2170 Federal Register / Vol. 90, No. 6 / Friday, January 10, 2025 / Rules and Regulations 

DEPARTMENT OF THE INTERIOR 

Fish and Wildlife Service 

50 CFR Part 16 

RIN 1018–BA77 

[Docket No. FWS–HQ–FAC–2015–0005; 

FXFR13360900000–245–FF09F14000] 

Injurious Wildlife Species; Listing 

Salamanders Due to Risk of 

Salamander Chytrid Fungus 

AGENCY: Fish and Wildlife Service, 

Interior. 

ACTION: Affirmation of interim rule as 

final; second interim rule and request 

for public comment. 

SUMMARY: The U.S. Fish and Wildlife 

Service is affirming as final the 2016 

interim rule that added all species of 

salamanders from 20 genera to the list 

of injurious amphibians. Under the 

injurious wildlife prohibitions of the 

Lacey Act, this final rule prohibits the 

importation into the United States and 

shipment between the continental 

United States, District of Columbia, 

Hawaii, Commonwealth of Puerto Rico, 

or any territory or possession of the 

United States of any live or dead 

specimen, including hybrids and parts, 

of those 20 genera of salamanders, 

except by permit for certain purposes or 

by Federal agencies solely for their own 

use. In addition to finalizing the listing 

of those 20 genera, we are publishing a 

new interim rule to add to the injurious 

amphibian list 16 genera that recent 

studies determined are also carriers of 

the fungus and to clarify some 

provisions from the final rule. This 

interim rule includes any live or dead 

specimen, hybrid, or parts of the 16 

genera and opens a public comment 

period. We take these actions to protect 

U.S. ecosystems from the introduction, 

establishment, and spread of the lethal 

chytrid fungus Batrachochytrium 

salamandrivorans, which infects and is 

carried by salamanders, and which is 

not yet known to be found in the United 

States. 

DATES: Effective date: The interim rule 

published at 81 FR 1534 on January 13, 

2016, was effective January 28, 2016. 

This final rule affirming the January 13, 

2016, interim rule and the interim rule 

set forth in this document are effective 

January 25, 2025. 

Comment submission: Interested 

persons are invited to submit written 

comments on the issues raised in the 

second interim rule as described below 

under Information Requested on or 

before March 11, 2025. 

ADDRESSES: Comment submission: You 

may submit comments by one of the 

following methods: 

(1) Electronically: Go to the Federal 

eRulemaking Portal: https://

www.regulations.gov. In the Search box, 

enter FWS–HQ–FAC–2015–0005, which 

is the docket number for this action. 

You may submit a comment by clicking 

on ‘‘Comment.’’ 

(2) By hard copy: Submit by U.S. mail 

to: Public Comments Processing, Attn: 

FWS–HQ–FAC–2015–0005, U.S. Fish 

and Wildlife Service, MS: PRB/3W, 

5275 Leesburg Pike, Falls Church, VA 

22041–3803. 

We request that you send comments 

only by the methods described above. 

We will post all comments on https:// 

www.regulations.gov. This generally 

means that we will post any personal 

information you provide us (see 

Information Requested, below, for more 

information). 

Supplementary materials: 

Background documents related to this 

rulemaking action, including the final 

economic analysis for the affirmation of 

the 2016 interim rule, are available at 

https://www.regulations.gov in Docket 

No. FWS–HQ–FAC–2015–0005. 

FOR FURTHER INFORMATION CONTACT: 

Kristen Sommers, Injurious Wildlife 

Listing Coordinator, U.S. Fish and 

Wildlife Service, Branch of Aquatic 

Invasive Species; MS: FAC; 5275 

Leesburg Pike; Falls Church, VA 22041– 

3803; 571–329–2214. Individuals in the 

United States who are deaf, deafblind, 

hard of hearing, or have a speech 

disability may dial 711 (TTY, TDD, or 

TeleBraille) to access 

telecommunications relay services. 

Individuals outside the United States 

should use the relay services offered 

within their country to make 

international calls to the point-ofcontact in the United States. 

SUPPLEMENTARY INFORMATION: 

Table of Contents 

I. Executive Summary 

II. Current Rulemaking Action 

III. Final Rule to the 2016 Interim Rule 

A. Background 

B. Summary of Comments Received on the 

2016 Interim Rule 

C. Affirmation of the 2016 Interim Rule 

D. Required Determinations 

IV. Second Interim Rule 

A. Species Information for Salamanders 

B. Species Information for 

Batrachochytrium salamandrivorans 

C. Population-Level and Ecosytem-Level 

Effects of Bsal 

D. Invasiveness of Salamanders and Bsal 

E. Pathway Analysis 

F. Risk Assessments of Bsal 

G. Factors That Contribute to Injuriousness 

of Salamanders 

H. Measures That Reduce or Remove 

Injuriousness of Salamanders 

I. Summary and Conclusion for Interim 

Rule 

J. Required Determinations 

K. Information Requested 

I. Executive Summary 

Why we need to publish a rule. We, 

the U.S. Fish and Wildlife Service 

(Service or FWS), are charged with 

administering 18 U.S.C. 42(a), as 

amended (commonly known as the 

injurious wildlife provision of the Lacey 

Act). Under this authority, the Secretary 

of the Interior may list by regulation 

those wild mammals, wild birds, fish, 

mollusks, crustaceans, amphibians, 

reptiles, and the offspring or eggs of any 

of the foregoing that are injurious to 

human beings, to the interests of 

agriculture, horticulture, or forestry, or 

to the wildlife or wildlife resources of 

the United States. 

We have determined that salamanders 

that can carry the fungus 

Batrachochytrium salamandrivorans 

(Bsal) are injurious to wildlife and 

wildlife resources of the United States. 

This determination was based on a 

review of the literature and an 

evaluation under the criteria for 

injuriousness by the Service. The 

purpose of listing these species as 

injurious wildlife is to prevent the 

introduction, establishment, and spread 

of Bsal in the wild in the United States. 

The fungus primarily affects 

salamanders, has lethal effects on many 

salamander species, and is not yet 

known to be found in ecosystems of the 

United States. 

What this document does. This 

document serves two purposes: It 

finalizes a rulemaking action initiated 

with publication of an interim rule in 

2016, and it promulgates a new and 

related interim rule. 

On January 13, 2016, we published an 

interim rule that amended our 

regulations pertaining to injurious 

wildlife (81 FR 1534). That interim rule 

(hereafter referred to as ‘‘the 2016 

interim rule’’) amended our regulations 

to add all species of salamanders from 

20 genera, of which there were 201 

species, to the list of injurious 

amphibians. Under the injurious 

wildlife prohibitions of the Lacey Act, 

the 2016 interim rule prohibited both 

importation into the United States and 

interstate transportation between States, 

the District of Columbia, the 

Commonwealth of Puerto Rico, or any 

territory or possession of the United 

States (the latter was clarified by a court 

decision in 2017) of any live or dead 

specimen, including parts, of these 20 

genera of salamanders, except by permit 


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for zoological, educational, medical, or 

scientific purposes (in accordance with 

permit conditions) or by Federal 

agencies without a permit solely for 

their own use. 

A second interim rule is now needed 

because of recent studies documenting 

additional genera that also share the 

same traits that make them injurious as 

carriers of Bsal. We are also revising 

some provisions from the 2016 interim 

rule in the final rule to make minor 

corrections to and improve clarity of the 

rule. 

The basis for our action. Defensible 

scientific evidence indicates that we 

need to list a total of 36 genera of 

salamanders as injurious wildlife to 

protect U.S. ecosystems. Therefore, we 

hereby affirm the injurious wildlife 

listings in the 2016 interim rule of all 

species in the following 20 genera: 

Chioglossa, Cynops, Euproctus, 

Hydromantes, Hynobius, Ichthyosaura, 

Lissotriton, Neurergus, Notophthalmus, 

Onychodactylus, Paramesotriton, 

Plethodon, Pleurodeles, Salamandra, 

Salamandrella, Salamandrina, Siren, 

Taricha, Triturus, and Tylototriton. We 

also add new injurious wildlife listings 

of all species in the following 16 genera 

through the second interim rule: 

Ambystoma, Andrias, Aneides, 

Aquiloeurycea, Calotriton, 

Chiropterotriton, Cryptobranchus, 

Desmognathus, Ensatina, Eurycea, 

Laotriton, Ommatotriton, Pachytriton, 

Proteus, Pseudobranchus, and 

Pseudotriton. 

The United States has the greatest 

diversity of salamanders in the world, 

salamanders are a vital part of native 

ecosystems, and numerous salamander 

populations are at risk of endangerment 

from Bsal. A risk assessment conducted 

by the U.S. Geological Survey (USGS) 

concluded that the potential for Bsal 

introduction into the United States is 

high, the United States has suitable 

conditions for Bsal survival, and the 

consequences of introduction into the 

United States are expected to be severe 

and occur across a wide geographic 

range within the United States. The 

most likely pathway of Bsal into the 

United States would be on the bodies of 

salamanders in the commercial 

salamander trade. Aside from our Bsal 

regulations, the ability and effectiveness 

of measures to prevent or control Bsal 

is currently low. Trade in wildlife 

occurs on a global scale, and 

amphibians are one of the most 

commonly traded animals. 

Therefore, listing the genera as 

determined in this rulemaking action 

will help to reduce the likelihood that 

Bsal enters the United States and 

presents a threat to native salamander 

species. 

II. Current Rulemaking Action 

This document does the following: 

• Affirms the current listing of 20 

genera of salamanders as injurious 

species by the 2016 interim rule as 

described above and any species within 

those genera. 

• Revises provisions in the preamble 

of the 2016 interim rule in response to 

a court decision that pertained to 

interstate transport of injurious wildlife 

as described below. 

• Removes the 201 itemized species 

names from the list in 50 CFR 16.14. 

This itemized list of scientific and 

common names is unnecessary because 

the regulations in 50 CFR 16.14(a) state 

that the prohibitions pertain to the 20 

genera ‘‘including but not limited to, the 

species listed in this paragraph.’’ We 

provided the itemized list of species in 

the 2016 interim rule largely for the 

convenience of the public and our law 

enforcement staff, but the taxonomy of 

salamanders is evolving, and the list is 

not static. However, while many 

scientific and common names have 

changed, all of the listed species remain 

in their same genera. 

• Clarifies prohibitions pertaining to 

hybrids and frozen specimens; clarifies 

what is not prohibited (including eggs 

or gametes; parts or tissues that have 

been chemically preserved, chemically 

treated, or heat treated so that the 

pathogen Batrachochytrium 

salamandrivorans, if present, is 

rendered non-viable; and molecular 

specimens consisting of only the nucleic 

acids from organisms). 

• Adds 16 genera to the list in 50 CFR 

16.14(a), as explained below, and 

solicits comments on these new genera. 

III. Final Rule to the 2016 Interim Rule 

A. Background 

On January 13, 2016, under the 

authority of 18 U.S.C. 42(a)(1), as 

amended, we, the U.S. Fish and Wildlife 

Service, published an interim rule in the 

Federal Register (81 FR 1534) to add all 

species from 20 genera to the list of 

injurious amphibians and announced 

the availability of the draft economic 

analysis and the draft environmental 

assessment of the 2016 interim rule. The 

rule took effect on January 28, 2016, and 

revised the lists of injurious wildlife in 

part 16 of title 50 of the Code of Federal 

Regulations (CFR), specifically the list 

of injurious amphibians at 50 CFR 

16.14. The 60-day public comment 

period closed on March 14, 2016. We 

solicited comments and supporting data 

to gain additional information. We also 

solicited peer review at the same time. 

In this document, we present our 

consideration of the public comments 

and peer review received on the 2016 

interim rule to inform our final 

determinations. We present a summary 

of the peer-review comments and the 

public comments and our responses to 

them in the ‘‘Summary of Comments 

Received on the 2016 Interim Rule’’ 

portion of the preamble to this final 

rule. The comments did not provide any 

substantive evidence that supported 

changing the genera in the interim rule. 

