Primate model

I S S U E S

The species dilemma and its potential impact on enforcing wildlife trade laws

Rachel L. Jacobs | Barry W. Baker

U.S. Fish & Wildlife Service, National Fish &

Wildlife Forensics Laboratory, Ashland,

Oregon

Correspondence

Rachel L. Jacobs and Barry W. Baker,

U.S. Fish & Wildlife Service, National Fish &

Wildlife Forensics Laboratory, 1490 East Main

Street, Ashland, Oregon 97520.

Emails: [email protected];

[email protected]

Abstract The varied answers to the question “What is a species?” provoke more than lively debates in

academic circles. They pose practical problems for law enforcement. Commercial wildlife trade

threatens many primate species and is regulated through such laws and international agree-

ments as the U.S. Endangered Species Act and the Convention on International Trade in Endan-

gered Species of Wild Fauna and Flora. Enforcing legislation relies on the ability to identify

when violations occur. Species-defining characters may not be preserved in wildlife trade items.

For example, pelage patterns and behavioral characters (e.g., vocalizations) are absent from

skulls. Accordingly, identifying victims of illegal trade can be difficult, which hinders enforce-

ment. Moreover, identifying new species and “splitting” of currently recognized species can

result in enforcement lags and regulatory loopholes. Although such negative consequences

should not hinder scientific advancement, we suggest that they be considered by primate taxon-

omists and provide recommendations to prevent unintended conservation consequences.

KEYWORDS

CITES, forensics, law enforcement, phylogenetic species concept, primate, taxonomy

1 | INTRODUCTION

Based on multiple taxonomic references, the latest book All the

World's Primates (AWP) identifies 505 primate species.1 Since its pub-

lication in 2016, more species have been described.2–7 The latter

group even includes a newly recognized species of great ape, Pongo

tapanuliensis.2 New species descriptions often reignite a debate that

has long plagued biological sciences: “What is a species?” How much

distinctiveness is enough to warrant a new name or to elevate a sub-

species to species rank?8–10 Species concepts are often revisited,8–10

and the implications of taxonomic changes on conservation are often

discussed.9,11–14

For primates, the increasing number of recognized species is

largely because of the wider acceptance of the Phylogenetic Species

Concept (PSC).8–10,14,15 This concept defines a species as a diagnos-

able group with a pattern of ancestry and descent.16,17 Some authors

suggest that the use of the PSC in primate taxonomy is largely driven

by conservation motives.8 Regardless of the motives for using the

PSC, there are potential positive and negative impacts for conserva-

tion. For example, the PSC arguably enhances biodiversity protection

by identifying many previously unrecognized species, thus allowing

conservation initiatives to be applied to more ecologically and evolu-

tionarily diverse populations.12,13 Conversely, conservation options

can become more limited as smaller units (e.g., single populations)

receive species status by precluding genetic rescue between newly

distinguished taxa and thus increase inbreeding.11

Unfortunately, one conservation implication is often overlooked:

the effect that changing taxonomy has on enforcing laws aimed at

protecting wildlife. Importantly, this issue applies to a broad range of

taxonomic groups, but primates are exemplary for (a) having a large

number of species newly recognized in recent years (e.g., between

1996 and 2016, the number of primate species increased from ~230

to ~500)1,18 and (b) experiencing large population declines due to

hunting for the wildlife trade.19,20 In discussing this issue, our goal is

not to argue in favor of one species concept over another, nor is to

express our opinions about whether certain taxonomic changes are

warranted. We are forensic scientists working in the United States on

wildlife crimes; these crimes often involve violations of trade laws that

include international species. As wildlife forensic scientists,21 we are

tasked with identifying victims of the crimes. We aim to give a victim

Received: 29 March 2018 Revised: 11 July 2018 Accepted: 21 September 2018

DOI: 10.1002/evan.21751

Published 2018. This article is a U.S. Government work and is in the public domain in the USA.

