Aquarium Biodiversity and Trade Flow online database httpswwwaquariumtradedataorg

This site acts as the public repository for the data it is graphically displayed with CSVdownload option

REFERENCESAppeltansW Bouchet P Boxshall G Fauchald K Gordon D Hoeksema B Poore G

Van Soest R Stoumlhr S Walter T Costello M 2011World register of marine speciesAvailable at httpwwwmarinespeciesorg (accessed on 23 June 2011)

Balboa CM 2003 The consumption of marine ornamental fish in the United States adescription from US import data In Cato JC Brown CL edsMarine ornamentalspecies Collection culture and conservation Ames Blackwell Publishing Company65ndash76

Bickford D Phelps J Webb EL Nijman V Sodhi NS 2011 Boosting CITES throughresearchndashresponse Science 331857ndash858 DOI 101126science3316019857-c

Blundell A Mascia M 2005 Discrepancies in reported levels of international wildlifetrade Conservation Biology 192020ndash2025 DOI 101111j1523-1739200500253x

Bruckner AW 2001 Tracking the trade in ornamental coral reef organisms the impor-tance of CITES and its limitations Aquarium Sciences and Conservation 3(1)79ndash94DOI 101023A1011369015080

Chan H-K Zhang H Yang F Fischer G 2015 Improve customs systems to monitorglobal wildlife trade Science 348(6232)291ndash292 DOI 101126scienceaaa3141

Chucholl C 2013 Invaders for sale trade and determinants ofintroduction of ornamen-tal freshwater crayfish Biological Invasions 15(1)125ndash141DOI 101007s10530-012-0273-2

Fautin DG Allen GR 1997 Anemone fishes and their host seaanemones a guide foraquarists and divers Perth Western Australian Museum 160 p

Firchau B Dowd S Tlusty MF 2013 Promoting a socially and environmentallybeneficial aquarium fish trade Connect 201316ndash18

Foster S Wiswedel S Vincent A 2016 Opportunities and challenges for analysis ofwildlife trade using CITES datamdashseahorses as a case study Aquatic ConservationMarine and Freshwater Ecosystems 26154ndash172 DOI 101002aqc2493

Francis-Floyd R Klinger R 2003 Disease diagnosis in ornamental marine fish aretrospective analysis of 129 cases In Cato JC Brown CL edsMarine ornamentalspecies collection culture and conservation Ames Blackwell Publishing Company93ndash100

Froese R Pauly D 2011 FishBase World Wide Web electronic publication Available athttpwwwfishbaseorg



Fujita R Thornhill DJ Karr K Cooper CH Dee LE 2014 Assessing and managingdata-limited ornamental fisheries in coral reefs Fish and Fisheries 15661ndash675DOI 101111faf12040

Garciacutea-Berthou E 2007 The characteristics of invasive fishes what has been learned sofar Journal of Fish Biology 71(Supplement D)33ndash55DOI 101111j1095-8649200701668x

Gopakumar G Ignatius B 2006 A critique towards the development of a marineornamental industry in India Sustain fish In Proceedings of the internationalsymposium on lsquoImproved sustainability of fish production systems and appropriatetechnologies for utilizationrsquo held during 16ndash18 March 2005 Cochi India 606ndash614

Green E 2003 International trade in marine aquarium species using the global marineaquarium database In Cato JC Brown CL edsMarine ornamental species collectionculture amp conservation Ames Blackwell Publishing Company 29ndash48

Holmberg RJ Tlusty MF Futoma E Kaufman L Morris JA Rhyne AL 2015 The800-pound grouper in the room asymptotic body size and invasiveness of marineaquarium fishesMarine Policy 537ndash12 DOI 101016jmarpol201410024

Jones R 2008 CITES corals and customs the international trade in wild coral InLeewis RJ Janse M eds Advances in coral husbandry in public aquariums ArnhemBurgersrsquo Zoo 351ndash361

Lunn K MoreauM 2004 Unmonitored trade in marine ornamental fishes the caseof Indonesiarsquos Banggai cardinalfish (Pterapogon kauderni) Coral Reefs 23344ndash351DOI 101007s00338-004-0393-y

Maison KA GrahamKS 2015 Status review report orange clownfish (Amphiprionpercula) Report to National Marine Fisheries Service Office of Protected ResourcesNOAA Silver Spring

Murray JMWatson GJ 2014 A critical assessment of marine aquarist biodiversity dataand commercial aquaculture identifying gaps in culture initiatives to inform localfisheries managers PLOS ONE 9(9)e105982 DOI 101371journalpone0105982