However, some comments did provide 

justification for modifying certain 

requirements stipulated in the 2016 

interim rule. 

The Service published an interim rule 

in 2016 instead of issuing a proposed 

rule for the listing under the 

Administrative Procedure Act (APA) (5 

U.S.C. 551 et seq.). As explained in the 

2016 interim rule, we had good cause to 

forgo notice and public comment on a 

proposed rule and instead take 

immediate action in the form of an 

interim rule to help prevent the fungus 

Bsal from being introduced, established, 

or spread in the United States for the 

reasons listed above. The fungus, lethal 

to many salamander species in the 

United States, is carried on the skin of 

salamanders and can be unintentionally 

imported by salamanders in trade. The 

2016 interim listing of the 20 genera of 

salamanders has prohibited the 

importation of high-risk species under 

the injurious wildlife prohibitions of the 

Lacey Act, and the fungus has remained 

absent from the United States. 

In this document, a clarification from 

the 2016 interim rule reflects a court 

decision in 2017. Under the D.C. Circuit 

Court of Appeals decision in United 

States Association of Reptile Keepers, 

Inc. v. Zinke, 852 F.3d 1131 (D.C. Cir. 

2017), import of injurious wildlife into 

the United States remains prohibited. In 

addition, transport of injurious wildlife 

between the enumerated jurisdictions in 

the shipment clause of 18 U.S.C. 42(a) 

(the continental United States, the 

District of Columbia, Hawaii, the 

Commonwealth of Puerto Rico, and any 

territory or possession of the United 

States) remains prohibited. However, 

the court held that 18 U.S.C. 42(a) does 

not prohibit interstate movement 

between States within the continental 

United States. This means that 

transportation of injurious wildlife 

between the 49 States within the 

continental United States (the 

contiguous 48 States and Alaska) is not 

prohibited by the statute, unless that 

movement of the wildlife is restricted 

due to conditions associated with issued 

injurious wildlife permits. We note that 

transport from the lower 48 States to 


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Alaska through Canada, or vice versa, 

remains prohibited, because that 

transport includes an import into the 

United States. 

The language in 50 CFR 16.14(a) was, 

and still is, correct in that it does not 

prohibit interstate transport between 

States within the continental United 

States. The final economic analysis 

affirming the 2016 interim rule reflects 

the clarification of interstate transport 

authority between States within the 

continental United States since the 

court decision on April 7, 2017. 

However, injurious wildlife unlawfully 

imported into the United States or 

transported between the enumerated 

jurisdictions is still unlawful to 

transport within the continental United 

States. Under the Lacey Act 

amendments of 1981, 16 U.S.C. 

3372(a)(1), it is unlawful for any person 

to import, export, transport, sell, 

receive, acquire, or purchase any 

wildlife transported in violation of any 

law of the United States. This includes 

transport of any injurious wildlife 

imported into the United States or 

transported between the enumerated 

jurisdictions in violation of 18 U.S.C. 

42. 

B. Summary of Comments Received on 

the 2016 Interim Rule 

The following comments were 

submitted during the peer and public 

comment period for the 2016 interim 

rule. Knowledge about Bsal has 

improved since then due to the many 

excellent studies by researchers and our 

own understanding of the disease. In the 

following responses to the comments 

under Peer Review Summary and Public 

Comments Summary, respectively, we 

have endeavored to answer the 

comments both as they related to the 

knowledge of Bsal, trade, and related 

issues at the time of the 2016 interim 

rule and as of the current knowledge as 

appropriate. The current knowledge 

from recent research affirms the 2016 

interim rule, supports the second 

interim rule set forth in this document, 

and can be found below in IV. Second 

Interim Rule. In our responses to the 

comments, when we refer to ‘‘this rule,’’ 

we mean that the information and the 

changes apply both to the final rule for 

the 2016 interim rule and the second 

interim rule. 

Peer Review Comments and Our 

Responses 

In accordance with peer review 

guidance of the Office of Management 

and Budget ‘‘Final Information Quality 

Bulletin for Peer Review,’’ released 

December 16, 2004, we solicited expert 

opinion on information contained in the 

2016 interim rule from three 

knowledgeable individuals selected 

from specialists in the relevant 

taxonomic group and ecologists with 

scientific expertise that includes 

familiarity with alien herpetological 

introductions and invasions, predictive 

tools for risk assessment, and invasion 

biology. We posted our peer review plan 

on the Service’s Science website 

(https://www.fws.gov/media/peerreview-plan-listing-salamandersinjurious-due-risk-salamander-chytridfungus), explaining the peer review 

process and providing the public with 

an opportunity to comment on the peer 

review plan. No comments were 

received regarding the peer review plan. 

The Service solicited independent 

scientific reviewers who submitted 

individual comments in written form. 

We avoided using individuals who had 

already expressed strong support for or 

opposition to the subject and 

individuals who were likely to 

experience personal gain or loss (such 

as financial or prestige) or otherwise 

could be perceived as having a conflict 

of interest as a result of the Service’s 

decision. We received responses from 

three peer reviewers. A scientist with 

the USGS served as one of the peer 

reviewers. 

We requested that the reviewers 

provide comments that were specific to 

the 2016 interim rule and the draft 

economic analysis. We reviewed all 

comments for substantive issues and 

any new information they provided. We 

consolidated their comments (without 

attribute) and our responses into key 

issues in this section. Some peer 

reviewer comments that called for 

technical changes or more minor 

corrections have not been noted, but we 

have made our best effort to correct 

those grammatical or biological errors 

and clarify certain ambiguous 

statements in the second interim rule 

and supporting documents. We 

prepared the second interim rule and 

second draft economic analysis to 

reflect peer reviewer comments and new 

scientific information where 

appropriate. 

The comments we received indicated 

support for the 2016 interim rule and for 

the analysis that we conducted given 

the need to prevent harm to native 

species from Bsal. All three peer 

reviewers concluded that the data and 

analyses we used in the interim rule 

were appropriate and the conclusions 

we drew were logical and reasonable. 

All three peer reviewers provided 

additional insights (which we used to 

clarify points in the second interim rule) 

or references to recently published 

studies, which support the final rule. In 

general, the peer reviews supported the 

Service’s conclusions and agreed that 

they were reasonable, though they did 

note that we generalized some of our 

findings, such as the average 

temperature of salamander habitats as 

an indicator of Bsal vulnerability with 

regards to salamanders nationwide. We 

have clarified these issues where 

practical in the second interim rule. The 

peer reviewers suggested that there was 

a need to expand our discussion 

regarding possible treatment options, 

which we have added to the second 

interim rule. The peer reviewers also 

acknowledged that, while the rule is 

important, research questions remain 

that could shed light on ways to better 

prevent the introduction of Bsal into the 

United States. 

General Comments 

(PR1) Comment: In support of the 

2016 interim rule, prevention provides 

an environmental and biodiversity 

benefit; the probability of introduction 

is at the very least reduced; and host 

species need not become established in 

the environment to transmit Bsal. If an 

owner houses multiple salamander 

species, transmission can occur in 

captivity to other species that may be 

able to establish a population in the 

wild. The commenter also agrees with 

the exclusion from the rule of tailless 

amphibians (frogs and toads), which 

were uniformly resistant in the tests by 

Martel et al. (2014). 

Response: We concur with the 

comments. At the time of the 2016 

interim rule publication, we lacked 

evidence of tailless amphibians as 

carriers, and this final rule simply 

affirms our findings on salamanders. 

(PR2) Comment: The average 

temperature of salamander habitats as 

an indicator of Bsal vulnerability with 

regard to salamanders nationwide 

would be difficult to defend. The rule 

provides a very broad average, and 

extrapolating it weakens the point. 

Response: We have clarified the rule 

under Bsal Risk Assessments in 

response to this comment to reflect 

more specifically the areas that the risk 

assessments identified as highest risk. 

We intended to demonstrate that most 

salamander species in the United States 

are not protected from Bsal by living 

outside of the Bsal optimal growth range 

or in areas beyond the threshold where 

Bsal can survive. 

(PR3) Comment: If time allowed, a 

few simple tests to improve the 

scientific foundation of the interim rule 

could have strengthened the decision to 

include or exclude species. 

Response: The Service is not a 

research agency, and we utilized 


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available research when we found in 

2016 that there was good cause to forgo 

notice and public comment on a 

proposed rule and instead take 

immediate action in the form of an 

interim rule to help prevent Bsal from 

being introduced, established, or spread 

in the United States. In this rule, we also 

utilize newly published or otherwise 

available research. In the years since the 

2016 interim rule was published, many 

of the relevant studies affirmed our 

interim findings, while others support 

additional genera as documented in our 

second interim rule. None substantively 

contradicted our findings. 

(PR4) Comment: Given the long time 

that Bsal has been around (150 years), 

the massive number of imported 

salamanders, and only recent 

characterization of Bsal, it may be 

possible that earlier Bsal was 

characterized as a related chytrid 

fungus, Batrachochytrium dendrobatidis 

(Bd). 

Response: As of the completion of this 

rule, resource managers, scientists and 

other researchers have been utilizing the 

latest scientific techniques to actively 

look for Bsal in nonresearch captive 

populations and in wild populations in 

the United States. The USGS conducted 

a massive sampling effort of 11,189 

samples from 594 sites in 223 counties 

within 35 U.S. States and 1 site in 

Mexico specifically for Bsal in wild 

populations (Waddle et al. 2020). The 

sites were chosen based on the species’ 

susceptibility (including some frog 

species) and highest risk geographically. 

No Bsal was found. As of the 

completion of this rule, we are unaware 

of any positive Bsal detections in the 

wild. Testing of archived samples by 

other laboratories has been done, and no 

Bsal has been detected to our 

knowledge. The evidence is not 

conclusive that Bsal has never been in 

the wild in the United States, but there 

is no evidence that it has. 

(PR5) Comment: When the rule 

repeats the information about invasive 

species, the point should be made that, 

even if a salamander found to be 

injurious could not establish a 

population in the wild, an infected 

salamander in captivity can still 

transmit Bsal to native populations if 

that salamander escapes or if material 

touching it is disposed of improperly. 

Response: We agree and have 

incorporated the suggested language 

into the second interim rule. 

(PR6) Comment: The 2016 interim 

rule states that the main pathway for the 

global spread of Bsal is the international 

trade in salamanders and that the most 

likely pathway of a salamander that is 

a host to Bsal into the United States 

would include a pet store or online 

retailer. Since neither the United 

Kingdom or Germany Bsal outbreaks 

were related to a pet store or online 

retailer, the commenter recommends 

stating, ‘‘The most likely pathway of a 

salamander that is a host to Bsal into the 

United States would be the captive 

salamander commercial trade’’ to cover 

the diverse salamander trade. 

Response: We have edited the second 

interim rule accordingly. 

(PR7) Comment: Given that Bd was 

probably introduced by release of 

laboratory animals as well as pets, 

institutional use of listed salamanders 

should also be regulated to protect U.S. 

ecosystems from Bsal. 

Response: This rule is intended to 

prevent the introduction of Bsal, 

whereas Bd was already widespread in 

the United States when that fungus was 

identified as the cause of major 

amphibian mortality. Importation of 

listed salamanders is regulated by this 

rule within the authorities under the 

injurious wildlife provisions of 18 

U.S.C. 42. Any listed salamanders that 

are imported under a permit exception 

for zoological, educational, medical, or 

scientific purposes are required to 

observe sanitary procedures and double 

containment to prevent escape and are 

not allowed to be released. The Service 

may also establish additional permit 

conditions if deemed appropriate to 

ensure responsible use, maintenance, 

and containment of injurious wildlife 

specimens posing a risk of pathogen 

transfer and continued protection of the 

public interest and health, under 18 

U.S.C. 42(a)(3) and the Service’s 

permitting regulations in 50 CFR part 13 

and part 16. 

