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The freshwater aquarium trade as a vector for incidental invertebrate fauna PDF

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BiolInvasions(2010)12:3757–3770 DOI10.1007/s10530-010-9768-x ORIGINAL PAPER The freshwater aquarium trade as a vector for incidental invertebrate fauna Ian C. Duggan Received:1February2010/Accepted:20April2010/Publishedonline:30April2010 (cid:2)SpringerScience+BusinessMediaB.V.2010 Abstract The aquarium trade has a long history of non-indigenous species had a greater probability of transportingandintroducingfish,plantsandsnailsinto beingrecordedinaquariacontainingaquaticplantsand regionswheretheyarenotnative.However,otherthan in those that were heated. Methods for disposal of snails, research on species carried ‘‘incidentally’’ aquariumwastesrangedfromdepositingwashingson rather than deliberately by this industry is lacking. I thelawnorgarden(alowriskforinvasion)todisposing sampledinvertebratesintheplankton,andfromwater of water into outdoor ponds or storm-water drains among bottom stones, of 55 aquaria from 43 New (ahigherrisk).Itisrecommendedthataquariumowners Zealandhouseholds.Irecorded55incidentalinverte- beencouragedtopouraquariumwastesontogardensor bratetaxa,includingcopepods,ostracods,cladocerans, lawns—already a common method of disposal—as molluscs, mites, flatworms and nematodes. Six were invasionriskwillbeminimisedusingthismethod. known established non-indigenous species, and eight others were not previously recorded from New Keywords Pet trade (cid:2) Craspedacusta (cid:2) Zealand.Ofthelatter,twoharpacticoidcopepodspe- Melanoides (cid:2) Exotic (cid:2) Invertebrates (cid:2) cies, Nitokra pietschmanni and Elaphoidella sewelli, Alternative vector are not native to or known from New Zealand, demonstrating the aquarium trade continues to pose aninvasionriskforincidentalfauna.Theremainingsix specieswerelittoral/benthicrotiferswithsubtropical/ tropicalaffinities;thesemayormaynotbenative,as Introduction research on this group is limited. A variety of behaviours associated with the set-up and keeping of The keeping of home aquaria is a popular hobby homeaquariawererecorded(e.g.,fishandplantsinany globally with, for example, 8% of households in the homeweresourcedfromstores,wildcaught,orboth, UnitedStatesestimatedtopossessfishtanks(AMVA and cleaning methods varied), which made predic- 2007). The aquarium trade has been responsible for tion of ‘‘high risk’’ behaviours difficult. However, the establishment of many fish, plant and mollusc species into areas where they are not native (e.g., Fuller et al. 1999; Mackie 1999; Rixon et al. 2005). I.C.Duggan(&) Despitethis,littleresearchhasbeenconductedonthe CentreforBiodiversityandEcologyResearch, role of the aquarium trade in transporting and DepartmentofBiologicalSciences,TheUniversityof introducing species into new habitats. That which Waikato,PrivateBag3105,Hamilton,NewZealand e-mail:[email protected] hasbeenconductedhasdealtalmostexclusivelywith 123 3758 I.C.Duggan taxa intentionally (i.e., deliberately) carried by the vector has been found carrying these species (Grig- aquarium trade (e.g., fish, aquatic plants and large orovichetal.2003). snail species; Rixon et al. 2005; Cohen et al. 2007; Theaimsofthisresearchareprimarilytoexamine Changetal.2009).Lessisknown,however,offauna the identity and abundance of small incidental inver- carried ‘‘incidentally’’, such as rotifers, micro-crus- tebratesinfreshwaterhomeaquaria,andtodetermine taceansandsnails,likelytobelivingintheaquarium whetherthisvectorprovidesaninvasionriskforthese water column, associated with the aquatic plants, animals. New Zealand is an ideal place to undertake and/or living amongst the bottom stones, sediments thisresearch,as:(1)anumberofnon-indigenoussnails and detritus. Nevertheless, the presence of some are known to have been introduced via this vector members of these taxa has been noted in home there,while(2)anumberofinvertebrateinvadersexist aquaria. For example, snail species not intentionally for which no vector has been definitively identified carried by the aquarium trade were noted in North (particularly crustaceans; e.g., Duggan et al. 2006b). American stores by Rixon et al. (2005) and Keller Further, I aim to elucidate behaviours or factors that and Lodge (2007), and members of this taxon are contribute tothe presence ofnon-indigenous inverte- well known to aquarium owners for ‘‘appearing’’ in bratesinhomeaquaria,whichmayprovidestrategies their tanks (Perera and Walls 1996). This is partic- forthemanagementoftheindustryorprovideadvice ularly the case for the Physidae, Thiaridae, Planor- for best practices for individual aquarium owners. bidae and Lymnaeidae, families with representatives Questionstoownersthereforeaimedto:(1)identifythe that have established non-indigenous populations in sourcesofbiotadeliberatelykeptintheaquaria,which many parts of the world (Perera and Walls 1996; mayinfluencethepresenceofincidentalfauna(e.g.,if Pointier 1999; Cowie and Robinson 2003). Despite addition of aquatic plants or live fish food affects their small size, aquatic invertebrate invaders can incidental faunal composition), (2) determine other have large ecological or economic effects on the factors that may affect the composition of incidental ecosystems they invade. For example, zebra mussels faunawithintanks(e.g.,presenceofaquariumheating, (Dreissena spp.) and the ctenophore Mnemiopsis filtration, and frequency of tank cleaning), and (3) leidyi were listed amongst the IUCN’s ‘‘100 of the provide information on potential mechanisms that world’s worst invasive species’’ (Lowe et al. 2000). mightleadtothereleaseofnon-indigenousfaunainto Theaccumulationofknowledgeofbiotacarriedby naturalwaters. any particular vector allows for the development of specificmanagementstrategiestoreduceintroduction rates.Forexample,studiesofbiotainballastwaterand Methods sediments(e.g.,Lockeetal.1993;Dugganetal.2005) hasledtothedevelopmentofballastwaterexchange, Sampling was undertaken over a one year period or partial ballast water exchange, to reduce invasion between 5 March 2007 and 8 March 2008 for risk. To date there have been no similar systematic incidental fauna in: (a) the open water of the tanks studiesthatquantifythenumbersofincidentalspecies and (b) the interstitial water among the bottom carriedbytheaquariumtradeand,besidesmolluscan sediments. Volunteer participants were recruited by taxa,littleefforthasbeenmadetodeterminewhether approaching aquarium owners known to the author, non-indigenousspeciesareevenbeingcarriedbythis using a bulk email message to all University of vector.Thus,itisnotyetknownwhethermanagement Waikato staff, through two newspaper articles techniques, outside of existing border control strate- describing the project that asked for participants, gies, are required to reduce invasion rates for this and by word of mouth. Most participants (37: 86%) vector.Thisisdespitesmallinvertebratesestablishing were in the Waikato region, while the remainder populations in regions for which no vectors are yet (6: 14%) was recruited in the Wellington region, known. In the North American Great Lakes, for approximately 500 km away. Participants were vis- example—arguablythebeststudiedfreshwatersystem ited at times convenient to them, and where possible globallyforinvadersandtheirvectors—theharpacti- atatimewhentheyweretocleantheirhomeaquaria, coid copepods Heteropsyllus cf. nunni and Nitokra so as to provide estimates of the faunal composition incerta have established populations but no specific and abundances that might be released from aquaria. 123 Thefreshwateraquariumtrade 3759 Samples were collected from the interstitial water owner; live food bought from a pet store), how often among bottom sediments using a modified 50 mL thetankwascleaned,howoldthetankwas,andhow volumetricpipettewith rubber bulb. The tapered end thewaterwasdisposedofaftercleaning.Itwasnoted of the pipette was removed to increase the bore size whether tanks were lit, had heating, and/or had to 5 mm. The end of the pipette was placed into the filtration.Arulerwasusedtoestimatethevolumesof aquarium sediment (usually comprised of aquarium water in the tank and area comprised of inorganic stones[5 mm), and water was sucked from among sediments (i.e., aquarium stones, rocks), as well as the stones in *50 mL aliquots while moving the the sizes of the sediments. pipette horizontally through the stones. Water from Samples were returned to the laboratory for enu- the pipette was emptied into a measuring cylinder merationandidentificationusingdissectingandcom- until a total of approximately 250 mL of water was pound microscopes. Particular attention was paid to collected for each tank. Two tanks contained no snailsandsmallcrustaceans,taxathathaveextensive bottom stones; in these the sampling was undertaken invasionhistoriesinNewZealandandelsewhere(e.g., similarly, running the base of the pipette across the Winterbourn 1973; Grigorovich et al. 2003; Duggan bottom of the glass. Water was typically sampled et al. 2006a). The volume of inorganic sediment, depending on the method used by the aquarium comprising of mainly uneaten food and faeces, was owner to clean out their tank, so as to replicate taxa estimatedfromeachsamplebymeasuringtheareaof likely to be washed from the tanks while cleaning; detritus in the sample containers of the preserved this involved either pouring off water after removal zooplanktonsamples. ofthefish(smalltanks),siphoning,orusingajug(or Because of the wide variety of behaviours and similar container) to remove water. Water samples environmentalvariablesassociatedwithaquaria,even were collected to a measured volume of approxi- withinindividualhouseholds(Table 1),trendsintank mately5 L,althoughsmallervolumeswerecollected fauna were analysed on a tank by tank basis, rather in the smallest tanks (i.e., the smallest tank sampled thanbyhousehold.Relationshipsbetween(a)species hadatotalvolumeofonly2.4 Labovethesediment). abundancesinthewaterandsedimentcomponentsof Samples were concentrated using a 40 lm sieve and tanks and (b) tank environmental factors recorded, the material retained was preserved in[50% ethanol were examined using Pearsons correlations (continu- (final concentration). A visual inspection was con- ousdata;e.g.,temperature)orMann–WhitneyUtests ductedforsnailsonplants,glassaquariumsides,and (categorical data; e.g., lighting). Mann–Whitney U among the sediments, and representative individuals tests were only carried out on categorical data where werecollected;thiswasbecauseonlysmall(\5 mm) samplessizesweresimilarbetweengroups.Datawere snails were collected using the pipette method. A transformed for statistical tests (usually log (x ? 1)) small volume of surface water was also collected, to improve normality. The frequencies of occurrence from which specific conductance, temperature (YSI ofcommonnon-indigenousspecies(i.e.,thosespecies 30 conductivity and temperature meter) and pH withfiveormoreoccurrences)wereexaminedrelative (Oakton Ultra Basic pH Testr) were immediately tofactorsthatmayhaveleadtotheirintroductioninto, measured. and their survival in, tanks. Questionnaires were used to elucidate how inci- dental species might be entering the tanks, and to understandhowtheymightbeintroducedfromhome Results aquaria into natural waterways. Each participant was questioned to determine the numbers and types of Tank attributes fish,plantsandsnailsdeliberatelyaddedtoeachtank over time since set up, those extant in the aquaria at Fifty-five aquaria belonging to forty-three people the time of sampling, and the sources of the biota were sampled. Thirty-four households had single intentionally placed in the tank (i.e., bought or wild tanks, while four had two tanks each; all of these collected). Participants were also asked what sorts of tanks were sampled. Where greater than two tanks food had been added to the aquarium (store bought were present, only a subset or representative tanks dry, dead or frozen food; live food caught by the were sampled; one household had three tanks (one 123 3760 I.C.Duggan Table1 Tanksetup, Pertankbasis(/55) Perhouseholdbasis(/43) environmentalvariables, andownerbehavioursfor Fish 31tropicalfishtanks 22tropicalfish sampledhomeaquariaona 20goldfishtanks 17goldfish pertankandperhousehold basis 4mixed(i.e.,tropicalfish 4mixed(i.e.,3householdshadtropicalfishand andgoldfishinthesame goldfishinthesametank,1indifferenttanks) tank) Plants 