Murray JMWatson GJ Giangrande A LiccianoM Bentley MG 2012Managing themarine aquarium trade revealing the data gaps using ornamental polychaetes PLOSONE 7(1)e29543 DOI 101371journalpone0029543

NOAA 2014 Endangered and threatened wildlife and plants 12-month finding for theEastern Taiwan Strait Indo-Pacific humpback dolphin dusky sea snake banggaicardinalfish Harrissonrsquos dogfish and three corals under the endangered speciesact Federal Register The Daily Journal of the United States 79(241) 32 p Availableat httpswwwgpogov fdsyspkgFR-2014-12-16pdf2014-29203pdf

Padilla DKWilliams SL 2004 Beyond ballast water aquarium and ornamental tradesas sources of invasive species in aquatic ecosystems Frontiers in Ecology and the Envi-ronment 2(3)131ndash138 DOI 1018901540-9295(2004)002[0131BBWAAO]20CO2

Rhyne AL Tlusty MF 2012 Trends in the marine aquarium trade the influence ofglobal economics and technology AACL Bioflux 599ndash102


Rhyne AL Tlusty MF Kaufman L 2012 Long-term trends of coral imports into theUnited States indicate future opportunities for ecosystem and societal benefitsConservation Letters 5(6)478ndash485 DOI 101111j1755-263X201200265x

Rhyne AL Tlusty MF Kaufman L 2014 Is sustainable exploitation of coral reefspossible A view from the standpoint of the marine aquarium trade Current Opinionin Environmental Sustainability 7101ndash107 DOI 101016jcosust201312001

Rhyne AL Tlusty MF Schofield PJ Kaufman L Morris Jr JA Bruckner AW2012 Revealing the appetite of the marine aquarium fish trade the volume andbiodiversity of fish imported into the United States PLOS ONE 7(5)e35808DOI 101371journalpone0035808

Smith KF Behrens MDMax LM Daszak P 2008 US drowning in unidentifiedfishes scope implications and regulation of live fish import Conservation Letters1103ndash109 DOI 101111j1755-263X200800014x

Smith KF Behrens M Schloegel LM Marano N Burgiel S Daszak P 2009 Reducingthe risks of the wildlife trade Science 324594ndash595 DOI 101126science1174460

Talbot R PedersenMWittenrichML Moe Jr M 2013 Banggai cardinalfish a guide tocaptive care breeding amp natural history Shelburne Reef to Rainforest Media

Tissot BN Best BA Borneman EH Bruckner AW Cooper CH DrsquoAgnes H FitzgeraldTP Leland A Lieberman S Mathews Amos A Sumaila R Telecky TMMcGilvrayF Plankis BJ Rhyne AL Roberts GG Starkhouse B Stevenson TC 2010How USocean policy and market power can reform the coral reef wildlife tradeMarine Policy341385ndash1388 DOI 101016jmarpol201006002

Tlusty M 2002 The benefits and risks of aquacultural production for the aquariumtrade Aquaculture 205(3ndash4)203ndash219 DOI 101016S0044-8486(01)00683-4

Tlusty MF Rhyne AL Kaufman L Hutchins M Reid GM Andrews C Boyle PHemdal J McGilvray F Dowd S 2013 Opportunities for public aquariumsto increase the sustainability of the aquatic animal trade Zoo Biology 321ndash12DOI 101002zoo21019

Vincent AC Sadovy deMitcheson YJ Fowler SL Lieberman S 2014 The role of CITESin the conservation of marine fishes subject to international trade Fish and Fisheries15(4)563ndash592 DOI 101111faf12035

Wabnitz C Taylor M Green E Razak T 2003 From ocean to aquarium CambridgeUNEP-WCMC

Wallace RD Bargeron CT Ziska L Dukes J 2014 Identifying invasive species in realtime early detection and distribution mapping system (EDDMapS) and othermapping tools Invasive Species and Global Climate Change 4219

Wood E 2001 Global advances in conservation and management of marine ornamentalresources Aquarium Sciences and Conservation 365ndash77DOI 101023A1011391700880

WoRMS Editorial Board 2015World Register of Marine Species Available at httpwwwmarinespeciesorg (accessed on 10 June 2015)


management plans for the traded species and ideally could be implemented at key tradeports to better assess the global trade of aquatic wildlife

Subjects Marine Biology Science PolicyKeywords Marine aquarium trade Wildlife trade Coral reef Data visualization