(PR8) Comment: What is the citation 

for the statements, ‘‘Based on scientific 

evidence, we know that the fungus is 

lethal to at least two salamander species 

native to the United States. Of the 190 

native U.S. species, we find that at least 

67 species are carriers and 20 are not 

carriers’’? 

Response: Martel et al. 2014 was the 

source for the first sentence, and we 

used a combination of sources for 

information about native species and 

the testing that was done. We stated our 

sources and findings for these 

statements in the relevant sections in 

this rule. 

Vulnerability and Carrier Status of 

Native Species 

(PR9) Comment: One of the 

considerations was that, even if a 

salamander listed by the 2016 interim 

rule could not establish a population in 

the wild, an infected salamander in 

captivity (or the water and soil in which 

it came into contact) can transmit Bsal 

to native populations. In addition to 

water and soil, how about if there is 

affected foliage or paper that was used 

in transit? 

Response: The commenter is correct 

that a variety of materials could become 

contaminated with fungal spores if in 

contact with infected salamanders. 

However, it is not possible to provide a 

complete list of these potential fomites 

(materials, such as water, that can act as 

passive carriers and can transfer 

pathogens) in the text of the rule. 

Rather, we believe that listing the 

salamander species that may be carriers 

of Bsal as injurious wildlife, thereby 

prohibiting the importation of 

potentially infected individuals, reduces 

the risk for pathogen spread by any 

substrate. 

(PR10) Comment: The considerations 

say that controlling Bsal is not practical. 

The peer reviewer recommends revising 

to note that, while there are control 

methods available for infected 

individual salamanders in captivity 

(Blooi et al. 2015a, 2015b), there are no 

practical control measures for freeranging salamanders. 

Response: We agree with the 

comment. In the second interim rule, we 

clarify that, while treatment options 

exist that may help reduce the threat 

posed by Bsal for imported and captiveheld specimens, those options have not 

been standardized and their 

effectiveness remains uncertain for 

large-scale application. 

Pathways 

(PR11) Comment: The pathway by 

which Bsal spreads is unknown, except 

that water is involved. Thus, the States 

should be responsible for implementing 

measures on waterways that prevent the 

introduction of Bsal. 

Response: We agree that the U.S. 

States should be strong partners in 

helping to prevent the introduction, 

establishment, and spread of Bsal. We 

conclude that the main pathway for the 

potential global spread of Bsal is the 

introduction into the United States 

through international trade in 

salamanders, and we are acting with 

this rule to reduce those risks. 

(PR12) Comment: The pathway 

analysis, epidemiology of the disease, 

and investigation of the origins of the 

outbreak need more investigation. In 

addition, no laws or regulations exist to 

control the disposal of untreated water 

from captive salamander enclosures. 

Given the virulence of the disease, how 

did Bsal enter the European 

environment? Was it the result of opensystem housing, such as outdoor pens or 

open-system water flow? Intentional 


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release of pets seems an unlikely source 

since the course of the disease is rapid 

with signs of infection within 8 days. 

Response: While it is possible that 

Bsal can be transmitted through other 

pathways and vectors, the comment 

does not provide any evidence that 

other pathways are more likely than 

international trade. Drawing on the 

evidence cited in this rule in the 

Pathway Analysis, we conclude that the 

primary potential pathway for the entry 

of Bsal into the United States is through 

the international trade in salamanders. 

Our analysis concludes that Bsal can 

survive on infected animals long enough 

for the pathogen to be introduced into 

the environment and transmitted to 

species that are negatively impacted by 

Bsal. 

(PR13) Comment: We suggest another 

pathway that should be addressed is 

that fishes, plants, and invertebrates 

may be co-cultured with newts. It is 

unknown if they can act as a fomite. For 

Bd, there is evidence for foliage and 

invertebrates as substrates. Amphibians 

can enter the United States as 

stowaways on agricultural and other 

imports. For example, the Cuban tree 

frog that invaded Florida hitchhiked in 

shipping crates coming from the 

Caribbean. It is also possible that Bsal 

could be transported in contaminated 

water that is entering the United States 

with imported fish for aquaculture or 

the pet trade. 

Response: We concur with the 

comment. Other pathways are a 

concern; however, the Service 

concludes that the trade pathway in 

salamanders is the most significant 

means by which Bsal could be 

introduced into the United States. The 

final and second interim rules will be 

protective because a co-cultured 

salamander that has also been found to 

be a carrier would be prohibited from 

importation into the United States. The 

Service will also continue to seek 

opportunities to work with partners to 

minimize the risk from other pathways. 

Species Additions 

(PR14) Comment: Some of the Bd 

infections observed in species from 

Ambystoma may have been Bsal. 

California tiger salamanders 

(Ambystoma californiense) can survive 

chytrid infections that would make 

them likely carriers. Another peer 

reviewer stated that the rule states, ‘‘At 

least four [native species] are resistant to 

Bsal infection, of which one is expected 

to be a carrier because Bsal was able to 

invade the skin of that species long 

enough to move or transmit the fungus 

to other salamanders.’’ This is a 

reasonable assumption. The commenter 

makes the same assumption for 

Ambystoma based on their ability to be 

infected by Bd. 

Response: The testing results 

available to the Service at the time of 

the 2016 interim rule provided no 

evidence that some species within 

Ambystoma are carriers of Bsal. 

However, subsequent research provides 

that evidence. Please see IV. Second 

Interim Rule below for that evidence. 

Prohibition on Interstate Transportation 

(PR15) Comment: Enforcing the 

interstate prohibition will be difficult. 

Also, it seems unnecessary if Bsal is not 

known to exist in the United States. 

While it is possible that Bsal may be 

present on a pet in the United States, 

the interstate transportation prohibition 

could prevent movement of that pet. 

Response: As stated above under A. 

Background in III. Final Rule to the 

2016 Interim Rule, the D.C. Circuit 

Court of Appeals held on April 7, 2017, 

that transportation of injurious wildlife 

between the 49 States within the 

continental United States (the 

contiguous 48 States and Alaska) is not 

prohibited by the Lacey Act, unless that 

movement of the wildlife is restricted 

due to conditions associated with issued 

injurious wildlife permits. The language 

in 50 CFR 16.14(a) was and still is 

correct in that it does not prohibit 

interstate transport between States 

within the continental United States. 

Transport of injurious wildlife between 

the enumerated jurisdictions set forth in 

the shipment clause of 18 U.S.C. 42 (the 

continental United States, the District of 

Columbia, Hawaii, the Commonwealth 

of Puerto Rico, and any territory or 

possession of the United States), 

codified in Federal regulations at 50 

CFR 16.3, remains prohibited. The 

Service will continue to seek 

opportunities to encourage affected 

members of the public to take 

responsible actions related to listed 

species. 

Prohibition on Preserved Specimens 

and Parts 

(PR16) Comment: There is little risk of 

Bsal transmission from chemically 

preserved specimens. Even if 

contaminated with Bsal DNA, it is 

unlikely that the chytrid would be 

viable or lead to introduction of Bsal 

into the United States. However, the 

prohibition should be maintained for 

live or frozen specimens, because it is 

unclear whether Bsal can survive 

freezing. Experimental studies are 

needed to elucidate the viability of Bsal 

after preservation and freezing. 

Response: In response to comments 

we received explaining that preserved 

salamanders or their preserved parts 

pose a low risk of transmitting Bsal, the 

Service is removing chemically 

preserved specimens and body parts 

from the injurious wildlife listing as 

long as chemical preservation is 

adequate to render the fungus inviable. 

Frozen specimens remain regulated as 

parts of injurious wildlife as clarified in 

A. 

Species Capable of Carrying Bsal 

(PR17) Comment: The 2016 interim 

rule states in the section ‘‘Invasiveness 

and Transmission of Bsal’’ that the 

rough-skinned newt and the eastern 

newt are capable of carrying Bsal. What 

is the evidence for this? 

Response: Martel et al. (2014) found 

that the eastern newt and rough-skinned 

newt were found to be lethally 

vulnerable to Bsal. Below in E. Pathway 

Analysis, Introduction Pathways, we 

added that Bsal can remain viable inside 

dead host tissue (Martel et al. 2013). We 

have concluded that lethally vulnerable 

salamanders are also carriers. More 

recent information affirms the newts as 

carriers (please see II. Current 

Rulemaking Action below for that 

evidence). 

Invasiveness of Salamanders 

(PR18) Comment: The rule states that 

Oriental fire-bellied newts (Cynops 

orientalis) and paddle-tailed newts 

(Paramesotriton (Pachytriton) labiatus 

or brevipes), which are native to China, 

have been found in the wild near an 

animal importer’s facility in Florida. 

Because they were found outside of the 

facility does not necessarily mean that 

they are a breeding, invasive, 

reproducing population. 

Response: We concur with the 

comment that being found outside of a 

facility does not necessarily mean that 

the species in question is actually 

invasive, although a released 

salamander could persist long enough in 

the ecosystem to transmit Bsal if the 

salamander was exposed to viable 

spores. 

(PR19) Comment: The rule states that 

Picco and Collins (2008) found that 

salamanders sold as bait were highly 

infected with both ranavirus and Bd, 

thereby increasing the likelihood of 

pathogen transmission into new areas of 

the United States through the act of 

fishing. Have declines from this 

pathway been documented? If not, the 

point is rather moot or at least weak. 

Response: That comment refers to the 

section on invasiveness of salamanders. 

The Picco and Collins (2008) reference 

demonstrates that anglers routinely 

release salamanders into the areas 

where they fish, which serves as one 


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pathway for salamanders being 

introduced into the environment, 

including nonnative habitats. This 

pathway may also serve as a vector for 

pathogens, including Bsal. 

(PR20) Comment: The 2016 interim 

rule states that the four salamander 

genera most commonly imported into 

the United States from 2004 to 2014 

were Cynops, Paramesotriton, Triturus, 

and Pachytriton. You should check 

Krysko et al. (2011) against the firebellied newt. 

Response: Krysko et al. (2011) was 

cited by the USGS in its fact sheet for 

the Oriental fire-bellied newt in 

reporting nonindigenous occurrences, 

although none have been reported since 

2010 (USGS Nonindigenous Aquatic 

Species (NAS) 2021 [CYOR]). 

(PR21) Comment: In evaluating the 

potential to eradicate or manage 

established populations, the 2016 

interim rule says that, while some 

introduced salamanders in the United 

States have been successfully 

controlled, others have not. However, 

evidence for control is sparse. There is 

a difference between a small population 

living in exceptional circumstances and 

an invasive species. In many cases, 

small populations of animals will 

persist but not spread. These are not 

invasive and should not be used as 

examples of the removal of invasives. 

Response: Executive Order 13751 

defines an invasive species as a 

nonnative organism whose introduction 

causes or is likely to cause economic or 

environmental harm or harm to human, 

animal, or plant health. Establishment 

and spread can increase the threat that 

a particular species causes, but 

establishment and spread are not in 

themselves mandatory criteria for 

defining a species as invasive or 

injurious. The analysis under Potential 

To Eradicate or Manage Established 

Populations in the 2016 interim rule 

was intended to show that there is a risk 

of harm from the introduction of Bsal 

even if a nonnative salamander 

population could be successfully 

controlled or eradicated in the 

environment. 

(PR22) Comment: The 2016 interim 

rule states that the total risk of Bsal to 

native salamanders is high. It should 

probably take Bsal invasive risk into 

account. 

Response: The total risk to native 

salamanders was based on a USGS risk 

assessment (Richgels et al. 2016). We 

took invasive risk into account in other 

sources. We discussed the issues related 

to invasion risk of Bsal under Likelihood 

of Release or Escape. To make our 

listing determination for salamanders, 

we drew upon the results of multiple 

independent risk assessments and our 

own analysis and found that Bsal is 

likely to be introduced into the United 

States if no additional risk mitigation 

steps were taken by the Service. 