44with 32with 11without 10without ?1withtankswithandwithout Tankvolume Ave:56.2L Ave:116.6L (excl. Min:2.4L Min:2.4L sediments) Max:303.4L Max:1157.7L Volumeof Ave:5.22L Ave:11.9L stones/gravel Min:0L Min:0L Max:31.92L Max:169.4L Heating 30heated 22heated 25notheated 19notheated ?2withtankswithandwithout Filter 39external/internalfilter 27external/internalfilter 4undergravelfilter 3undergravelfilter 12nofiltration 11nofiltration ?2withtankswithandwithout Lighting 31lit 22lit 14notlit 20notlit ?1withtankswith/without sampled), one-four tanks (two samples), one-five proportionstockedapplesnails(Pomaceabridgesior tanks (three sampled), one-nine tanks (four sampled) P.canaliculata;9.1%).Allownersfedtheirfishwith and the other fourteen tanks (three sampled). Thus, processedfoodboughtfrompetstores,althoughsome the average number of tanks present in each house- supplemented this with live bought (e.g., Daphnia; hold was 1.8, although the typical household 18.2%) or live caught food (e.g., mosquito, chiron- (median) had only one. Each aquarium was sampled omidorotherinsectlarvae;38.2%),whichcouldalso once only during the study. allow introduction of incidental fauna to aquaria. There was a great variety of behaviours involved Variousfactorsweremeasuredthatmightaffectthe with the keeping of aquaria that might affect the survival and abundance of animals in tanks. The species composition of incidental fauna, or the temperature of heated tanks ranged from 18.7 to presence of non-indigenous species, in the tanks 28.8(cid:3)C(average25.4(cid:3)C),whileunheatedtanksranged (Table 1). Most owners (67.4%) sourced their aquar- from10.2to25.1(cid:3)C(average18.3(cid:3)C);thelattervaried ium inhabitants from stores only, with only a small seasonally.Specificconductancerangedfrom99.9to proportion having everything in their tanks wild 4170.0 lS cm-1 (average 621.3 lS cm-1), and pH caught(4.7%;oneownerhadcollectedallnativefish rangedfrom4.4to9.5(average6.7).Mostaquariahad and plants, while the other had collected all non- filtrationandlighting,andmoretankswereheatedthan indigenousfish,plants,andsnailsfortheiraquarium). not(Table 1). Theremainderhadamixtureofstoreboughtandwild caught inhabitants (usually bought fish with wild- Species composition and abundance in aquaria collectedplants).Alltankscontainedfish,whilemost (80.0%) had also added plants that may aid in the Fifty-five incidental invertebrate species were introductionofincidentalfaunatotanks.Onlyasmall recorded (Table 2). Species belonged to a variety of 123 Thefreshwateraquariumtrade 3761 Table2 Listofspecies Water Sediments Notknown Established identifiedfromaquariumwater fromNZ NIS andwaterfrominterstitial sedimentsfromNewZealand Rotifers aquaria Cephalodellaforficula X X Colurellauncinata X Cupelopagisvorax X Dicranophorusepicharis X X Keratellavalga X Lecaneaculeata X X Lecanebulla X X Lecaneclosterocerca X X Lecanedecipiens X X Lecanehamata X X Lecaneinermis X X X Lecaneludwigiiludwigii X X Lecaneludwigiiohioensis X X Lecanelunaris X X Lecanenana X X X Lecanepyriformis X X Lecanerhytida X Lecanesignifera X X X Lepadellaacuminata X X Lepadellaovalis X Lepadellapatella X Limnaismelicerta X X Proalessp. X X Ptygurasp. X X cf.Sphyriaslofuana X X Squatinellagellii X Squatinellamutica X X Testudinellapatina X X Bdelloids X X Copepods Acanthocyclopsrobustus X X Eucyclopsserrulatus X X Microcyclopsvaricans X Paracyclopssp. X Nitokrapietschmanni X X X Phylognathopusvigueri X X Elaphoidellasewelligp X X Epactophanesrichardi X Cyclopoidnauplii X X Harpacticoidindet X X Harpacticoidnauplii X X Cladocerans Alonacambouei X X 123 3762 I.C.Duggan Table2 continued Water Sediments Notknown Established fromNZ NIS Chydorussphaericusgp X Molluscs Ferrisiaspp. X Gyrauluscorinna X Lymnaeastagnalis X X Melanoidestuberculata X X Physaacuta X X Planorbariuscorneus X X Radixauricularia X X Othertaxa Ostracods X X Mites X X Oligochaetes X X Flatworms X Springtails X Chironomid:Paratanytarsusgrimmii X Indeterminatechironomids X Nematodes X X Craspedacustasowerbii X X taxonomic groups, and included rotifers, copepods, were typically similar (usually \10% difference), cladocerans, molluscs, ostracods, mites, flatworms, exceptformolluscs,whichwerenotfoundamongthe chironomids, nematodes and cnidaria. These were water fraction (but in 43% of tanks overall), mites, primarily mobile benthic species (e.g., substrate- alsomorefrequentlyfoundamongsediments(78.2%) associatedrotiferspecies,cyclopoidandharpacticoid than water (30.9%), and rotifers, more frequently copepods,snails,oligochaetes),withfewtrulyplank- occurring in water (94.5%) than sediments (76.4%). tonic representatives (e.g., Keratella valga, recorded Averageabundancesoftaxaweregreatlyaffectedby as a single individual in one tank; Table 2). Average having high abundances in few or single tanks; for densities of animals in the water component were example, cladocerans were on average the most 499.9 ind. L-1 and in sediments 966.1 ind. L-1, abundant taxon in the water fraction across tanks although these numbers were influenced by some (average 374.0 ind. L-1), but were present only in 5 tanks containing many individuals (i.e., the data was of 55 tanks; one tank had over 25,000 ind. L-1. non-normally distributed). Thus, the median abun- Rotifers were next most abundant, with average dance of animals in the water fraction was 13.3 ind. numbersof79.2ind.L-1,butthemedianacrosstanks L-1, and in sediment 241.8 ind. L-1, indicating that was only 5.5 ind. L-1. Rotifers dominated the the sediments typically have far greater abundances sediments, averaging 402.9 ind. L-1 (median 47.2), thantheopenwater.Onaverage,theareacomprising followed by cladocerans, mites and copepods. the bottom sediments was 10.6% of that in the water Relationshipsbetweentotalnumbersoforganisms fraction. Rotifers were the most frequently occurring present in the water and sediment fractions and tank taxon overall (present in 96.4% of the 55 tanks variables were weak or non-existent. A significant sampled), followed by mites (85.5%), oligochaetes negative relationship was determined only for the (56.4%), copepods (49.1%) and molluscs (43%); number of animals in the water column and the these were the only taxonomic groups present in volume of organic sediments present (i.e., the more greaterthan20%oftanks.Percentageoccurrencesof faecesandfoodremaininginthebottomofthetanks, each taxon between the water and sediment fractions the less animals were found in the water column); 123 Thefreshwateraquariumtrade 3763 however, this had a low correlation coefficient Examining the data for the 36 aquaria where all (R = -0.329; P = 0.014). No relationships were inhabitants were derived from aquarium stores (no found for temperature, specific conductance, pH, wild caught components), all of the common non- time since last clean, age of tank, tank volume, indigenous species were found, indicating that stores sediment size (inorganic), number of fish, or the have acted as a route for these species to enter presence or lighting (all P[0.05). aquaria. Too few data were found for aquaria containing only wild caught specimens; however, Species notpreviously known from New Zealand noneofthethreetankscontainedanyofthecommon non-indigenousspecies,althoughonetankhadPhysa Eight species not previously recorded from New acuta, now a very common species throughout New Zealand, and six known established non-indigenous Zealand. Lymnaea stagnalis was also only found species,wererecorded.Fivesnailspecieswereknown once, in one of the wild caught tanks, indicating this invaders in New Zealand: Planorbarius corneus species may not be spread via stores. The other (11tanks), Melanoides tuberculata(10tanks), Physa variable that seemingly related to the presence of acuta (6 tanks), Radix auricularia (3 tanks), and common non-indigenous species was heating; all Lymnaea stagnalis (1 tank). Additionally, a small groupswererecordedmorefrequentlyinheatedtanks cnidarian medusa, likely Craspedacusta sowerbii, than non-heated (Fig. 1). No discernable patterns was also recorded on one occasion. Species recorded wereobservedforfiltration,additionoflivefood,and thataredefinitelynotnativetoNew Zealand,butare lighting (graphs not presented). not yet recorded as having non-indigenous popula- tions, were the