INTRODUCTIONThe live aquarium fish trade faces a multitude of potential threats including incidences ofreduced biodiversity from over-extraction habitat destruction in some source countries(Francis-Floyd amp Klinger 2003 Gopakumar amp Ignatius 2006) and negative impacts ofspecies invasions in the US and elsewhere (Chucholl 2013 Garciacutea-Berthou 2007Holmberget al 2015 Padilla amp Williams 2004) Despite these threats the aquarium trade can be aunique and significant positive force in reef-side communities (Rhyne Tlusty amp Kaufman2014) Benefits include but are not limited to saving threatened species from the brinkof extinction through the development of captive breeding programs (Tlusty 2002) andcatalyzing habitat preservation through sustainable supply-side practices (Firchau Dowd ampTlusty 2013) These sustainable practices include stewardship mechanisms for sustainablelivelihoods via poverty alleviation and the protection of threatened ecosystems (RhyneTlusty amp Kaufman 2014) Finally consumer education of aquarium trade sustainabilitycan promote widespread public appreciation for the worldrsquos aquatic ecosystems with theultimate goal of minimizing negative impacts of this trade (Tlusty et al 2013) While aproactive stance can transform a large consumer base into a powerful agent for biodiversityconservation increased sustainability and human well-being inaction will likely amplifythe deleterious threats currently faced by the trade Currently the lack of oversight leadingto a poor concept of trade volume and subsequent regulatory inefficiency has greatlyhampered the development of a sustainable industry Increasing the sustainability of themarine aquarium animal industry should be considered a primary initiative for the entireaquarium industry transport chain (Tlusty et al 2013) Increasing the sustainability ofthe transport chain of marine aquarium animals is achieved through a more thoroughunderstanding of the magnitude of the trade (Fujita et al 2014) which begins by assessingthe scale of imports into the US (the primary destination) (Rhyne et al 2012) Once theannual volume of US imports is gauged other relevant issues that lead to environmentaland economic benefits can then be tackled including animal quality and shipping survival

There is no clear picture of the number of live marine fish and invertebrate species orindividuals involved in the aquarium trade primarily due to insufficient global trackingof the import and export of these animals (Bruckner 2001 Fujita et al 2014 Green 2003Lunn amp Moreau 2004 Tissot et al 2010 Wabnitz et al 2003) Multiple sources of datahave been used to monitor this trade (Green 2003 Smith et al 2008 Wabnitz et al 2003Wood 2001) However not all of these data systems are sufficient for or were even intendedfor monitoring the aquarium trade For example compulsory data are maintained underfederal mandates for species listed by the Convention on International Trade in Endangered















(nPr)minusn





A

B





A

B

C








ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)


26 3 22121 863 19 4 28944 772 48 8 34272 967 52 8 93872 865




106 6 42846 999 101 3 30187 995 73 3 29011 99 144 7 102182 995



Rhyne


1036

Table 1 (continued)

ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)



New Caledo-nia

2 0 84 1000 2 0 75 1000 17 0 387 997 17 0 546 998




141 5 24574 964 109 4 22093 989 67 11 34360 100 174 19 81275 986


96 6 37972 940 138 9 115686 751 139 13 142068 787 227 19 334174 788


134 8 47262 965 133 6 34773 949 138 10 41673 925 180 15 125588 947



United King-dom



Rhyne


1136

Table 1 (continued)

ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)


Rhyne


1236



Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)


6 2 93781 999 5 2 133056 1000 18 4 107103 963 19 4 333940 988







Rhyne


1336

Table 2 (continued)


Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(N

o)

Kno

wn(

)

Papua NewGuinea



3 0 95 211 2 0 686 26 ndash 890 00 4 0 1671 23


3 2 47362 1000 7 5 200088 421 24 3 39588 688 29 6 287038 554


17 2 12521 514 10 1 4084 674 8 2 16753 437 21 3 33358 495



United King-dom



Rhyne


1436






Rank 2008 2009 2011





parasema47 6




44 7

4 Dascyllusaruanus



37 10




36 8




30 8




30 10




19 5




19 9




16 9




16 3




15 7




15 3




15 5




14 7


13 15




bispinosa12 14


Rhyne


1636

Table 3 (continued)

Rank 2008 2009 2011





11 11




09 10




cadenati140 4




123 8




73 3




46 3


33 4

6 Trochusmaculatus


27 2

7 Nassariusvenustus




9 Dardanusmegistos


24 12


24 6


23 5



22 1





16 1



16 4


Rhyne


1736

Table 3 (continued)

Rank 2008 2009 2011




16 1





19 Enginamendicaria



14 2



Rhyne


1836











Rhyne


2036

Table 4 (continued)