Evol Anthropol. 2018;27:261–266. wileyonlinelibrary.com/journal/evan 261

a name, a species name when possible. Species-level identification is

often necessary to determine whether or not a crime has been com-

mitted, as well as if and how perpetrators will be prosecuted. Changes

in taxonomy can therefore change the name we are able to give a vic-

tim, which can hinder prosecution, because legislation often lags

behind scientific advances. Moreover, when new species are named,

researchers might not yet know the impact (if any) that hunting pres-

sure for the trade has (or has had) on populations. Accordingly, data

might be lacking to make appropriate recommendations for trade

regulations, resulting in inadequate protections. By making primate

taxonomists aware of these issues, and of the challenges we face in

wildlife forensic science, we hope that the academic community will

be able to help prevent potential undesired conservation threats.

2 | WILDLIFE REGULATIONS AND SPECIES NAMES

Trade in all primates is regulated internationally under the Convention

on International Trade in Endangered Species of Wild Fauna and Flora

(CITES), which was established in 1973 to ensure that international

trade in specimens of wild animals and plants does not threaten their

survival.22,23 Decisions to list species under CITES currently involve

183 Parties (i.e., 182 countries and one regional economic integration

organization, the European Union), which are also tasked with enfor-

cing the regulations.23 These CITES listings do not reflect the species'

conservation status as determined by the International Union for Con-

servation of Nature (IUCN) Red List of Threatened Species. CITES

focuses only on species impacted by trade, whereas IUCN assesses

species' conservation statuses (e.g., Vulnerable, Endangered, Critically

Endangered) based on a wide range of threats (http://www.

iucnredlist.org/about/overview). There is collaboration between these

two entities.24,25 For example, IUCN is involved in reviews of pro-

posed CITES listing amendments for consideration prior to CITES

decisions.25

Many species are also specifically listed under the

U.S. Endangered Species Act (ESA), which regulates, among other

activities, national and international trade in threatened and endan-

gered species involving U.S. jurisdiction;23,26 the list includes non-

native species (https://ecos.fws.gov/ecp/; note that “threatened” and

“endangered” status in ESA also differs from that of IUCN, see

ESA definitions at https://www.fws.gov/endangered/laws-policies/).

Finally, many primates may receive protections under laws within

their range countries.27–29 Importantly, legislation related to conserva-

tion and wildlife trade generally defines protections and regulations

based on species names (or in some cases subspecies names). Among

primates, trade regulations differ across species. For example, under

CITES, international commercial trade in live primates, as well as their

parts and derivatives, is either strictly prohibited (appendix I) or per-

mitted but regulated (appendix II).22,30 The difference between these

two categories has important implications for prosecuting violations

of wildlife trade laws. However, the primate remains we often observe

in the trade, including skulls, body parts (e.g., hands and feet), bush-

meat, and fragmentary postcranial material, can be difficult to identify

to species level based on morphology and/or genetics (Figure 1).21

Accordingly, when closely related taxa are split listed, that is, when

they contain both appendices I and II taxa (e.g., Alouatta, Ateles, and

Saguinus),22,30 enforcing appendix I violations can be difficult. ESA vio-

lations, where only a limited number of species are listed,26 can be

equally problematic to enforce.

3 | WILDLIFE FORENSICS AND SPECIES IDENTIFICATION

If species can be identified as requiring some form of legal protection,

then how can it be that identifying animals to species is so difficult?

From a forensics point of view, there are multiple related reasons. First,

identifications must be ironclad and defensible in a court of law. The

livelihoods and liberty of the accused are on the line, so identifications

must be certain. If an evidence item does not exhibit clear characters

diagnostic to species, then identifications must be made to the next

lowest taxonomic grouping possible (e.g., genus or family). Second,

most evidence items are not associated with accurate geographic infor-

mation, and thus, such data cannot be used in identifications. Third,

primate species are defined based on a variety of criteria. The “diagno-

sability” aspect of the PSC is not limiting;16,17 different species may be

considered diagnosable based on genetic, morphological, or behavioral

characters. If those criteria are not preserved in an evidence item, as

frequently happens (e.g., pelage coloration is generally not preserved in

bushmeat material), then identification to species level is not possible.