harpacticoid copepods Nitokra pie- Release tschmanni(9tanks)andElaphoidellasewelli(1tank). Additionally, rotifer species not recorded from New A variety of methods were used for disposal. Of the Zealand, Lecane aculeata (1 tank), Lecane inermis 55 tanks examined, 19 disposed of their washings (3 tanks), Lecane nana (1 tank), Lecane signifera exclusively on the garden (16 of 43 owners), 27 put (3tanks),Limnaismelicerta(3tanks),andaspecimen theirwashingsexclusivelydownthesink(19owners) conformingmostcloselytoSphyriaslofuana(1tank), and 5 tanks had their washings either poured on the were also recorded. garden or poured down the sink (4 owners). Two As all tanks contained fish, the frequency of the tanks had never had water removed, one because it more common ([5 occurences) non-indigenous taxa wasanewtank(1 monthold),whiletheotherowner wasrelatedtootherfactorsthatmayhaveledtotheir did not feel the need to remove water, even after introduction and survival to tanks; all snails were having the tank for 2 years. Of the remaining tanks, combined and treated as one group, but Melanoides one was disposed of in an external fish pond, while tuberculata, Planorbarius corneus and Physa acuta another was emptied into a storm-water drain acces- werealsocommonenoughtobetreatedindividually. sible outside the window from the aquarium. Tank All six rotifer species were treated as a single group, cleaning behaviours varied greatly among partici- as none were found in greater than five tanks pants, ranging from tanks never having water individually. The introduction of aquatic plants removed to regular complete cleanings (including showed clear patterns of introduction (Fig. 1). Snails rock scrubbing). On average, 48.3% of any tanks were only found in aquaria with aquatic plants, contents were expected to be emptied during each whereas they were absent from all aquaria without cleaning, and tanks were cleaned on average every aquaticplants.Therotifersnotpreviouslyrecordedin 2.7 months. With an average tank volume of 56.2 L NewZealandshowedasimilarpattern.However,the (Table 1), it is estimated that 118.7 L per tank per harpacticoid Nitokra pietschmanni was found in year(or159.7 Lperhouseholdperyear)aredisposed relativelyequalproportionsintankswithandwithout of. Assuming approximately 8% of households pos- introduced macrophytes; as adults and nauplii of this sesshomeaquaria(AMVA2007),andthatthereare1.6 taxon were commonly found in the water column, million households in New Zealand (December 2009 they are perhaps introduced to tanks with the water estimate from Statistics New Zealand: www.stats. surrounding fish. govt.nz),itisestimatedthat12.8 9 104householdsin 123 3764 I.C.Duggan Fig.1 Percentages of tanks in which common (C5 occur- Zealand home aquaria, for a aquariums with and without rences) non-indigenous species, or species not known from aquariumplants,binaquariawhereallconstituentswerestore New Zealand, were recorded, from a survey of 55 New bought,andcinheatedandnon-heatedaquaria 123 Thefreshwateraquariumtrade 3765 New Zealand own home aquaria. Thus, 20.4 million remainingtankshadadditionsofspeciesfromnatural litres of aquarium waste might be expected to be waters,butestablishedpopulationsofN.pietschman- disposed ofperyearinNew Zealand,fromacountry ni are not known from New Zealand, despite a of only 4.4 million people. At an average concen- number of surveys for this group (Chapman and tration of animals of 539.6 ind. L-1 in each tank, Lewis 1976). The second harpacticoid, Elaphoidella 11.09 109 animals might be expected to be washed sewelli, is known from Asia, Africa, and India, and fromtanksperyear.Acrossalltankstheharpacticoid was found in only one aquarium. All inhabitants in copepod N. pietschmanni, the most common and that aquarium were also store bought. The common abundantofnon-indigenousspeciespresent,averaged Asian native distribution of N. pietschmanni and E. 56.9 ind. L-1, and therefore 7.7 9 108 individuals sewelli is consistent with the predominant global might beexpectedtobewashed fromtanksper year. regions for aquarium fish culture facilities and Assuming2of43ownersreleasewaterintopondsor exports (Cheong 1996), which indicates the likely storm-water drains, more ‘‘high risk’’ methods of pathway for introduction. disposal, it is estimated that 5.2 9 108 animals in Some or all of the rotifer species not previously total, and 3.6 9 107 N. pietschmanni, might be recorded may potentially be native to New Zealand, released into external waterways without treatment. as species found in this study typically have broad This is likely an overestimate for freshwaters as, distributions,whilelittleresearchhasbeenconducted although in the Waikato most storm-water is drained inNewZealandofthelittoralorbenthicfauna(Shiel to lakes and rivers, a high proportion of storm-water et al. 2009). However, of the four Lecane species in the country will be drained to the ocean. recorded, L. nana, L. signifera and L. aculeata are noted by Segers (1995) to be more common in ‘‘tropicalandsubtropicalwaters’’,or‘‘warmwaters’’. This suggests that these species may well not be Discussion nativetoNewZealand,unlesstheyaretobefoundin the warmer far north of the country. Taxonomic composition and non-indigenous A number of known established non-indigenous species invertebrate species were identified, which shows a strong signal for the historical importance of this I recorded a variety of invertebrate species from the vector. Those recorded were primarily snail species: water column and sediments of home aquaria, Radixauricularia,Planorbariuscorneus,Melanoides including species possibly present in New Zealand tuberculata,PhysaacutaandLymnaeastagnalis.Most but not previously recorded, non-indigenous species havebeenestablishedinNewZealandforsometime, known to have already invaded, and non-indigenous withtheaquariumtradealreadyinferredasthevector species not yet known to have established. As such, for introduction (Winterbourn 1973; Duggan 2002). the aquarium trade appears to remain a significant However, some of these taxa still have fairly limited risk for the establishment and spread of non-indige- distributions in New Zealand, particularly M. tuber- nous species. culata, R. auricularia and P. corneus, and the aquar- Of greatest concern, two harpacticoid copepods iumtradethereforestillprovidesopportunitiesforthe and six rotifer species were identified that are not spreadofthesespecies.Asmallcnidarianmedusawas known from New Zealand. The harpacticoid Nitokra also recorded;nofreshwater cnidarian native toNew pietschmanniwasthemostcommon,recordedinnine Zealand, and few globally, are capable of producing aquaria belonging to seven owners, and was also medusa, but the non-indigenous Craspedacusta sow- highly abundant. This suggests that N. pietschmanni erbiiisknownfromanumberoflakesinthenorthern is widespread and abundant in the New Zealand halfoftheNorthIsland(e.g.,Boothroydetal.2002). aquarium trade. To date, this species is known from Sampling in the current study was not designed to Asia (Japan, China, South Korea) and the Hawaiian collectbenthicattachedspecies;assuch,thepolypsof Islands (Chang and Yoon 2008). In four tanks, all thisspeciescouldbewidespreadintheaquariumtrade, fish, plants and snails were store bought, indicating withthesamplingofthismedusaafortuitousevent.For that stores are the probable source to the tanks; the example, medusa of C. sowerbii have been noted in 123 3766 I.C.Duggan aquaria previously, both in New Zealand and else- study,theremustbesomelevelofregionalism(14of where(Fish1971;RahatandCampbell1974). 43ownershadeithersomeorallcomponentsoftheir I did not attempt identifications of many oligo- tankscollectedfromthewild).However,withrespect chaetes during this study, due to most specimens to non-indigenous species in aquaria, there appear to consisting of juveniles, but these were common and be both similarities and differences between taxa occasionally abundant. However, many tanks were recordedinNewZealandandthoseknowntooccurin observed to contain Pristina species of which, for the aquarium trade elsewhere. Snail species such as example, three species are believed to have invaded Physa acuta and Melanoides tuberculata are known the North American Great Lakes (Grigorovich et al. within, and are considered to have