Rhyne


2136





Rhyne


2236

10 100 1000 1000000

01

02

03

04

05

06

07

08

09

10

Cum

ulat

ive

Sum

mat

ion

( in

divi

dual

s)


A

10 100 100000

01

02

03

04

05

06

07

08

09

10


B










Rhyne


2436

Table 6 (continued)




Rhyne


2536

Table 6 (continued)



Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native


























































































(nPr)minusn





A

B





A

B

C








ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)


26 3 22121 863 19 4 28944 772 48 8 34272 967 52 8 93872 865




106 6 42846 999 101 3 30187 995 73 3 29011 99 144 7 102182 995



Rhyne


1036

Table 1 (continued)

ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)



New Caledo-nia

2 0 84 1000 2 0 75 1000 17 0 387 997 17 0 546 998




141 5 24574 964 109 4 22093 989 67 11 34360 100 174 19 81275 986


96 6 37972 940 138 9 115686 751 139 13 142068 787 227 19 334174 788


134 8 47262 965 133 6 34773 949 138 10 41673 925 180 15 125588 947



United King-dom



Rhyne


1136

Table 1 (continued)

ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)


Rhyne


1236



Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)


6 2 93781 999 5 2 133056 1000 18 4 107103 963 19 4 333940 988







Rhyne


1336

Table 2 (continued)


Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(N

o)

Kno

wn(

)

Papua NewGuinea



3 0 95 211 2 0 686 26 ndash 890 00 4 0 1671 23


3 2 47362 1000 7 5 200088 421 24 3 39588 688 29 6 287038 554


17 2 12521 514 10 1 4084 674 8 2 16753 437 21 3 33358 495



United King-dom



Rhyne


1436






Rank 2008 2009 2011





parasema47 6




44 7

4 Dascyllusaruanus



37 10




36 8




30 8




30 10




19 5




19 9




16 9




16 3




15 7




15 3




15 5




14 7


13 15




bispinosa12 14


Rhyne


1636

Table 3 (continued)

Rank 2008 2009 2011





11 11




09 10




cadenati140 4




123 8




73 3




46 3


33 4

6 Trochusmaculatus


27 2

7 Nassariusvenustus




9 Dardanusmegistos


24 12


24 6


23 5



22 1





16 1



16 4


Rhyne


1736

Table 3 (continued)

Rank 2008 2009 2011




16 1





19 Enginamendicaria



14 2



Rhyne


1836











Rhyne


2036

Table 4 (continued)




Rhyne


2136





Rhyne


2236

10 100 1000 1000000

01

02

03

04

05

06

07

08

09

10

Cum

ulat

ive

Sum

mat

ion

( in

divi

dual

s)


A

10 100 100000

01

02

03

04

05

06

07

08

09

10


B










Rhyne


2436

Table 6 (continued)




Rhyne


2536

Table 6 (continued)



Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native













































































A

B





A

B

C








ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)


26 3 22121 863 19 4 28944 772 48 8 34272 967 52 8 93872 865




106 6 42846 999 101 3 30187 995 73 3 29011 99 144 7 102182 995



Rhyne


1036

Table 1 (continued)

ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)



New Caledo-nia

2 0 84 1000 2 0 75 1000 17 0 387 997 17 0 546 998




141 5 24574 964 109 4 22093 989 67 11 34360 100 174 19 81275 986


96 6 37972 940 138 9 115686 751 139 13 142068 787 227 19 334174 788


134 8 47262 965 133 6 34773 949 138 10 41673 925 180 15 125588 947



United King-dom



Rhyne


1136

Table 1 (continued)

ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)


Rhyne


1236



Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)


6 2 93781 999 5 2 133056 1000 18 4 107103 963 19 4 333940 988







Rhyne


1336

Table 2 (continued)


Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(N

o)

Kno

wn(

)

Papua NewGuinea



3 0 95 211 2 0 686 26 ndash 890 00 4 0 1671 23


3 2 47362 1000 7 5 200088 421 24 3 39588 688 29 6 287038 554


17 2 12521 514 10 1 4084 674 8 2 16753 437 21 3 33358 495



United King-dom



Rhyne


1436






Rank 2008 2009 2011





parasema47 6




44 7

4 Dascyllusaruanus



37 10




36 8




30 8




30 10




19 5




19 9




16 9




16 3




15 7




15 3




15 5




14 7


13 15




bispinosa12 14


Rhyne


1636

Table 3 (continued)