Bsal Infection 

(P23) Comment: The 2016 interim 

rule states that Bsal can also be 

introduced into the environment 

through the improper disposal of 

contaminated water or other materials 

used to transport salamanders and that 

the fungus can likely persist in such 

materials independent of whether a 

salamander is present. Although the 

fungus can persist in the environment, 

it may not be at infectious levels. 

Response: The number of fungal 

spores required to initiate a Bsal 

infection has not been well researched, 

and this number may vary among host 

species and with other disease-related 

factors (environmental and Bsal-specific 

factors). The Service’s analysis was 

based on whether the available evidence 

showed that a given genera was capable 

of carrying Bsal and introducing it into 

U.S. environments. 

(PR24) Comment: The 2016 interim 

rule states that the discharge of 

untreated water used to house infected, 

captive animals could be a pathway for 

releasing infective zoospores into the 

environment and exposing native 

salamanders to Bsal. There is some 

evidence for Bd, such as the 30,000 

zoospores detected after 10 hours in 

DiRenzo et al. (2014), but a more direct 

experiment occurred in Carey et al. 

(2006), where the solutions that had 

housed toadlets being tested were used 

to infect other toads. This is strong 

evidence and should be included. 

Response: We concur with the 

comment and have added the reference 

to Carey et al. 2006 in the second 

interim rule. 

Likelihood of Release or Escape 

(PR25) Comment: An outbreak of Bsal 

in Germany was discovered in a captive 

salamander collection (Sabino-Pinto et 

al. 2015). 

Response: The comment is correctly 

stated, and this point is now referenced 

in the second interim rule. 

Ability To Prevent or Control the Spread 

of Pathogens or Parasites 

(PR26) Comment: The two treatments 

from Blooi et al. 2015(a) should be 

changed from ‘‘in development’’ to 

‘‘available.’’ A sentence explaining that 

this treatment is limited in feasibility 

and applicability (that is, not all 

salamander species can tolerate the 

temperature treatment recommended) 

should be added. 

Response: We concur with the 

commenter’s statements as reflected in 

Blooi et al. 2015(b) and modified the 

second interim rule consistent with the 

comment. 

(PR27) Comment: Control measures 

are available for Bsal-infected 

salamanders, but these would be more 

relevant for captive salamanders rather 

than free-ranging salamanders. 

Response: We have edited the second 

interim rule accordingly by recognizing 

that control measures for Bsal-infected 

salamanders are more relevant for 

captive rather than free-ranging 

salamanders. 

Impacts on Wildlife Resources or 

Ecosystems 

(PR28) Comment: The 2016 interim 

rule states that, ‘‘[i]f rough-skinned 

newts were to experience severe 

declines from Bsal infection, a result 

could be significant additional inputs of 

carbon to the atmosphere as has been 

observed with other species. The 

commenter recommends modifying the 

sentence to read, ‘‘If rough-skinned 

newt populations were to experience 

severe declines from Bsal infection, 

atmospheric inputs of carbon may be 

altered, as has been observed with other 

species (Wyman 1998; Best and Welsh 

2014).’’ 

Response: We have edited the second 

interim rule consistent with the peer 

reviewer’s comment. 

(PR29) Comment: The 2016 interim 

rule noted that scientists and diagnostic 

laboratories are working to standardize 

laboratory protocols. What happens if 

they do? 

Response: Standardized laboratory 

protocols are an important part of 

disease management, but the ability to 

validate, document, and enforce disease 

testing requirements is also necessary. 

Additionally, the capacity to implement 

quarantines and live-animal inspections 

may be required. Publication of the final 

rule does not preclude future regulatory 

action based on emerging science and 

increased capabilities. 

Economic Analysis, Regulatory 

Flexibility Analysis, and Effect on 

Industry 

(PR30) Comment: Alternative 3 was 

preferred over Alternative 4 in the draft 

economic analysis. It was not clear 

whether salamanders were excluded 

from Alternative 3 because they were 

not tested or whether all of the ones 

tested showed no infection. If they 

simply were not tested, Alternative 4 

seems like the more responsible option 

given a precautionary approach since 

many salamander genera appear to be at 

risk and given that the difference in cost 


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between Alternatives 3 and 4 seemed 

relatively small ($3.8 million versus $4 

million). Moreover, untested genera may 

become substitutes when the genera 

under Alternative 3 are no longer 

available, which remains a problem if it 

is reasonable to expect some risks 

associated with the untested salamander 

genera. 

Response: Under Alternative 3, we 

listed genera for which we have 

evidence that at least one species in a 

genus is a carrier of Bsal with no 

conclusive countervailing evidence that 

other species in that genus are not 

carriers. Alternative 3 does not include 

species from genera that have not been 

tested for Bsal vulnerability. Alternative 

4 would include the listing of all 

salamander species. The expected 

increase in cost from Alternative 3 to 

Alternative 4 was not considered in our 

determination about the injuriousness of 

the species because the determination is 

based on defensible scientific evidence. 

The Service determined that there was 

unknown risk from genera where no 

species have yet been tested for Bsal 

and, therefore, could not list those 

genera at this time. 

(PR31) Comment: It was not clear in 

Alternative 5 whether there would be 

administrative costs associated with 

health certificates and whether there is 

a probability of making a mistake. 

Response: While most of the testing 

costs of administering a healthcertificate program may not fall on the 

government or public, there would still 

be costs to the Service involved in staff 

time to set up the program and oversee 

it, as well as wildlife inspectors 

checking import shipments for the 

additional requirements. The draft 

economic analysis lumps the 

administrative costs with costs of 

testing, and both are mentioned as a 

concern in sections 4.1.1.5 and 5.0. A 

health-certificate program was not our 

preferred alternative for a nationwide 

regulatory program by the Service at this 

time because of uncertainties with its 

effectiveness, including the 

effectiveness and sensitivity of current 

testing methods (including the return of 

false negatives); lack of validation and 

sufficient testing capacity; lack of 

standardized treatment methods; and 

lack of agency resources required to 

conduct inspections, interpret results, 

and issue health certificates. 

(PR32) Comment: Has inflation been 

taken into account in the analyses of 

economic costs to adjust costs of today’s 

dollar values? 

Response: All prices in the draft 

economic analysis were updated for the 

final economic analysis to the 2021 

price level Consumer Price Index for All 

Urban Consumers that was used for all 

indexing purposes (see section 3.1.2). 

Salamander retail price data was 

received in 2015 dollars during the 

course of the study. Tables labeled as 

2021$, or 2015$, have either been 

adjusted for inflation or did not need 

adjustment. The original price level is 

the year for the citation unless 

otherwise noted. Tables without a price 

level or data origin year have been 

amended. 

(PR33) Comment: The economic costs 

appear to reflect the maximum costs 

since it does not appear that alternative 

sales were considered. For example, if 

buyers cannot buy salamanders, would 

they buy other amphibians instead or 

would they simply buy nothing? Only 

the latter would result in the estimated 

costs. Similarly, the market for ‘‘local’’ 

salamanders may increase as a response. 

Response: Section 3.1.2 of the 

analysis of economic costs explains 

three points. Estimated importation 

losses are stated as maximums due to 

the unknown effect on domestic 

breeding and consumer substitution. 

Domestic losses are also estimated at the 

maximum (loss of entire industry) due 

to the lack of data on transport between 

the enumerated jurisdictions in the 

shipment clause of 18 U.S.C. 42 

(codified in Federal regulations at 50 

CFR 16.3). We also added detail to this 

section to clarify why the losses may 

range from zero to the maximums stated 

in each alternative analysis section later 

in the report. 

(PR34) Comment: The pet industry 

will not be altered significantly by this 

rule, but Bsal would likely impact wild 

populations of salamanders. Perhaps 

with the exception of breeders, pet sales 

would probably shift to another animal 

with little loss of revenue. 

Response: Exit from an industry or 

substituting a legal product is 

dependent on multiple factors as 

discussed in the report beginning in 

section 3.1.2 of the economic analysis. 

We added details and clarification to 

this section in the final economic 

analysis. Substitution can occur with 

other salamander species, an animal 

from another order, or another category 

of goods altogether. 

(PR35) Comment: The draft economic 

analysis states under ES 1.1 Economic 

Analysis, ‘‘In addition, we used data 

from IMPLAN® (Minnesota IMPLAN 

Group, 2013) to estimate the direct 

effects of this rulemaking.’’ MIG 

changed their name. They now go by 

IMPLAN Group LLC. In addition, what 

data year did you use? 

Response: We changed the name to 

‘‘IMPLAN Group LLC’’ in the final 

economic analysis. We used study area 

data from 2013 for the economic 

analysis. 

(PR36) Comment: Regarding the draft 

economic analysis under ES 1.4.5 

Alternative 5, does the cost estimate of 

the loss of revenue to companies or 

individuals involved in the importation 

or interstate movement of any 

salamander species consider the cost of 

health-certificate examinations and 

inspections by veterinarians affiliated 

with the Department of Defense, zoos, 

and industry as well as private 

practitioners? 

Response: Due to data limitations, the 

final economic analysis does not 

explicitly estimate the cost of health 

certificates. The analysis assumes that 

any additional costs for examinations 

and inspections would be absorbed by 

the importer or passed on to the 

consumer, but data limitations restrict 

the analysis from estimating whether 

the importer or consumer would pay. 

Thus, we assume the estimated losses 

for all alternatives including Alternative 

5 is the average sales price of a 

salamander. On average, we assume the 

estimated maximum loss (sales price) 

would include all testing costs. 

(PR37) Comment: The commenter 

believes a job in IMPLAN is annualized. 

IMPLAN’s definition is ‘‘A job in 

IMPLAN = the annual average of 

monthly jobs in that industry.’’ 

Response: We have changed the final 

economic analysis to reflect the above 

definition. 

(PR38) Comment: The draft economic 

analysis, at ES 1.6 Conclusion, states 

that it is unclear how much testing, 

treatment, and the health certification 

processes would cost. 

Response: It is correct that these costs 

are unknown and could not be 

estimated unless a compliance method 

is developed. 

(PR39) Comment: The commenter 

found locating the tables and figures in 

the draft economic analysis to be 

challenging. For example, figure 1 is not 

shown for several pages after first being 

noted in 2.2 Salamander Market. 

Response: Placement of tables and 

figures was determined by the 

progression of the analysis. As many 

numbers are referred to many times, 

they will not always be near all 

discussions. Table and figure numbers 

are given to allow the reader to find 

them. 

(PR40) Comment: The draft economic 

analysis states in Table 3–Pet Stores 

Industry that the annual payroll for all 

is less than the annual payroll for small 

business. That does not seem right. 

Response: We corrected the table in 

the final economic analysis. 


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(PR41) Comment: The draft economic 

analysis states in 2.3.5 U.S. Bred 

Salamanders, ‘‘Domestically bred 

salamanders would represent less than 

one percent of the United States 

salamander sales between 2012 through 

2014 if this data depicts the entire 

domestic supply.’’ This is confusing 

because table 10 states that 76 percent 

of commercial salamanders are U.S. 

bred. 

Response: The 1 percent refers to all 

salamanders, whereas the 76 percent 

refers to the species and genera listed in 

table 10. We amended table 10 in the 

final economic analysis to clarify this 

point. 

(PR42) Comment: The draft economic 

analysis states under 3.1.1 Analysis of 

Economic Benefits, ‘‘Fewer outdoor 

recreationists could lead to a decrease in 

expenditures; to demonstrate we use 

$25,000. Implementing a fictional 

alternative, Alternative Y would reduce 

the probability of Bsal establishment to 

10 percent from 80 percent. The 

expected costs in the current situation 

would be $20,000 ($25,000 × 0.8); with 

Alternative Y, the expected costs would 

be $2,500 ($25,000 × 0.1). Net avoided 

costs would be $17,500 

($20,000¥$2,500), one measure of the 

benefits of Alternative Y.’’ The 

commenter finds this example to be 

confusing and suggests omitting. 