Finally, species descriptions are often inadequate for forensic identifica-

tions. There are a number of reasons why this might be the case. Spe-

cies names are tied to the type specimen(s),31 and thus the full range of

variation within a described species may be unknown. For morphologi-

cal analyses, there may be no (or low quality) illustrations of species-

diagnostic differences. When measurements are critical, raw data may

be unavailable (species means are of limited value), and often, it is

unclear what is considered to be a description compared to a diagnosis,

especially when synapomorphic and autapomorphic traits are not

clearly defined. For genetic analyses, internal reference specimens or

published sequences of congeners may be unavailable for comparison.

4 | TAXONOMIC CHANGES AND ENFORCING CURRENT REGULATIONS

Many of these challenges to making species identifications are com-

pounded when there are taxonomic splits, in which multiple species

are recognized from what was previously recognized as one or few

species. Again, we do not aim to imply that such changes should not

be made; science should not have an agenda. Although some

researchers have argued for greater oversight over taxonomic

changes,32,33 others have emphasized the dangers such oversight

could have on scientific progress.31 Our aim is to simply point out that

such splits have consequences beyond those related to academic

research. One major forensic complication that arises from taxonomic

splits is related to interpreting previously described variation. As men-

tioned earlier, forensic wildlife evidence often includes modified

and/or partial remains. Because species descriptions may be

262 JACOBS AND BAKER

inadequate for species-level identification of such items, we often

incorporate comparative research of particular elements/features into

our evaluations. It can be difficult to assess whether variation that

was previously observed and described in a species is valid after a tax-

onomic split. If this is not explicitly addressed in the literature, or if

raw data and their associated specimens are not available, then previ-

ously published data under earlier taxonomies are of minimal use for

forensic identifications. This issue not only impacts our use of pub-

lished research but also our use of forensic comparative collections.

By the nature of evidence items, forensic reference collections are

often atypical compared to museum collections. Similarly, however,

species names must be reassessed in light of taxonomic changes,

which can result in the same challenges as evaluating evidence items.

Importantly, beyond forensic identification, changing taxonomy

can have additional consequences related to prosecution and species

protection. Changing taxonomy can introduce potential loopholes in

law enforcement, because new species names are not necessarily rec-

ognized by current laws. In such cases, prosecution might be hindered

and/or it might be necessary to reevaluate the intent of a law, which

can take time. For example, the scientific community has recognized

Sumatran and Bornean orangutans as separate species, rather than two

subspecies, since around 2001.34 It was not until 2018 that the inter-

pretation of the ESA listing, Pongo pygmaeus, was addressed to also

include Pongo abelii.35 Fortunately, given the potential new species

Pongo tapanuliensis,2 the statement further addresses protection of any

additional species that might be recognized in the future,35 a welcome

addition to the legislation that we would like to see become more wide-

spread. CITES appears to update species names more regularly.31 For

example, taxonomic revisions to the genus Pithecia in 201436 are recog-

nized in the online CITES Species+ website, www.speciesplus.net30.

When taxonomic revisions are accepted, then these species receive

similar listing status as the taxon from which it was split, which may or

may not be appropriate. It should also be noted that outside of the

United States and CITES, range country laws can also be impacted by

changing taxonomy. For example, a recent article indicated that Trachy-

pithecus and Semnopithecus (formerly Presbytis) are not formally

protected under Chinese law, which recognizes the genus Presbytis

only, introducing a potential line of defense within the court system.27

We are unaware of whether or not the laws have changed since the

publication. Given that many countries suffer from very low levels of

prosecution for wildlife crimes,37–39 such obstacles could make prose-

cution even more difficult. Because there can be an extensive amount

of time between a potential violation and actual prosecution, taxonomic

changes could even impact current and ongoing cases.