Rank 2008 2009 2011





11 11




09 10




cadenati140 4




123 8




73 3




46 3


33 4

6 Trochusmaculatus


27 2

7 Nassariusvenustus




9 Dardanusmegistos


24 12


24 6


23 5



22 1





16 1



16 4


Rhyne


1736

Table 3 (continued)

Rank 2008 2009 2011




16 1





19 Enginamendicaria



14 2



Rhyne


1836











Rhyne


2036

Table 4 (continued)




Rhyne


2136





Rhyne


2236

10 100 1000 1000000

01

02

03

04

05

06

07

08

09

10

Cum

ulat

ive

Sum

mat

ion

( in

divi

dual

s)


A

10 100 100000

01

02

03

04

05

06

07

08

09

10


B










Rhyne


2436

Table 6 (continued)




Rhyne


2536

Table 6 (continued)



Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native













































































A

B

C








ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)


26 3 22121 863 19 4 28944 772 48 8 34272 967 52 8 93872 865




106 6 42846 999 101 3 30187 995 73 3 29011 99 144 7 102182 995



Rhyne


1036

Table 1 (continued)

ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)



New Caledo-nia

2 0 84 1000 2 0 75 1000 17 0 387 997 17 0 546 998




141 5 24574 964 109 4 22093 989 67 11 34360 100 174 19 81275 986


96 6 37972 940 138 9 115686 751 139 13 142068 787 227 19 334174 788


134 8 47262 965 133 6 34773 949 138 10 41673 925 180 15 125588 947



United King-dom



Rhyne


1136

Table 1 (continued)

ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)


Rhyne


1236



Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)


6 2 93781 999 5 2 133056 1000 18 4 107103 963 19 4 333940 988







Rhyne


1336

Table 2 (continued)


Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(N

o)

Kno

wn(

)

Papua NewGuinea



3 0 95 211 2 0 686 26 ndash 890 00 4 0 1671 23


3 2 47362 1000 7 5 200088 421 24 3 39588 688 29 6 287038 554


17 2 12521 514 10 1 4084 674 8 2 16753 437 21 3 33358 495



United King-dom



Rhyne


1436






Rank 2008 2009 2011





parasema47 6




44 7

4 Dascyllusaruanus



37 10




36 8




30 8




30 10




19 5




19 9




16 9




16 3




15 7




15 3




15 5




14 7


13 15




bispinosa12 14


Rhyne


1636

Table 3 (continued)

Rank 2008 2009 2011





11 11




09 10




cadenati140 4




123 8




73 3




46 3


33 4

6 Trochusmaculatus


27 2

7 Nassariusvenustus




9 Dardanusmegistos


24 12


24 6


23 5



22 1





16 1



16 4


Rhyne


1736

Table 3 (continued)

Rank 2008 2009 2011




16 1





19 Enginamendicaria



14 2



Rhyne


1836











Rhyne


2036

Table 4 (continued)




Rhyne


2136





Rhyne


2236

10 100 1000 1000000

01

02

03

04

05

06

07

08

09

10

Cum

ulat

ive

Sum

mat

ion

( in

divi

dual

s)


A

10 100 100000

01

02

03

04

05

06

07

08

09

10


B










Rhyne


2436

Table 6 (continued)




Rhyne


2536

Table 6 (continued)



Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native


















































































ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)


26 3 22121 863 19 4 28944 772 48 8 34272 967 52 8 93872 865




106 6 42846 999 101 3 30187 995 73 3 29011 99 144 7 102182 995



Rhyne


1036

Table 1 (continued)

ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)



New Caledo-nia

2 0 84 1000 2 0 75 1000 17 0 387 997 17 0 546 998




141 5 24574 964 109 4 22093 989 67 11 34360 100 174 19 81275 986


96 6 37972 940 138 9 115686 751 139 13 142068 787 227 19 334174 788


134 8 47262 965 133 6 34773 949 138 10 41673 925 180 15 125588 947



United King-dom



Rhyne


1136

Table 1 (continued)

ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)


Rhyne


1236



Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)


6 2 93781 999 5 2 133056 1000 18 4 107103 963 19 4 333940 988







Rhyne


1336

Table 2 (continued)


Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(N

o)

Kno

wn(

)

Papua NewGuinea



3 0 95 211 2 0 686 26 ndash 890 00 4 0 1671 23


3 2 47362 1000 7 5 200088 421 24 3 39588 688 29 6 287038 554


17 2 12521 514 10 1 4084 674 8 2 16753 437 21 3 33358 495



United King-dom



Rhyne


1436






Rank 2008 2009 2011





parasema47 6




44 7

4 Dascyllusaruanus



37 10




36 8




30 8




30 10




19 5




19 9




16 9




16 3




15 7




15 3




15 5




14 7


13 15




bispinosa12 14


Rhyne


1636

Table 3 (continued)

