The LEMIS data from 2000-01-01 to 2022-06-30 included 8.7 million entries (listed import groupings), of which we examined 3,479,466 entries, where the items were imported (93% of the 8.7 million entries), constituted whole individuals (52%) live or dead (e.g., excluding parts), and was reported as a count (i.e., quantity of individuals, not mass, length or volume). 99% of these entries were cleared for import, and 84% of 8.7 million entries were listed to a taxonomic level of genus or species. Once outliers were removed, these data record the import of 2,847,052,429 items that constituted a whole individual, 815,572,384 (29%) of which were vertebrates. Records included 21,135 species from 10,452 genera (Data_S22), of which 6,689 species were invertebrates (4,496 genera), 11,243 were terrestrial vertebrates (3,968 genera), 2,885 were marine/aquatic vertebrates (1,664 genera), and 280 were plants. Small numbers of other taxa (such as fungi) were also noted (38; Figure 1A; Data_S9). Entries that lacked genus/species level data, (and were often listed under “Exemption 4” -commercial interests barring release of species level data) were not included in our analysis (Figure 1A).
We merged LEMIS data with data for CITES imports into the US over the same period and the number of species increased to 29,445 with the greatest increase being 8,116 plant species (in addition to 232 animal species) that were only recorded in CITES, not present in the LEMIS data. This difference is attributed to the jurisdiction of plant imports in the US by the United States Department of Agriculture's Animal and Plant Health Inspection Service (USDA-APHIS). Likewise, animals inspected by the USDA (cattle, sheep, swine, goats, horses, mules, domesticated chickens, turkeys, ducks, geese, and squabs) are rarely included in LEMIS even though some may come from the wild, and siluriformes (catfish) and ratites may be declared to USFWS or USDA, and their records may not be reliable (see supplements).
Invertebrate groups showed the highest numbers of whole individuals traded (both live and dead). Miscellaneous (i.e., those organisms not confidently included in more specific ad-hoc groupings) included 580,769,084 individuals and represented the most traded of our groupings (60% of total individuals; Figure 2; Data_S4), followed by Crustaceans and Molluscs at 119,859,391 (12%) and Fish 84,544,098 (8.7%).
By number of entries, Terrestrial Mammals dominated, with more than 1,348,125 entries (35%), followed by Echinoderms and Cnidaria 886,973 (23%; Figure 1B-C). The differences between the numbers of individuals traded and the number of entries highlights differences in trade dynamics and/or recording (Text S1), where for some groups individuals are more likely to be imported in bulk shipments (potentially enabled by their size or form, e.g., eggs/slings).
1.1 Dimensions of trade: what is traded, what they are traded for, and where are they from?
Most species in trade, even if predominantly captive bred, will have some individuals from the wild (Table S1), though the veracity of listings of source (wild or captive) may vary and is hard to assess 11 (Data_S21).
Overall 65% of individuals were declared as wild-sourced, but this ranged dramatically depending on the group (Figure 2). For example, Marine Mammals show 96% and Echinoderms and Cnidaria show 94% of individuals coming from the wild. Lepidoptera and Arachnids are reported to fall below 15% wild-sourced individuals, thus skewing overall percentages 3. Terrestrial Mammals (21%), Birds (18%), and Arachnids (14%) have higher rates of ranched individuals. Ranching refers to individuals collected from the wild as eggs or juveniles, then raised in captivity; however, the listing of some species (particularly certain invertebrates, as well as Terrestrial Mammals) as ‘ranched’ suggests inconsistent use, as the collection of some of these species as juveniles or eggs from the wild seems improbable, especially in invertebrates. However, it should be noted that if weight was considered the percentage wild may increase for certain taxa such as fish, but inconsistent units as well as body-sizes means this measure is challenging.
Mammals exhibit the most wild-sourced entries (94%), followed by Birds (89%) (Figure 2). This result potentially indicates that bulk imports are less likely to be wild-sourced, while those smaller or more specialist shipments are wild-sourced. The wild sourcing of individuals is not evenly spread, but impacts the vast majority of traded species to some extent (Figure S6). All marine mammal species listed as traded have wild-sourced individuals, and every other group except Lepidoptera and Arachnids have over 80% of the species being wild-sourced at some point (Data_S22). The lower overall percentage from the wild is influenced by a small number of species being traded in high volumes not originating from the wild, whilst conversely the majority of other taxa are imported in smaller numbers, but largely from the wild (Figure S6). In vertebrates in particular, we find species traded in high quantities with most being wild-sourced. Similar patterns exist when examined by order, where many orders see high percentages of individuals wild-sourced (Figure S7). When looking by order, the skew is towards wild sourcing: over 64% of orders have over 90% of their individuals originating from the wild (Figure S7).