Response: We deleted the example in 

the final economic analysis. 

(PR43) Comment: In the Executive 

Summary of the draft economic 

analysis, you describe some of the 

potential costs of the regulation (for 

example, lost consumer surplus for pet 

owners). It seems in the discussion 

under 3.1.2 Analysis of Economic Costs 

that the costs in the cost/benefit sense 

are being conflated with lost revenue 

and the economic analysis. This is okay, 

but this section could be more clear. 

Response: The Executive Summary 

indicates that consumer surplus cannot 

be estimated under the scope of this 

report and that an alternate 

methodology will be used. Sections 

3.1.1–3.1.2 explain how the analysis 

uses the maximum sales as a proxy for 

the direct economic losses. No 

economic benefits are evaluated for the 

existence of a species in this report. 

(PR44) Comment: The draft economic 

analysis states under 3.1.2.2 Small 

Business, ‘‘Estimates using the unique 

importers (average of 5 a year), or one 

breeder, yield the maximum adverse 

impacts; no fewer entities would be 

impacted under the status quo. 

Applying these two methods brackets 

the impacts on importers and pet 

stores.’’ It is unclear what ‘‘average of 5 

a year’’ means. 

Response: We changed ‘‘average of 5 

a year’’ to ‘‘annual average’’ in the final 

economic analysis. The analysis was 

also updated to an annual average of six 

importers. 

(PR45) Comment: In the draft 

economic analysis, the numbers in the 

sectors columns of tables 12–14 do not 

seem to correspond to anything. Could 

this column be omitted? 

Response: The columns were deleted 

in the tables. 

Public Comments and Our Responses 

We reviewed all 280 comments we 

received during the public comment 

period for the 2016 interim rule (81 FR 

1534, January 13, 2016). We received 

comments from Federal agencies, State 

agencies, commercial and trade 

organizations, conservation 

organizations, nongovernmental 

organizations, and private citizens. The 

comments provided a range of views on 

the proposed listing as follows: (1) 

Unequivocal support for the listing with 

no additional information included; (2) 

unequivocal support for the listing with 

additional information provided; (3) 

equivocal support for the listing with or 

without additional information 

included; (4) unequivocal opposition to 

the listing with no additional 

information included; and (5) 

unequivocal opposition to the listing 

with additional information included. 

While all comments were reviewed 

and considered, several comments did 

not contain information that was new 

compared to other comments or 

included substantial information that 

required analysis. Comments included 

individual ideas, data, 

recommendations, or suggestions on the 

interim listing and the draft economic 

analysis. Some commenters addressed 

the 14 questions we posed in the 2016 

interim rule. We consolidated 

comments and responses into key issues 

in this section. We edited some 

comments for brevity or grammar while 

maintaining the intent. We combined 

comments that expressed similar 

perspectives. 

Use of Scientific and Common Names 

(1) Comment: The Service asked, for 

the species being listed in the 2016 

interim rule, if the scientific and 

common names are the most 

appropriate ones accepted by the 

scientific community. Most of the 

herpetological community uses the 

Society for the Study of Amphibians 

and Reptiles joint societies-endorsed list 

(Crother 2012); both the Association of 

Fish and Wildlife Agencies (AFWA) and 

Partners in Amphibian and Reptile 

Conservation (PARC) use this 

nomenclature in our formal 

publications. However, some States use 

other nomenclature, while some others 

use older nomenclature simply due to 

the inability to update frequently. 

Response: The comment identifies the 

disparate use of scientific and common 

names used among herpetological and 

management entities. We believe this 

approach supports our decision to 

remove the enumerated list of species 

within each genus in 50 CFR 16.14 for 

the second interim rule. Each species 

within each genus will therefore be 

included as injurious wildlife in the list 

of injurious amphibians. 

(2) Comment: There are quite a few 

errors (some species listed twice under 

different Latin names) in the proposal. 

Response: The commenter did not 

provide specific examples, so we cannot 

check this comment with additional 

references. The comment does support, 

however, our decision to remove the 

enumerated list of species within each 

genus in 50 CFR 16.14 for the second 

interim rule. 

Listing of Preserved Specimens, Parts, 

and Eggs and Gametes 

(3) Comment: Scientific specimens of 

salamanders that are desiccated or have 

been fixed or preserved in formalin or 

alcohol should be exempt from this rule 

because Bsal is no longer viable. 

Response: We concur that preserved 

specimens do not pose a risk for 

pathogen transmission as long as 

chemical preservation is adequate to kill 

Bsal, and we have removed chemically 

preserved specimens from the reach of 

this final rule. 

(4) Comment: What is included in the 

definition of ‘‘parts of salamanders’’ and 

why? Listing swabs makes testing for 

disease more difficult, which could 

adversely affect the intended effect of 

the rule. Please provide an exemption 

for tissue samples (including 

histological samples), molecular 

extractions, swabs, and other parts. 

Response: Any item that contains 

cells or genetic material from a listed 

species is considered a ‘‘part’’ of the 

listed animal. This definition is not 

unique to the salamander rule but is 

consistent with standard regulatory 

definitions used by the Service. 

Specimens, such as skin swabs and 

tissue samples for microscopic analysis 

(histology), are included as ‘‘parts’’ in 

the rule consistent with the definition of 

‘‘fish or wildlife’’ outlined in 50 CFR 

10.12, which includes ‘‘any part, 

product, egg, or offspring thereof.’’ Also, 

50 CFR 10.12 states that ‘‘amphibians’’ 

means a member of the class, Amphibia, 

including, but not limited to, frogs, 

toads, and salamanders; including any 


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part, product, egg, or offspring thereof, 

or the dead body or parts thereof 

(excluding fossils), whether or not 

included in a manufactured product or 

in a processed food product. Specimens 

such as swabs intended for culture or in 

transport or growth media will require 

permits. We may issue permits to 

facilitate all of the above-described 

activities. For purposes of this rule, eggs 

and gametes and purified extracted 

genetic material of salamanders are 

excluded from the prohibitions as 

‘‘parts’’ because they are unable to cause 

pathogen transmission. However, swabs 

and histological samples that are 

preserved or fixed in appropriate 

concentrations of ethanol or 

formaldehyde-based solutions are also 

not injurious as long as chemical 

preservation is adequate to kill Bsal as 

described in current peer-reviewed 

literature. The appropriate 

concentration and minimum exposure 

time for a given chemical preservative 

or fixative to render any Bsal organisms 

non-viable varies with the precise 

chemical formulation and should be 

utilized as described in association with 

such actions in the peer-reviewed 

literature. Please also refer to IV. Second 

Interim Rule. 

Purpose of Listing as Injurious 

(5) Comment: Several comments 

provided feedback on whether eggs and 

gametes should be included in this rule. 

As a comment noted, specimens require 

transport with some form of medium, 

such as water or plant materials, to 

remain viable, and that medium could 

harbor Bsal, thus constituting a threat by 

indirectly moving disease vectors with 

the eggs or gametes and increasing the 

risk of indirect Bsal transmission. 

Further, eggs at certain stages of 

development could contain keratinized 

tissues (for example, Xie and Yu (1992)), 

which could transmit the Bsal pathogen. 

However, other comments noted that if 

entire genera are excluded from the 

listing because they cannot be infected, 

then the relative risk from the transport 

of eggs is no greater. 

Response: Our authority does not 

include the listing of the medium, such 

as water or plant materials, that the 

specimens are transported in. As noted 

in this rule, there is no evidence that 

salamander reproductive material also 

contains keratin that might harbor Bsal. 

Therefore, eggs and gametes are not 

listed by this rule. 

Effect of Rule on Scientific Research 

(6) Comment: The rule will have a 

negative impact on scientific research, 

especially on native taxa. The 

prohibition should not apply to 

scientific research, providing that the 

biologist in question is in possession of 

an approved permit from the State 

where the specimen(s) were originally 

collected. 

Response: Permits from the Service 

for injurious listed species can be 

obtained for scientific, zoological, 

educational, and medical use for 

importation, shipment between the 

enumerated jurisdictions in 18 U.S.C. 

42(a)(1) (codified in Federal regulations 

in 50 CFR 16.3), and transport for a 

previously permitted salamander. The 

statute does not cover collection of 

native species or transport of injurious 

listed salamanders across State lines 

within the continental United States 

(see PR15). 

Species Not on the List 

(7) Comment: Several commenters 

advocated for adding various genera or 

listing at the family level, such as 

Salamandridae, while others advocated 

for listing all species. 

Response: The salamander species 

listed by this final rule and the second 

interim rule are those found within a 

genus for which we have confirmation 

that at least one species in that genus is 

a carrier of Bsal, and there is no 

conclusive countervailing evidence 

suggesting that some species within the 

genus are not carriers. Although 

additional salamander species could be 

at risk from Bsal infection or could serve 

as a carrier, we are not listing species in 

those genera because they had not yet 

been tested. We considered listing more 

species based on the comments we 

received. However, the logic we used for 

listing at the genus level breaks down at 

the family level for one family. In the 

family Plethodontidae, the genus 

Gyrinophilus is not known to be a 

carrier, but the genera Hydromantes and 

Plethodon are carriers. As a result, we 

cannot list all species within 

Plethodontidae. We also cannot list a 

species without science-based 

documentation. We can list for the 

statutorily defined purposes under the 

statute codified at 18 U.S.C. 42(a); any 

other purpose is beyond the scope of 

this rulemaking. Please see IV. Second 

Interim Rule below for additional genera 

we have documented as injurious and 

are therefore listing. 

(8) Comment: Some comments noted 

that while some salamander species 

appeared to be resistant to Bsal in 

infection experiments, it is unclear how 

strong this resistance will be outside of 

the optimal husbandry conditions found 

in laboratory settings. 

Response: As part of the justification 

for listing, the Service acknowledges 

that salamander species known to be 

tolerant of Bsal infection under 

experimental conditions may 

demonstrate more severe clinical 

disease when infection is combined 

with additional stressors in the wild, as 

has been found for other diseases, 

including those in amphibians (Wobeser 

2007; Kerby et al. 2011; Kiesecker 2011). 

However, the Service needed evidence 

that a species was a carrier or likely to 

be a carrier before listing the genus as 

injurious. 

(9) Comment: Tylototriton podichthys 

was recently described and should be 

added to the list (Phimmachak et al. 

2015). 

Response: All species in a genus are 

also listed as injurious even if they are 

not specifically identified in the rule. 

Because we identified Tylototriton as 

one of the genera listed in the 2016 

interim rule and hereby affirmed, T. 

podichthys is one of the species listed 

as injurious. The comment supports our 

decision to remove the enumerated list 

of species within each genus in 50 CFR 

16.14 for the second interim rule. 

(10) Comment: The Service should 

establish an expedited process by which 

additional salamander species can be 

added to the list as new information 

becomes available. 

Response: Rulemaking under 18 

U.S.C. 42 is governed by the APA, under 

which we promulgated the 2016 interim 

rule and this final rule. The Service is 

adding new genera to the list with the 

second interim rule in this document. 

Species Should Be Removed From the 

List 

(11) Comment: Species from the 

genera Cynops, Salamandra, 

Pleurodeles, Siren, Notophthalmus, and 

Triturus should be removed. They are 

the most commonly kept species and 

listing will significantly affect those 

who raise, study, or keep animals from 

these species. 

Response: Due to shared 

characteristics by species within a 

genus, other species within these genera 

are also likely to be carriers of Bsal. The 

Service found that species from the 

genera Cynops, Salamandra, 

Pleurodeles, Siren, Notophthalmus, and 

Triturus can carry Bsal and, therefore, 

pose a substantive risk to native 

salamander populations. The listing of 

these species as injurious wildlife does 

not regulate possession, transport, 

breeding, or sale within the continental 

United States unless regulated under 

permit. Other Federal, State, Tribal, or 

Territorial laws may apply. 