Rank 2008 2009 2011





11 11




09 10




cadenati140 4




123 8




73 3




46 3


33 4

6 Trochusmaculatus


27 2

7 Nassariusvenustus




9 Dardanusmegistos


24 12


24 6


23 5



22 1





16 1



16 4


Rhyne


1736

Table 3 (continued)

Rank 2008 2009 2011




16 1





19 Enginamendicaria



14 2



Rhyne


1836











Rhyne


2036

Table 4 (continued)




Rhyne


2136





Rhyne


2236

10 100 1000 1000000

01

02

03

04

05

06

07

08

09

10

Cum

ulat

ive

Sum

mat

ion

( in

divi

dual

s)


A

10 100 100000

01

02

03

04

05

06

07

08

09

10


B










Rhyne


2436

Table 6 (continued)




Rhyne


2536

Table 6 (continued)



Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native













































































Table 1 (continued)

ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)



New Caledo-nia

2 0 84 1000 2 0 75 1000 17 0 387 997 17 0 546 998




141 5 24574 964 109 4 22093 989 67 11 34360 100 174 19 81275 986


96 6 37972 940 138 9 115686 751 139 13 142068 787 227 19 334174 788


134 8 47262 965 133 6 34773 949 138 10 41673 925 180 15 125588 947



United King-dom



Rhyne


1136

Table 1 (continued)

ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)


Rhyne


1236



Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)


6 2 93781 999 5 2 133056 1000 18 4 107103 963 19 4 333940 988







Rhyne


1336

Table 2 (continued)


Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(N

o)

Kno

wn(

)

Papua NewGuinea



3 0 95 211 2 0 686 26 ndash 890 00 4 0 1671 23


3 2 47362 1000 7 5 200088 421 24 3 39588 688 29 6 287038 554


17 2 12521 514 10 1 4084 674 8 2 16753 437 21 3 33358 495



United King-dom



Rhyne


1436






Rank 2008 2009 2011





parasema47 6




44 7

4 Dascyllusaruanus



37 10




36 8




30 8




30 10




19 5




19 9




16 9




16 3




15 7




15 3




15 5




14 7


13 15




bispinosa12 14


Rhyne


1636

Table 3 (continued)

Rank 2008 2009 2011





11 11




09 10




cadenati140 4




123 8




73 3




46 3


33 4

6 Trochusmaculatus


27 2

7 Nassariusvenustus




9 Dardanusmegistos


24 12


24 6


23 5



22 1





16 1



16 4


Rhyne


1736

Table 3 (continued)

Rank 2008 2009 2011




16 1





19 Enginamendicaria



14 2



Rhyne


1836











Rhyne


2036

Table 4 (continued)




Rhyne


2136





Rhyne


2236

10 100 1000 1000000

01

02

03

04

05

06

07

08

09

10

Cum

ulat

ive

Sum

mat

ion

( in

divi

dual

s)


A

10 100 100000

01

02

03

04

05

06

07

08

09

10


B










Rhyne


2436

Table 6 (continued)




Rhyne


2536

Table 6 (continued)



Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native













































































Table 1 (continued)

ExportCountry

2008 2009 2011 2008ndash2011

Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)


Rhyne


1236



Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)


6 2 93781 999 5 2 133056 1000 18 4 107103 963 19 4 333940 988







Rhyne


1336

Table 2 (continued)


Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(N

o)

Kno

wn(

)

Papua NewGuinea



3 0 95 211 2 0 686 26 ndash 890 00 4 0 1671 23


3 2 47362 1000 7 5 200088 421 24 3 39588 688 29 6 287038 554


17 2 12521 514 10 1 4084 674 8 2 16753 437 21 3 33358 495



United King-dom



Rhyne


1436






Rank 2008 2009 2011





parasema47 6




44 7

4 Dascyllusaruanus



37 10




36 8




30 8




30 10




19 5




19 9




16 9




16 3




15 7




15 3




15 5




14 7


13 15




bispinosa12 14


Rhyne


1636

Table 3 (continued)

Rank 2008 2009 2011





11 11




09 10




cadenati140 4




123 8




73 3




46 3


33 4

6 Trochusmaculatus


27 2

7 Nassariusvenustus




9 Dardanusmegistos


24 12


24 6


23 5



22 1





16 1



16 4


Rhyne


1736

Table 3 (continued)