established popu- 2003). Combined, this data shows that the aquarium lations from, home aquaria in a number of locations trade has likely been responsible for a large number globally (e.g., New Zealand and North America; of invasions into New Zealand’s freshwaters to date PereraandWalls1996;Mackie1999;Duggan2002). and—althoughmostnon-indigenousspeciesrecorded However,otherspecies,suchasthetropicalAmerican had already invaded—still poses somefuture risk for species Elminia floridensis, have been observed on both establishment and spread. plants in aquarium stores in North America (Rixon The composition of the aquarium fauna was etal.2005),butwerenotdocumentedinmystudy.The distinctive not only by what was present, but also incidentalfaunainhomeaquariamaydifferdepending bywhatwasabsent.WhilealargeproportionofNew ontheregionsfromwhichplantsandfisharesourced Zealand’s historic aquatic invertebrate invaders were from;however,asSingaporeistheprimarysourceof recorded duringthisstudy(snails,C. sowerbii),most aquarium fishes to global markets, including North of the recently established invaders are relatively AmericaandEurope(Cheong1996),itisprobablethat large planktonic species, including Daphnia and some incidental species are widespread. Surveys of calanoid copepod species (e.g., Duggan et al. faunasinhomeaquariaelsewhere,andinthedominant 2006a). These taxa are not likely to survive in home aquariumfishandplantculturefacilitiesglobally,are aquaria containing fishes, as they are large bodied, imperativefordeterminingthesepatterns. and would be easily seen by these visual predators Overall,mostofthetaxaweretypicaloflittoralor (e.g., Ortega-Salas et al. 2009). Indeed, 5 of the 43 benthic habitats, and not of the plankton. This may households sampled had bought live Daphnia cul- reflectthehighdegreeofzooplanktivoryinthewater tures from stores to feed their fish, but no live fraction of tanks, but also that these species possess Daphnia were recorded in tanks. One of the recent adaptations for attaching to substrates that might calanoidcopepodinvadersrecordedinNewZealand, allow them to fasten to the aquarium sides during the North American Skistodiaptomus pallidus, has cleaning (e.g., Duggan 2001), or to the manner in beenrecordedaslivingwithinsuchcultures(Duggan which they are introduced into the tanks (below). et al. 2006a; Banks and Duggan 2009). If the aquarium trade has been the source of these intro- Arrival, survival and abundance in tanks ductions, they have perhaps been released directly into waterways from culture samples, rather than It was difficult to determine any single factor, or being released from home aquaria themselves. Alter- subset of factors, responsible for high abundances of natively,diapausingeggs,whichbothofthesespecies taxa in tanks; this makes the development of man- are known to produce (e.g., Dowell 1997), could be agement strategies for owners to reduce the preva- deposited in tank bottom sediments and hatch lence of species in their tanks difficult. I observed a following release into natural waters. vastarrayofpractices thatownersused tosetupand Fewdataexistoftheincidentalfaunacarriedbythe maintain their aquaria, which led to a multitude of aquarium trade. Although it is probable that benthic possible important factors responsible for survival in rather than planktonic forms will dominate, it is not tanks,evenamongaquariaownedbyasingleperson. knownwhetherthesamespeciesexistelsewhere(i.e., Even categorical factors measured had complicating whether there is a cosmopolitan aquarium fauna). factors. For example, filtration devices used differed, Certainly, due to the widespread collection of plants which could be set at a variety of different strengths, and fish from local lakes and ponds in the current and their efficiency will have been affected by the 123

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The aquarium trade has a long history of transporting and introducing fish, plants and . It was noted whether tanks were lit, had heating, and/or had filtration. TFH Publications, Inc. Neptune City, New Jersey, USAGoogle Scholar.
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