5 | TAXONOMIC CHANGES AND POTENTIAL NEED FOR NEW REGULATIONS

Additional impacts related to species protection include the unfortu-

nate potential for increased pressure on newly recognized species as

a consequence of their perceived “novelty.”40 Rare primates are some-

times targeted in wildlife trade (the pet trade in particular) as status

FIGURE 1 Examples of confiscated primate material. (a) Hand of Pan sp. (b) Skull of immature Pongo sp. (c) Dried infant Nycticebus sp. (d) Partial

skin/pelage of Nycticebus sp. Scale bars = 1 cm

JACOBS AND BAKER 263

symbols.28,41–43 New species, regardless of their population numbers,

could be similarly targeted, and this trend has been noted to occur in

other, nonprimate taxa.40 Finally, as mentioned earlier, trade in newly

recognized species might not be appropriately regulated. In cases of

taxonomic splits, the impact of wildlife trade on previously recognized

widely distributed species might be minimal, but newly recognized

species with smaller populations could be (or could have been) more

severely impacted, as has been suggested for the genus Aotus.28,29

Aotus (night monkeys) has undergone several taxonomic revisions,

from which one species is now often recognized as 11.29 Legal trade

in Aotus species (CITES appendix II; nine species recognized) is rela-

tively high among primates and might impart unsustainable pressure

on rarer species.29 Importantly, it is difficult to actually assess the

impact of trade on Aotus species, in part, because not all species are

currently recognized by CITES and not all traded animals (legal and

illegal) are identified to species.29 Moreover, identification to Aotus

species can be difficult for law enforcement due to morphological sim-

ilarities.29 In such cases, if a change in regulation status is decided for

some but not all species, it could be difficult to enforce. Accordingly,

changes to trade regulations may need to occur at the genus level

(or higher), as was successfully done for the genus Nycticebus. In

recognition of the large impact that the wildlife trade has on slow

lorises, as well as the ongoing taxonomic changes within this group,

the genus Nycticebus was elevated to appendix I under CITES in 2007

(i.e., all trade prohibited).44,45

When legislative changes appear warranted, members of the aca-

demic community can be advocates for change. For example, they can

regularly check and comment on Federal Register documents (https://

www.federalregister.gov/). They can provide concerns about CITES

listing statuses to the governments of primate range countries in

which they work or nongovernmental agencies that are observers at

CITES conferences; changes to regulations require proposals from

Parties.23 It is important to note, however, that recommendations for

legislative changes should be carefully considered. For example, spe-

cies that are deemed to be threatened with extinction by the IUCN

(e.g., Endangered, Critically Endangered) might not be severely threat-

ened by wildlife trade. Decisions to elevate species under the CITES

trade agreement could have negative conservation consequences by

stimulating/increasing trade.46

Clearly, the relationships among changing taxonomy, legislation,

and law enforcement are complex. In recognition, some have argued

for refinements to legislation that accommodate changing

TABLE 1 Suggestions for researchers to aid forensic identification of species in the wildlife trade

General guidelines

Be aware that taxonomic changes can have law enforcement implications.

Be aware of range country laws and taxonomy used in those legal systems.

Ensure range countries are aware of any previously unknown occurrences of species within their borders, so that protective legislative action can be considered.27 However, be aware that public geographic information can pose significant threat to species.40

Consider that changes to legislation and trade agreements might be better made prior to publishing taxonomic revisions/species descriptions.53

Be aware that within many countries, there are limited resources for wildlife law enforcement. Accordingly, many forensic analyses (e.g., genetics) might not be possible.29 Morphological identification when feasible is faster and cheaper. That said, requirements for forensic identifications might differ among countries, with certain techniques being favored or required over others. Researchers should communicate with local governments and international trade authorities to discuss the impacts of species identifications and taxonomic changes.29

Be aware that morphological and genetic reference samples are needed for forensic species identification. Lack of access to such material is often a limiting factor in successful law enforcement efforts.