Rank 2008 2009 2011




16 1





19 Enginamendicaria



14 2



Rhyne


1836











Rhyne


2036

Table 4 (continued)




Rhyne


2136





Rhyne


2236

10 100 1000 1000000

01

02

03

04

05

06

07

08

09

10

Cum

ulat

ive

Sum

mat

ion

( in

divi

dual

s)


A

10 100 100000

01

02

03

04

05

06

07

08

09

10


B










Rhyne


2436

Table 6 (continued)




Rhyne


2536

Table 6 (continued)



Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native















































































Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(N

o)

gt1000(N

o)

Total(N

o)

Kno

wn(

)


6 2 93781 999 5 2 133056 1000 18 4 107103 963 19 4 333940 988







Rhyne


1336

Table 2 (continued)


Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(N

o)

Kno

wn(

)

Papua NewGuinea



3 0 95 211 2 0 686 26 ndash 890 00 4 0 1671 23


3 2 47362 1000 7 5 200088 421 24 3 39588 688 29 6 287038 554


17 2 12521 514 10 1 4084 674 8 2 16753 437 21 3 33358 495



United King-dom



Rhyne


1436






Rank 2008 2009 2011





parasema47 6




44 7

4 Dascyllusaruanus



37 10




36 8




30 8




30 10




19 5




19 9




16 9




16 3




15 7




15 3




15 5




14 7


13 15




bispinosa12 14


Rhyne


1636

Table 3 (continued)

Rank 2008 2009 2011





11 11




09 10




cadenati140 4




123 8




73 3




46 3


33 4

6 Trochusmaculatus


27 2

7 Nassariusvenustus




9 Dardanusmegistos


24 12


24 6


23 5



22 1





16 1



16 4


Rhyne


1736

Table 3 (continued)

Rank 2008 2009 2011




16 1





19 Enginamendicaria



14 2



Rhyne


1836











Rhyne


2036

Table 4 (continued)




Rhyne


2136





Rhyne


2236

10 100 1000 1000000

01

02

03

04

05

06

07

08

09

10

Cum

ulat

ive

Sum

mat

ion

( in

divi

dual

s)


A

10 100 100000

01

02

03

04

05

06

07

08

09

10


B










Rhyne


2436

Table 6 (continued)




Rhyne


2536

Table 6 (continued)



Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native













































































Table 2 (continued)


Species

Individu

als

Species

Individu

als

Species

Individu

als

Species

Individu

als

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(No)

Kno

wn(

)

Total(No)

gt1000(N

o)

Total(N

o)

Kno

wn(

)

Papua NewGuinea



3 0 95 211 2 0 686 26 ndash 890 00 4 0 1671 23


3 2 47362 1000 7 5 200088 421 24 3 39588 688 29 6 287038 554


17 2 12521 514 10 1 4084 674 8 2 16753 437 21 3 33358 495



United King-dom



Rhyne


1436






Rank 2008 2009 2011





parasema47 6




44 7

4 Dascyllusaruanus



37 10




36 8




30 8




30 10




19 5




19 9




16 9




16 3




15 7




15 3




15 5




14 7


13 15




bispinosa12 14


Rhyne


1636

Table 3 (continued)

Rank 2008 2009 2011





11 11




09 10




cadenati140 4




123 8




73 3




46 3


33 4

6 Trochusmaculatus


27 2

7 Nassariusvenustus




9 Dardanusmegistos


24 12


24 6


23 5



22 1





16 1



16 4


Rhyne


1736

Table 3 (continued)

Rank 2008 2009 2011




16 1





19 Enginamendicaria



14 2



Rhyne


1836











Rhyne


2036

Table 4 (continued)




Rhyne


2136





Rhyne


2236

10 100 1000 1000000

01

02

03

04

05

06

07

08

09

10

Cum

ulat

ive

Sum

mat

ion

( in

divi

dual

s)


A

10 100 100000

01

02

03

04

05

06

07

08

09

10


B










Rhyne


2436

Table 6 (continued)




Rhyne


2536

Table 6 (continued)



Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native


















































































Rank 2008 2009 2011





parasema47 6




44 7

4 Dascyllusaruanus



37 10




36 8




30 8




30 10




19 5




19 9




16 9




16 3




15 7




15 3




15 5




14 7


13 15




bispinosa12 14


Rhyne


1636

Table 3 (continued)

Rank 2008 2009 2011





11 11




09 10




cadenati140 4




123 8




73 3




46 3


33 4

6 Trochusmaculatus


27 2

7 Nassariusvenustus




9 Dardanusmegistos


24 12


24 6


23 5



22 1





16 1



16 4


Rhyne


1736

Table 3 (continued)