(12) Comment: No native species 

should be listed. Listing native species 

as injurious wildlife solely on the basis 

of their vulnerability or capacity to carry 


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an absent foreign pathogen is 

concerning. Additionally, most of the 

animals tested that were lethally 

vulnerable were dead within about a 

month, as per Martel et al. (2014), and 

the odds of any of these animals being 

available for sale while carrying the 

disease are almost nonexistent. 

Response: We listed native species in 

the 2016 interim rule partly because 

some native species that we concluded 

can be carriers of Bsal are raised outside 

the United States and imported into the 

country and partly because listing 

would prohibit transport of injurious 

salamanders between the enumerated 

jurisdictions in the shipment clause of 

18 U.S.C. 42 (codified in Federal 

regulations at 50 CFR 16.3), in order to 

prevent introduction, establishment, 

and spread of the pathogen in U.S. 

ecosystems. Several native species of 

newts were already known to be highly 

susceptible to dying from Bsal. Not all 

species die immediately upon exposure 

to Bsal, and there is no evidence that 

lethally vulnerable species cannot 

survive long enough for Bsal to be 

transmitted within the United States if 

they are infected prior to their 

movement. At the time of the drafting of 

the 2016 interim rule, most of the 

research was being conducted on Asian 

and European species to find out where 

the fungus may have originated and 

why wild European salamanders were 

dying. After the 2016 interim rule 

published, many studies by U.S. 

researchers began to provide 

information for the conservation of 

native species in the event Bsal is 

introduced into the American 

environment. These studies demonstrate 

that many native salamanders are 

susceptible and can be Bsal carriers. 

(13) Comment: Many of the listed 

species in some genera, such as 

Plethodon, Taricha, and 

Notophthalmus, have never been found 

to carry Bsal. These species should be 

delisted. 

Response: New information confirms 

that species from the genera Plethodon, 

Taricha, and Notophthalmus can carry 

Bsal based on laboratory studies. As of 

the drafting of the second interim rule, 

all three species of Notophthalmus have 

been found to be lethally susceptible to 

Bsal (Gray et al. 2023), and two of the 

four Taricha species are carriers (Gray et 

al. 2023). 

(14) Comment: The listing of the 

entire genus Plethodon is based on the 

Martel et al. (2014) study from a sample 

of two wild-caught P. glutinous 

imported to Europe. Under the 

circumstances, the evidence suggests 

that all species in the genus Plethodon 

should be removed from the list. 

Response: We disagree with the 

comment. While Martel et al. (2014) 

classified the slimy salamander 

(Plethodon glutinous) as resistant to 

infection, the study also demonstrated 

by histology that Bsal could invade the 

skin of the slimy salamander, even 

though it apparently cleared the 

infection and did not show signs of 

clinical disease. Our examination of the 

supplementary data of Martel et al. 

(2014), including histology (microscopy) 

tests and subsequent discussions with 

the authors, indicates that there is 

sufficient evidence that Bsal was able to 

invade the skin of this species long 

enough to move or transmit the 

infection to other salamanders (Martel et 

al. 2014; A. Martel, University of Ghent, 

pers. comm. 2015; K. Lips, University of 

Maryland, pers. comm. 2015). Because 

we expect all species within a genus to 

respond in a similar way for Bsal carrier 

status, we conclude that all species of 

Plethodon are potential carriers. Since 

the 2016 interim rule published, 

additional studies have shown multiple 

species in the genus Plethodon can be 

carriers (DiRenzo et al. 2021); see IV. 

Second Interim Rule. 

(15) Comment: Some species from the 

genus Neurergus have been bred over 

many generations and are in private 

collections (N. crocatus, N. kaiseri, and 

N. strauchii). N. kaiseri, which is listed 

under the Convention on International 

Trade in Endangered Species of Wild 

Fauna and Flora (CITES) Appendix I, 

has not been imported for years, and 

most of the animals in the United States 

are descendants of zoo colonies and 

hobbyist captive-bred animals. Because 

they are protected by other laws and not 

imported, they do not have any risk of 

transmitting Bsal and there is no need 

to list them. 

Response: Regardless of protection 

level under other laws, these species are 

still injurious wildlife under 50 CFR 

16.14 as part of the genus Neurergus. 

Their protection level under these other 

laws does not change the characteristics 

of the species that we find to be 

injurious to wildlife and the wildlife 

resources of the United States by reason 

of their potential to serve as vectors for 

the pathogen Bsal. Also, there is no way 

to confirm that captive-bred 

salamanders have not been exposed to 

Bsal through contact with other 

individuals. Neurergus has been 

confirmed to carry Bsal in a European 

collection (Fitzpatrick et al. 2018), and 

there is a chance a co-housed 

salamander of a different species could 

be imported into the United States. 

Need for Rule 

(16) Comment: The rule is 

unnecessary. The prohibition can be 

justified only if Bsal is found to be 

present in the United States. Other 

commenters stated that the rule is 

unnecessary because Bsal must be here 

already given the number of 

salamanders imported annually and 

their belief in the low likelihood of a 

captive salamander coming into contact 

with the wild populations and 

transmitting Bsal. 

Response: Since the publication of the 

2016 interim rule, Waddle et al. (2020) 

conducted a large-scale surveillance for 

Bsal across 594 counties in 35 States 

and 1 site in Mexico with 11,189 swab 

samples of wild salamanders and some 

frogs and toads, with no positive results 

for Bsal. The purpose of listing these 

species as injurious wildlife is to 

prevent the introduction of the Bsal 

fungus in the wild in the United States. 

A species does not need to be already 

present in trade or in the environment 

to be listed as injurious wildlife. In fact, 

it is often difficult to achieve a 

prevention outcome once a species or 

pathogen occurs in the environment. To 

make the listing determination for 

salamanders, we drew upon the results 

of multiple independent risk 

assessments and our own analysis and 

found that Bsal is likely to be 

introduced into the United States if no 

additional risk mitigation steps are 

taken. Additional discussion on this 

topic can be found in IV. Second Interim 

Rule under the section Likelihood of 

Release or Escape. 

(17) Comment: To list a native species 

of wildlife as injurious simply because 

it may act as a host to a rare but 

potentially devastating pathogen that 

has not been detected in the United 

States is an unmanageable proposition. 

Every native species of wildlife fits this 

criterion and would need to be listed as 

injurious for some rare pathogen 

detected in a very isolated outbreak on 

another continent, as has occurred with 

Bsal. 

Response: The purpose of listing these 

species as injurious wildlife is to act 

preemptively to prevent the 

introduction, establishment, and spread 

of the Bsal fungus in the wild in the 

United States. The fungus affects many 

native salamanders, with lethal effects 

on many salamander species, and it is 

not yet known to be found in the wild 

in the United States. There is an existing 

pathway for the fungus to arrive by 

importation of salamanders, including 

species native to the United States that 

are raised in captivity outside of the 

United States and then imported back 


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into the continental United States and 

the enumerated jurisdictions of the 

shipment clause. This regulatory action 

is being taken to prevent Bsal’s arrival 

through the organisms-in-trade 

pathway. If we wait until the fungus 

arrives, it will likely be impossible to 

eradicate. We will continue to evaluate 

other species for possible risks and 

consider injurious wildlife actions as 

appropriate and authorized under 18 

U.S.C. 42(a). 

(18) Comment: Bsal can be treated and 

cured in captivity, so there is no reason 

to limit availability of the species in 

question. 

Response: Voluntary actions, such as 

applying heat therapy as described in 

Blooi et al. (2015a) and Blooi et al. 

(2015b), may help reduce the threat 

posed by Bsal for specimens held in 

captivity. However, at this time it is not 

possible to determine the likelihood of 

success of such measures for all species 

or of achieving compliance with 

prophylactic treatment or treatment 

following the onset of symptoms. 

Therefore, it is unknown how effective 

treatment will be in preventing Bsal’s 

introduction, establishment, and spread 

in the United States, and no Bsal control 

is known for salamanders in the wild. 

(19) Comment: If a species that is not 

a carrier is similar in appearance with 

another species, neither species should 

be removed from the list unless both 

species are confirmed that they are not 

susceptible to or carriers of Bsal. 

Response: The Service does not have 

the authority under the statute to list a 

species based solely on its similarity of 

appearance to another species. We list 

based on our determination of 

injuriousness. 

(20) Comment: There is no case in the 

United States where salamanders, native 

or nonnative, have been proven as 

invasive or injurious. The 2016 interim 

rule does not substantiate injury by 

transplanted or exotic salamanders. 

Response: The salamanders are listed 

because they are carriers of a fungus that 

makes them harmful to other 

salamanders, not because the 

salamanders are invasive. We concluded 

that even if the salamander species 

listed by this rule do not become 

established, some species capable of 

carrying Bsal and listed by this rule can 

survive long enough in the wild to 

transmit Bsal. Our findings are 

discussed in Potential To Survive, 

Become Established, and Spread in IV. 

Second Interim Rule. 

Listing Purpose Is To Regulate Disease 

or Manage Native Species 

(21) Comment: Listing salamanders as 

injurious is not an appropriate means to 

regulate an animal disease. The 

injurious wildlife provisions of 18 

U.S.C. 42 pertain to animals and not 

diseases or pathogens. The focus of the 

2016 interim rule is Bsal, a fungus that 

the Service possesses no authority to 

regulate as acknowledged in the interim 

rule. The law provides no provisions for 

testing, surveillance, or certification of 

health to allow for movement in trade. 

Response: As we described in Listing 

Species That Carry Pathogens in the 

2016 interim rule, the Service can list as 

injurious any member of the 

enumerated taxa that are hosts to or 

carriers of pathogens that cause the host 

or carrier to be harmful by its presence 

to one or more of the interests listed in 

the statute. We have previously listed 

species that serve as hosts to or carriers 

of pathogens, as in the case of fishes in 

the salmon family (Salmonidae) (32 FR 

20655, December 21, 1967). We noted in 

the 2016 interim rule that there are 

concerns regarding the effectiveness and 

sensitivity of current testing methods 

(including the return of false negatives), 

lack of validation and sufficient testing 

capacity, and agency resources required 

to conduct inspections, interpret results, 

and issue health certificates. If these 

issues are resolved, it may be possible 

to establish a health certificate for 

salamanders that are free of Bsal. A 

health certificate was established for 

import of salmon under the authority of 

18 U.S.C. 42. While the concerns 

remain, and therefore a Bsal health 

certification has not been established, 

this does not mean that there is no 

authority to establish a health 

certification if circumstances were to 

change. Appropriate conditions may 

also be included in injurious wildlife 

permits under the authority of and 

consistent with the purposes of 18 

U.S.C. 42. 

(22) Comment: Several commenters 

noted that, by definition, ‘‘pathogens’’ 

are injurious and are regulated under 

the authority of other agencies. The 

World Trade Organization and the 

United States Department of Agriculture 

(USDA) recognize the World 

Organisation for Animal Health 

[WOAH, formerly OIE] as the proper 

body to set animal health standards. The 

WOAH helps develop and revise 

international standards for the safe trade 

of animals and animal products. The 

proper course to prevent the 

importation of salamanders carrying 

Bsal is to list the pathogen as a WOAH 

reportable disease, and instead of the 

2016 interim rule, there should be a 

cooperative effort to respond to the 

disease threat as provided through the 

WOAH, World Trade Organization, and 

the National Aquaculture Health Plan 

and Standards (formerly called the 

National Aquatic Animal Health Plan) 

for the United States. 

Response: The USDA and the Centers 

for Disease Control and Prevention have 

authority to regulate wildlife pathogens 

when those pathogens pose a risk to 

agriculture or human health, 

respectively. No such effects are 

currently known in the case of Bsal. The 

Service has authority to regulate the 

importation of certain species that pose 

a risk to wildlife and the wildlife 

resources of our country. This authority 

has been applied in the present case in 

response to a clear and immediate risk. 