1.2 Changing patterns of trade
For most vertebrate taxa the number of species and genera traded annually increased until 2017 after which they declined and species counts fluctuated. Birds (at 1,876 species) and reptiles (at 1,249 species) reached peaks earlier, in 2015 (Figure 3; Data_S10; Data_S7). Invertebrate groups show similar patterns, with a continued increase throughout the LEMIS data time frame; Crustaceans and Molluscs reached over 973 species in 2021, the most species traded in a single year for invertebrates (Figure 3, S1). Increases in invertebrates during 2020 may relate to heightened trade during the pandemic (Marshall et al., 2022). Declines in imports in recent years for some taxa may relate to more restrictive import regulations 24, and increased domestic trade from captive breeding. LEMIS recording methods also changed in 2016, which affected how data was split (records were split by purpose), which may be responsible for some of the rapid changes in trade. Notice of species that may be listed in upcoming CITES meetings could also increase trade in such groups prior to such meetings (such as 2019).
Cumulative sums of species over time reveal an increase in the number of species traded, however, part of the increase may originate from splitting of species, or creating notation to enable species to be more accurately listed (such as noting species individually rather than within aggregates such as “tropical fish”). Patterns largely follow the counts of species per group, with birds and fish showing the steepest increases (Figure S1; Data_S5). By contrast marine mammals have remained stable over the past two decades. The higher species numbers of birds and fish appear largely due to high numbers of species traded in individual years (Figure S1; Data_S8), while other groups (e.g., Terrestrial Mammals, Echinoderms and Cnidaria) reveal a considerable number of species consistently traded over the entire period.
For some groups the percentage of wild caught individuals decreased between 2020 and 2022 (Figure S2; Data_S13), but this time period may include issues with harvesting and transport access, or a lack of monitoring resources during the COVID-19 pandemic. Patterns of ranching also varied over time, with increasing percentages of species listed as ranched for mammals, arachnids and amphibians, especially in recent years. Other groups fluctuate, possibly marking changes in regulations, demand, and purpose, or possible delays in clearing in 2018 for vertebrate taxa.
There are also differences between the number of individuals imported as whole vs. live. While all live individuals were obviously whole, whole specimens may also be imported dead for other purposes (compare numbers in Figures S3 and S4; Data_S11). This highlights differences between likely trophies or specimens and animals imported live as pets or for research, education, and live exhibition. This difference is particularly noticeable in mammals, birds, and lepidopterans, who see large trade in dead whole individuals. For mammals and birds, this disparity is likely due to the import of hunting trophies (see section 1.3). Live bird imports have decreased, whereas imports of live lepidopterans have increased. Conversely, Reptiles, Amphibians, Crustaceans and Molluscs, see little difference in the live versus whole individual trends, indicating the predominance of live trade in these groups.
1.3 Imported for what purpose?
Commercial trade was the dominant purpose stated for the trade of whole individuals (Figure S5, Data_S16). All groups show over 77% of individuals are traded for commercial purposes, with the exception of marine mammals that are primarily imported for “scientific/research” purposes. The focus on commercial purposes is particularly acute in 8 of 15 groups (Figure 4A), with over 98% of individuals imported for commercial purposes.
Much of the wild sourcing of individuals is listed as being for commercial purposes (Figure S5). For example, 98% of species listed as wild-sourced within Echinoderms and Cnidaria are identified as being for commercial purposes, similarly over 90% of listed wild-sourced Porifera are labelled for commercial purposes; however, terrestrial invertebrates (at least those listed) are largely captive bred (Figure 4). Reptiles show the greatest intersection for a vertebrate group, where 54% of wild-sourced individuals were for commercial purposes, likely for the pet trade 2. However, the reptile percentage (both live and whole) has likely declined over time due to an increase in the percentage of captive bred individuals imported, as well as a slight decrease in overall trade volumes (Figure S2, 3; 4).
Much of the non-commercial trade is for scientific/research purposes, with a few notable exceptions (Figure 4): non-commercial trade of birds and mammals appears connected to hunting trophies. The primary non-commercial reason for amphibian import is listed as education, and this appears linked to the live import of these individuals. Amphibians have the highest percentage of wild sourcing when looking at non-commercial live imports for vertebrate groups (Figure S5; Data_S19), but the percentage of live reptiles and marine mammals is greater than amphibians when commercial trade is included. Live imports largely follow the trends of whole individuals when commercial trade is included. Notable exceptions are Terrestrial Mammals (25% wild dead or alive; 2% live only), Porifera Sponges, Bryozoa, and Squirts (91%; 62%), and Marine Mammals (96%; 77%) where we see less wild sourcing for live individuals.
1.4 Risks of invasion from wildlife trade
Of the thousands of species listed in LEMIS, many potentially pose a biological threat to native ecosystems by acting as invasive species. Of the world’s top 100 invasive species 25, 28 were recorded as imported live into the US within LEMIS. This includes continued (i.e., 2020-2022) trade of species like zebra mussels (Dreissena polymorpha) that have already led to detrimental ecological, environmental, and economic consequences in the US 26. In total, 203 known invasive species were imported into the US based on species listed as invasive in the US by the IUCN invasive species database (see methods). In addition, the Lacey Act has a shortlist of injurious invasive species 27 that includes 57 genus level listings and 34 species level listings. Of these, at least 23 species were listed as being imported live into the US on at least some occasions after regulatory changes. For example, 12 of the listed fish genera continue to be imported live, and of the 20 amphibian genera listed as injurious in 2016, seven were imported live after that date (and in addition to these imports, exports of all listed “injurious invasive” groups increased). Whilst this may have been for conservation or educational purposes, it underscores the need for further data in such systems.