Rank 2008 2009 2011




16 1





19 Enginamendicaria



14 2



Rhyne


1836











Rhyne


2036

Table 4 (continued)




Rhyne


2136





Rhyne


2236

10 100 1000 1000000

01

02

03

04

05

06

07

08

09

10

Cum

ulat

ive

Sum

mat

ion

( in

divi

dual

s)


A

10 100 100000

01

02

03

04

05

06

07

08

09

10


B










Rhyne


2436

Table 6 (continued)




Rhyne


2536

Table 6 (continued)



Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native













































































Table 3 (continued)

Rank 2008 2009 2011





11 11




09 10




cadenati140 4




123 8




73 3




46 3


33 4

6 Trochusmaculatus


27 2

7 Nassariusvenustus




9 Dardanusmegistos


24 12


24 6


23 5



22 1





16 1



16 4


Rhyne


1736

Table 3 (continued)

Rank 2008 2009 2011




16 1





19 Enginamendicaria



14 2



Rhyne


1836











Rhyne


2036

Table 4 (continued)




Rhyne


2136





Rhyne


2236

10 100 1000 1000000

01

02

03

04

05

06

07

08

09

10

Cum

ulat

ive

Sum

mat

ion

( in

divi

dual

s)


A

10 100 100000

01

02

03

04

05

06

07

08

09

10


B










Rhyne


2436

Table 6 (continued)




Rhyne


2536

Table 6 (continued)



Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native













































































Table 3 (continued)

Rank 2008 2009 2011




16 1





19 Enginamendicaria



14 2



Rhyne


1836











Rhyne


2036

Table 4 (continued)




Rhyne


2136





Rhyne


2236

10 100 1000 1000000

01

02

03

04

05

06

07

08

09

10

Cum

ulat

ive

Sum

mat

ion

( in

divi

dual

s)


A

10 100 100000

01

02

03

04

05

06

07

08

09

10


B










Rhyne


2436

Table 6 (continued)




Rhyne


2536

Table 6 (continued)



Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native























































































Rhyne


2036

Table 4 (continued)




Rhyne


2136





Rhyne


2236

10 100 1000 1000000

01

02

03

04

05

06

07

08

09

10

Cum

ulat

ive

Sum

mat

ion

( in

divi

dual

s)


A

10 100 100000

01

02

03

04

05

06

07

08

09

10


B










Rhyne


2436

Table 6 (continued)




Rhyne


2536

Table 6 (continued)



Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native













































































Table 4 (continued)




Rhyne


2136





Rhyne


2236

10 100 1000 1000000

01

02

03

04

05

06

07

08

09

10

Cum

ulat

ive

Sum

mat

ion

( in

divi

dual

s)


A

10 100 100000

01

02

03

04

05

06

07

08

09

10


B










Rhyne


2436

Table 6 (continued)




Rhyne


2536

Table 6 (continued)



Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native

















































































Rhyne


2236

10 100 1000 1000000

01

02

03

04

05

06

07

08

09

10

Cum

ulat

ive

Sum

mat

ion

( in

divi

dual

s)


A

10 100 100000

01

02

03

04

05

06

07

08

09

10


B










Rhyne


2436

Table 6 (continued)




Rhyne


2536

Table 6 (continued)



Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native













































































10 100 1000 1000000

01

02

03

04

05

06

07

08

09

10

Cum

ulat

ive

Sum

mat

ion

( in

divi

dual

s)


A

10 100 100000

01

02

03

04

05

06

07

08

09

10


B










Rhyne


2436

Table 6 (continued)




Rhyne


2536

Table 6 (continued)



Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native

















































































Rhyne


2436

Table 6 (continued)




Rhyne


2536

Table 6 (continued)



Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native













































































Table 6 (continued)




Rhyne


2536

Table 6 (continued)



Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native













































































Table 6 (continued)



Rhyne


2636

2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native













































































2005

2008

2009

2011

0

5

10

15

Milli

on F

ish

Year

-1

LEMISMABTF



300100000

120000

140000

160000

Fish

Yea

r-1

Wild Captive-bred

2013

2005200820092011




Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native













































































Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

02000400060008000

100001200014000160001800020000

Fish

mon

th-1

20132014

2012







100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native














































































100000

200000

300000

400000

500000

600000

700000

Tot

al F

ish

Impo

rted

NativeNon-Native













































































expanding our understanding of the trade in marine aquarium … · 2017-01-26 · submitted 12 june...

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