After the 2016 interim rule took effect, 

the WOAH did add Bsal as a reportable 

disease, but that action does not prevent 

importation. We work through such 

mechanisms as those provided by the 

WOAH and National Aquaculture 

Health Plan and Standards, and we 

support all efforts by the international 

community to participate in the global 

response to this pathogen. The Service, 

operating within its relevant regulatory 

authority to list injurious wildlife, took 

action through the 2016 interim rule 

due to the urgent need required to 

manage the threat Bsal poses to 

salamanders in the United States. 

(23) Comment: This salamander rule 

not only prevents safe commerce, it 

eliminates any incentive for industry to 

pursue research into the detection and 

treatment of Bsal. Other comments 

expressed similar issues and asked 

whether it would be possible to make 

testing mandatory to allow interstate 

movement. 

Response: While the Service 

acknowledges that some economic 

incentive may have been removed due 

to the prohibitions imposed by the 

injurious wildlife provisions of the 

Lacey Act as a result of listing species 

of salamanders as injurious wildlife 

under this rule, many salamander 

genera were not listed due to 

insufficient evidence at the time as 

carriers, and they remain a possible 

threat. Furthermore, research for 

detection and treatment of Bsal has 

increased considerably in the United 

States since the rule took effect. Permits 

allowing importation can be obtained 

for zoological, educational, medical, and 

scientific use. This final rule explains 

that interstate transportation between 

States within the continental United 

States is not prohibited as of 2017; 

however, the injurious wildlife listing 

still prohibits import into the United 

States, and transport of injurious 

wildlife between the enumerated 

jurisdictions in the shipment clause of 

18 U.S.C. 42(a)(1) (the continental 

United States, the District of Columbia, 


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Hawaii, the Commonwealth of Puerto 

Rico, and any territory or possession of 

the United States), codified in Federal 

regulations at 50 CFR 16.3. 

(24) Comment: The Lacey Act does 

not provide authority to list native 

species. The Lacey Act has been 

examined and critiqued over the last 

few decades within a variety of peerreviewed and gray-literature 

publications (Dentler 1993, U.S. 

Congress 1993, Anderson 1995, Whalen 

1998, Biber 1999, Jenkins et al. 2007, 

Alexander 2013). In no instance did 

these authors construe Lacey Act 

provisions to allow the listing of native 

animals as injurious. Notably, the U.S. 

Congress, Office of Technology 

Assessment, recommended in 1993: 

Congress could provide the Service with 

increased guidance on the purpose of 

this [injurious] list and the specific 

criteria for adding species to it. 

Response: The provisions of 18 U.S.C. 

42(a)(1) do not limit wildlife subject to 

the law to species not native to the 

United States. Under the law, the 

Service may list species that are 

indigenous to the United States if they 

cause injury to the interests enumerated 

in the law. The publications mentioned 

reflect the interpretations of the authors. 

Congress has also listed native species 

as injurious by statute, such as the 

Mariana fruit bat (Pteropus mariannus), 

further demonstrating that the authority 

of 18 U.S.C. 42 is not limited to 

nonnative wildlife. 

Additional Science and Data for Rule 

(25) Comment: In the 2016 interim 

rule, the Service asked what species 

listed as threatened or endangered by 

one or more States would be affected by 

the introduction of Bsal. AFWA and 

several States indicated that several 

salamander species are of interest to 

them, though it is not yet evident how 

Bsal would affect all of these species. A 

number of State threatened and 

endangered or protected species 

(restricted or prohibited from take, 

possession, sale, or other activities) 

were provided during the publiccomment period. 

Response: We appreciate the 

additional information on State 

threatened and endangered species. 

While the Service concluded that some 

species identified by the States are not 

carriers, others are, such as species in 

the genus Plethodon. The carrier status 

of several species, at the time the 

public-comment period closed, had not 

yet been identified. However, more have 

been identified since then, including 

affirming the genus Plethodon in this 

final rule. This additional information 

helps provide additional justification for 

listing species that are capable of 

carrying Bsal, as Bsal presents a risk to 

wildlife and wildlife resources of the 

United States, including those identified 

by the States as in need of protection. 

Pathways and Spread 

(26) Comment: The Service asked the 

question, ‘‘Are there other pathways for 

Bsal into the United States that we 

should address? If so, what are they?’’ 

According to AFWA, a pathway of 

concern that appears to have little or no 

Federal regulatory authority or 

enforcement pertains to biological 

supply companies. Others include 

internet sales involving small shipments 

using couriers such as FedEx or UPS, 

traditional medicine or foreign food 

markets, and ceremonial uses of these 

species. AFWA is aware of interstate 

shipments of some salamanders, though 

not necessarily the currently included 

species, for the purposes of the bait 

trade, but AFWA would like to see some 

exploration of whether there are imports 

for this purpose. 

Another comment noted that, while 

the pet trade is an important pathway, 

salamanders may stow away in nursery 

stock, as was observed with 

northwestern salamanders (Ambystoma 

gracile) in Christmas trees (Rochford et 

al. 2015). In addition to terrestrial 

nursery stock, the aquatic plant and 

animal trade may also spread Bsal in 

shipment water. 

Response: The Service’s pathway 

analysis found that the main pathway 

for the global spread of Bsal is the 

international trade in salamanders, such 

as Martel et al. (2014) noted. While not 

explicitly discussed, that international 

trade could include the uses noted in 

the comment, whether intentional or as 

a hitchhiker. Biological supply and bait 

companies are commercial entities. 

These companies have always had to 

comply with import and export 

regulations under 50 CFR part 14. With 

this injurious listing, these commercial 

businesses will be subject to the same 

prohibitions as other entities. Likewise, 

animals unintentionally imported or 

transported between the enumerated 

jurisdictions in the shipment clause of 

18 U.S.C. 42(a) (also set forth at 50 CFR 

16.3) through nursery stock or other 

pathways would also be in violation of 

the injurious wildlife listing’s 

prohibitions. As explained in A. 

Background in Final Rule to the 2016 

Interim Rule, interstate transport 

between States within the continental 

United States is not prohibited by the 

current prohibitions of 18 U.S.C. 42(a). 

(27) Comment: Bsal is known to 

persist in a moist environment for up to 

7 weeks, even without an amphibian 

host. This ability creates an alarming 

pathway for the potential spread of Bsal 

into the United States through a variety 

of means not fully addressed by the 

2016 interim rule. This unchecked 

pathway of Bsal into the United States 

presents a major limitation in our ability 

to prevent introduction of this 

potentially devastating infectious 

wildlife disease. 

Response: Materials that can transmit 

pathogens, such as water, represent a 

potential pathway. However, the Service 

does not have authority under the 

injurious wildlife listing provisions in 

18 U.S.C. 42(a) to prohibit importations 

of water and fomites that may be 

infected with the Bsal pathogen. Listing 

the species that can carry Bsal is 

expected to limit the movement of such 

materials, but they do remain a concern. 

The Service will continue to explore 

opportunities to address this issue with 

partners and stakeholders. 

(28) Comment: A comment suggested 

that it is premature to discount frogs 

and toads (anurans) and caecilians from 

getting Bsal. 

Response: Under this final rule, we 

listed salamanders for which we had 

affirmation at the time of the rule 

drafting that they could carry Bsal into 

the United States, and subsequent 

evidence confirms the determination. 

We do know about positive Bsal test 

results for several species of anurans 

and will continue to monitor research 

on them and caecilians and on 

salamanders for which data is currently 

unavailable. 

(29) Comment: The rule is 

unnecessary and will only hurt 

hobbyists. Hobbyists who keep 

salamanders may be tempted to release 

them into the wild if they cannot find 

alternatives and do not want to 

euthanize them. If so, the risk of Bsal 

being introduced into the wild might be 

increased. 

Response: We believe this regulatory 

action will safeguard the health of wild 

salamanders and those kept in captivity. 

We have taken action with this rule to 

list salamanders that we find can carry 

Bsal. Pet owners will still be allowed to 

keep their salamanders and sell or give 

them away within the enumerated 

jurisdictions of 18 U.S.C. 42, also set 

forth at 50 CFR 16.3. In addition, many 

States have laws making it illegal to 

release certain animals into the wild, 

and injurious listed species cannot be 

released into the wild under Federal 

law. Some States have amnesty 

programs that accept unwanted pets. 

The Service believes that the majority of 

pet owners and hobbyists would not 

intentionally release their animals into 

the wild; however, the pet trade was 


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identified as the major vector for a 

potential Bsal invasion. To assist pet 

owners who might need to find homes 

for their animals, we posted information 

about responsible alternatives to 

releasing salamanders on our website 

when we published the 2016 interim 

rule. That updated information can be 

found at https://www.fws.gov/node/ 

266100. 

(30) Comment: The interstate 

prohibition will not help prevent the 

spread as the zoospores are most likely 

going to be spread through moving 

water. Also, many wildlife diseases are 

moved by wildlife themselves, 

including migratory birds. Without 

evidence of infected animals in the 

trade, it is inappropriate to indict an 

industry or to blockade any trade based 

on speculation. Additional studies are 

needed to determine sources and causes 

for outbreaks. Without further 

surveillance and supportive data, it 

cannot be substantiated that the 

international and interstate trade is the 

vector for spread of this disease. 

Response: As we note in the final 

rule, the interstate prohibition has been 

clarified. In the 2016 interim rule, we 

did not indicate that the absolute cause 

of the spread of Bsal is the wildlife 

trade, although we concluded that the 

most likely pathway of Bsal into the 

United States is on the bodies of 

salamanders in the commercial 

salamander trade. We cited peerreviewed journal articles that suggest 

the spread of Bsal has been human 

mediated due to the discontinuity of the 

global distribution of Bsal between Asia 

and Europe, and we cite the detection 

of the pathogen in imported captive 

exotics. Both of these pieces of 

information suggest the spread of Bsal 

has been human mediated. Other 

pathways for Bsal introduction are not 

expected to be as significant compared 

to the international-trade pathway. 

While the Service is concerned about 

contaminated water, Bsal is not yet 

known to be present in the United 

States. Listing is intended to prevent the 

introduction, establishment, and spread 

of Bsal. Salamanders would have to 

come into contact with Bsalcontaminated water for the pathogen to 

be introduced. If no infected 

salamanders are here, they cannot 

transmit the pathogen to waters that can 

further spread the pathogen. 

Research suggests that waterfowl can 

carry Bd on their toes, although Bd 

could not survive more than 60 minutes 

of desiccation on the scale tissue 

(Garmyn et al. 2012). As a result, while 

Bd could be transmitted from one 

habitat to another on short flights, 

transmission is unlikely to be an 

intercontinental threat. Given the 

similarities between Bd and Bsal, Bsal is 

not likely to be introduced to the United 

States through bird migrations. 

Border Interstate Transportation 

(31) Comment: The prohibition on 

importation will help to prevent the 

movement of Bsal into the United States 

provided that it is also prevented from 

entering Canada. If an infected 

salamander enters through Canada, Bsal 

could be transported via water and 

waterfowl into the United States, 

negating the prohibition’s benefits. 

Response: In 2017, after the 2016 

interim rule was published, Canada 

passed a law prohibiting importation of 

all species of the order Caudata, alive or 

dead, and their gametes (ECCC 2017, 

2018). Canada, Mexico, and the United 

States actively coordinate in wildlife 

conservation issues through the Canada/ 

Mexico/U.S. Trilateral Committee of 

Wildlife and Ecosystem Conservation 

and Management meetings. 

(32) Comment: The interstate 

prohibition will make it harder to 

acquire scarce animals. Prohibiting 

interstate movement will hurt honest 

hobbyists who are working hard to find 

or produce healthy captive-bred 

animals. The prohibition should apply 

only to wild-caught animals or 

importation only but allow for 

movement of captive-bred animals in 

the United States that have been tested 

and found to be free of Bsal, especially 

since Bsal has not been found in the 

United States. 

Response: As explained under A. 

Background in II. Final Rule to the 2016 

Interim Rule, the interstate prohibition 

has been clarified. Under 18 U.S.C. 42, 

the Service does not have the authority 

to selectively prohibit the importation of 

wild-caught or captive-bred animals for 

a species listed as injurious wildlife. 