Recommendations

Species descriptions

Be explicit and detailed in describing, illustrating, and publishing morphological diagnoses.

Clearly distinguish between morphological descriptions and morphological diagnoses. Synapomorphies and autapomorphies should be clearly described and illustrated.

Note species that might be confused with a newly described/elevated species so that the need for broader protection/regulation (e.g., genus level) might be better assessed.

Be explicit on synonymy to help legal systems interpret your publications.

Make vouchered photographic databases publicly available.a

Make diagnostic vouchered mitochondrial and nuclear sequences publicly available (e.g., through online databases, such as GenBank).a

Create identification guides for species based on taxonomic revisions in the languages of species range countries, as well as those countries potentially involved in trade import and transit.

General research

Publish research on basic morphological description and genetic characterization. These might not seem to be the most ground-breaking publications, but they are important. In fact, basic descriptive morphological or genetic differences are often more useful in a forensic context than are functional/evolutionary explanations of the characters observed.b

If possible, fully document variability for species, sexes, and age ranges. Morphological outliers are often submitted as forensic evidence because their appearance is so unusual.

Make raw morphometric data and their associated specimen numbers publicly available.a

a Many researchers and publishers are already providing public access to published data. Publishers can help ensure data are publicly available and accessi- ble by requiring deposition prior to official publication and release of new species names.

b We acknowledge that publishers may be a barrier if they deem such studies as low impact or narrow in scope. Journals specifically aimed at publishing data (e.g., BMC Research Notes) might help overcome this obstacle.

264 JACOBS AND BAKER

taxonomy.31 It is also helpful if legislative decisions consider the

unique impacts taxonomic changes can have on wildlife forensic iden-

tification (e.g., whether species-level identification is feasible). Unfor-

tunately, legislative change can be slow. In the meantime, there are

steps that researchers can take to help reduce the challenges related

to forensic identification (Table 1).

In closing, we note again that this issue can be more broadly

applied to other biological taxa. As wildlife forensic scientists, we are

tasked with identifying a much broader group of animals (RLJ—all

mammals; BWB—all herpetofauna), and changing taxonomy, in partic-

ular taxonomic splitting, is widespread.47–52 Through increased aware-

ness and continued discussion, we hope to bridge the gap between

alpha taxonomy and its applied use in a forensic context. Many pri-

mate species face staggering challenges to their survival.19,20 It is

therefore imperative that academic and applied researchers work

together when possible to help reduce some of these threats.

The findings and conclusions in this article are those of the

author(s) and do not necessarily represent the views of the U.S. Fish

and Wildlife Service.

ACKNOWLEDGMENTS

The authors thank Pepper Trail and Thomas Leuteritz for helpful com-

ments on earlier versions of this manuscript.

CONFLCIT OF INTEREST

The authors declare that there are no conflicts of interest.

ORCID

Rachel L. Jacobs https://orcid.org/0000-0001-6075-7767

Barry W. Baker https://orcid.org/0000-0001-7037-5682

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AUTHOR BIOGRAPHIES

RACHEL JACOBS is a forensic mammalogist with a background in biologi-

cal anthropology. Her past and ongoing research has focused on the

behavior, ecology, population genetics, and morphology of living

primates.

BARRY BAKER is a wildlife forensic scientist with a background in

zooarchaeology and biological anthropology. His past and ongoing

research has focused on skeletal morphology and the forensic identifi-

cation of reptiles and mammals in the wildlife trade.

How to cite this article: Jacobs RL, Baker BW. The species

dilemma and its potential impact on enforcing wildlife trade

laws. Evol Anthropol. 2018;27:261–266. https://doi.org/10.

1002/evan.21751

266 JACOBS AND BAKER