Permits can be acquired for zoological, 

educational, medical, or scientific 

purposes. 

Effect on Hobbyists 

(33) Comment: Captive-breeding 

should be legal for private and hobbyist 

purposes. 

Response: Captive-breeding is not 

prohibited by the injurious wildlife 

provisions of the Lacey Act as a result 

of listing species of salamanders as 

injurious wildlife under the rule. 

(34) Comment: The rule will have a 

direct effect on both amphibian business 

owners and hobbyists as well as native 

ecosystems. The species that are listed 

are those most important to the hobby— 

animals that are easy to breed and that 

do well in captivity. The rule effectively 

transitions the hobby almost entirely 

away from captive-breeding. Captivebred animals are healthier, less likely to 

carry diseases, more likely to thrive in 

captivity, and do not harm wild 

populations. Commercially wildcollecting animals can cause long-term 

damages to populations and has been 

known to play a role in disease 

transmission as collectors travel 

between areas and do not disinfect their 

equipment. 

Response: The commenter states that 

captive-bred animals are healthier and 

less likely to carry diseases but does not 

provide evidence to support this 

statement. State wildlife agencies are 

responsible for regulating the collection 

of most wild salamanders, including 

injurious listed ones, and State 

authorities can be used to protect 

populations from overharvest. 

(35) Comment: The science is wrong 

on the number of salamanders crossing 

State lines. The commenter knows one 

individual who sold 1,500 captive-bred 

tiger salamanders last year outside their 

State. The interstate prohibition will 

cause a drop in the diversity of captivebred species and related expertise in the 

country. This prohibition will severely 

limit many forms of research since 

expert American salamander keepers 

will be unable to maintain and share 

their experience through captivebreeding programs. Researchers will be 

limited largely to axolotls (Ambystoma 

mexicanum), which may not work for 

their needs. Even Martel et al. (2014) 

was largely dependent on captive-bred 

animals; in a few years, a similar study 

will be impossible from the United 

States. 

Response: The rule will not end 

scientific endeavors that would benefit 

the injurious listed species. 

Additionally, as explained in this final 

rule, the prohibition on interstate 

movement between States within the 

continental United States has been 

clarified. 

(36) Comment: The rule interferes 

with educational opportunities and 

exposes exhibitors, nature centers, 

wildlife rehabilitators, private citizen 

hobbyists, and commercial breeders to 

Federal prosecution and penalties under 

the Lacey Act. 

Response: The rule is intended to 

protect native species, which will help 

ensure that the public maintains the 

opportunity to enjoy them in their wild 

habitats. Also, the injurious wildlife 

provisions of the Lacey Act do not 

prohibit ownership or breeding of 

injurious wildlife, unless unlawfully 

imported or transported between the 

enumerated jurisdictions or otherwise 

restricted due to conditions associated 

with issued permits. People and 


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zoological institutions can still own 

salamanders where consistent with 

other Federal, State, and Tribal laws and 

regulations applicable to the species. 

The listing also will not prevent the 

continued use of these species for 

education, and prohibited activities may 

be authorized by permit for zoological, 

educational, medical, or scientific 

purposes (in accordance with permit 

conditions). Finally, as explained in the 

final rule, the interstate prohibition 

between States within the continental 

United States has been clarified. 

Effect on Conservation Efforts 

(37) Comment: Captive-breeding has 

been proven to be the most reliable way 

of ensuring the survival of endangered 

(or common) species. Furthermore, 

captive-breeding provides a backup 

gene pool for wild populations that may 

be drastically reduced from Bsal. Also, 

the listing would make it illegal to 

transport listed salamander species 

across State lines and would devastate 

conservation programs across the 

United States. The permitting process 

will keep many zoos and aquariums 

from participating in propagation efforts 

of salamander species on the list, many 

of which need help. 

Response: While captive-breeding is 

useful in many cases to ensure survival, 

it is less so when a novel, lethal 

pathogen is the cause. Listing the 

species as injurious in this rule will not 

affect legitimate conservation efforts 

that U.S. breeders can carry out for the 

species. The law allows for the issuance 

of permits authorizing otherwise 

prohibited movement or imports for 

scientific or zoological purposes, 

including non-commercial conservation 

breeding operations. The Service has 

provided information online to help 

people apply for a permit (see 

Permitting Difficulties below in this 

comment discussion for additional 

details). Finally, as explained in the 

final rule, the current prohibition on 

interstate transport in 18 U.S.C. 42(a) 

has been clarified and does not apply to 

interstate transport between States 

within the continental United States. 

(38) Comment: When scientists collect 

tissues or specimens for lab 

experiments, the animals are never 

released into the wild and therefore 

pose no threat to the spread of Bsal or 

any other pathogen. The Service’s 

imposition of increased Federal 

permitting will inhibit scientists who 

are studying the biology of regulated 

species and may dissuade graduate 

students or other biologists from such 

work. This type of regulatory change 

can hinder conservation efforts before 

their need can even be evaluated. 

Response: This listing should not 

adversely affect any valid conservation 

efforts. In general, all wildlife species 

must be declared at the time of 

importation (see 50 CFR part 14), but 

most do not require special permits. 

Prior to this rule, only species of 

salamanders listed under the 

Endangered Species Act (ESA) or CITES 

required import permits under those 

wildlife laws implemented by the 

Service. For injurious wildlife, permits 

are not needed for interstate transport 

between the States within the 

continental United States (except into or 

out of the District of Columbia), and 

permits to allow import and transport 

between the enumerated jurisdictions in 

the shipment clause of 18 U.S.C. 42(a) 

may be granted for bona fide scientific 

purposes. This rule should have no 

significant effect on any conservation 

efforts that are currently being or will be 

carried out. 

(39) Comment: One commenter has 

never owned a pet frog or salamander 

yet has educated more than 3,000,000 

people about amphibians via online and 

printed educational materials and 

through live presentations and hikes to 

amphibian habitats to see local, native 

wild amphibians. The commenter states 

that truly inspirational amphibian 

experiences occur when humans come 

across wild amphibians, not captive 

amphibians. 

Response: The Service encourages 

visitors to the Service’s national wildlife 

refuges and other public lands to 

appreciate salamanders in their natural 

environments. The purpose of listing 

these salamander species as injurious 

wildlife is to prevent the introduction, 

establishment, and spread of Bsal in the 

wild in the United States to protect 

wildlife and wildlife resources, 

including native salamanders in the 

wild. 

(40) Comment: The rule prevents the 

ability of salamander owners to further 

test their collections and, therefore, 

could unintentionally increase the 

spread of this disease rather than 

decrease it, if it arrives in this country. 

Another commenter noted that the 

current prohibition, especially on 

interstate movement, will discourage 

cooperation to get domestic collections 

tested for the disease. 

Response: As explained in the final 

rule, the current prohibition on 

interstate transport in 18 U.S.C. 42(a) 

has been clarified and does not apply to 

interstate transport between States 

within the continental United States. 

Treatment and testing that does not 

involve import into the United States, 

transport between the enumerated 

jurisdictions in 18 U.S.C. 42(a) (also set 

forth at 50 CFR 16.3), or injurious 

wildlife permits are not regulated by 

this rule. 

(41) Comment: The rule does not list 

members of the Ambystoma genus, so 

this omission may increase the chances 

of legal and illegal collection of 

Ambystoma. 

Response: Listing a species as 

injurious wildlife results in prohibitions 

on import into the United States and 

shipment between the enumerated 

jurisdictions in 18 U.S.C. 42(a), codified 

in Federal regulations at 50 CFR 16.3. 

Neither listing a species as injurious nor 

not listing it results in a prohibition on 

collection. It is the responsibility of a 

person who may be engaged in 

salamander collection to be aware of 

any Federal, State, Tribal, or territorial 

law or regulation that applies to such 

activity. For example, some salamanders 

are federally protected from take 

(including, but not limited to, 

collection) under the Endangered 

Species Act, and other laws or 

regulations may otherwise prohibit or 

regulate collection of other salamanders 

in national wildlife refuges, national 

parks, or other Federal lands, or in 

accordance with State or Tribal laws. 

While it is possible that some people 

will switch to Ambystoma spp. in place 

of a listed species if they want to keep 

salamanders, they may currently do so 

in States where it is legal under State 

law. We are listing the genus 

Ambystoma with the second interim 

rule as a way to prevent the potential 

introduction of the fungus. 

Permitting Difficulties 

(42) Comment: Multiple commenters 

expressed concern that the listing would 

complicate research efforts or breeding 

programs for recovery efforts for some 

native salamanders due to extended 

permit-application processing time and 

limited Federal resources to adequately 

address an increased number of 

applications. 

Response: As explained in the final 

rule, the current prohibition on 

interstate transport in 18 U.S.C. 42(a) 

has been clarified and does not apply to 

interstate transport between States 

within the continental United States. 

Fewer permit requests will be required 

because interstate transport between 

States within the continental States is 

not prohibited. 

(43) Comment: The Service should 

consider adopting a cooperative 

agreement or memorandum of 

agreement to allow easier movement of 

prohibited species for certain purposes. 

Response: Several commenters 

suggested memoranda of understanding 

(MOUs) or other mechanisms in lieu of 


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permits. Those arrangements cannot be 

used to authorize import or transport 

between the enumerated jurisdictions in 

the shipment clause of 18 U.S.C. 42 (the 

continental United States, the District of 

Columbia, Hawaii, the Commonwealth 

of Puerto Rico, and any territory or 

possession of the United States), which 

are codified in Federal regulations at 50 

CFR 16.3. Other interstate transport 

between States within the continental 

United States is not prohibited by 18 

U.S.C. 42(a). The text of 18 U.S.C. 

42(a)(3) requires that exceptions to 

otherwise prohibited activities with 

injurious wildlife be authorized by 

permit, and only if there has been a 

proper showing of responsibility and 

continued protection of the public 

interest and health. The regulations at 

50 CFR 16.22 specifically provide that 

the Service may issue a permit 

authorizing the importation into or 

shipment between the continental 

United States. Thus, MOUs cannot be 

utilized for authorizing import or 

shipment between the enumerated 

jurisdictions. We have provided 

information online that helps people 

who are requesting a permit understand 

and navigate the process at https://

www.fws.gov/node/266100. The Service 

is committed to processing permit 

applications as quickly as possible to 

minimize any delay or disruption of 

legitimate activities. Permit applications 

can be found here: https://fws.gov/ 

service/3-200-42-importacquisitiontransport-injurious-wildlifeunder-lacey-act. 

(44) Comment: A commenter 

recommends that, to receive a permit to 

transport potentially infectious (noninactivated) material, be it live or dead 

salamanders, parts of dead salamanders, 

or biological samples, one of the 

requirements should be proving absence 

of infection with Bsal. To acquire a 

permit, the sender or receiver or both 

would have to quarantine the 

salamanders or other material (and 

demonstrate that the quarantine 

measures are adequate to contain spread 

of the pathogen), sample a percentage of 

the total number of animals or biologic 

materials to be shipped, and submit 

those samples, such as skin swabs from 

live or dead non-fixed salamanders, to 

a diagnostic laboratory for PCR testing. 

Permit granting would depend upon 

confirmation of the negative status of 

the animals or biologic materials. 

Response: While testing of specimens 

and live animals before moving them 

would be advisable, testing could not be 

a prerequisite for receiving a permit at 

this time because the details of reliable 

testing from all exporting countries have 

not been confirmed. And, as mentioned 

in IV. Second Interim Rule below, 

interstate transport between States 

within the continental United States is 

not prohibited under the current 

prohibitions of 18 U.S.C. 42(a) for the 

listed salamanders, making the 

requirement not necessary for many 

domestic shippers. We recommend that 

salamander transporters conduct best 

practices to reduce the risk of 

introducing, transporting, or spreading 

Bsal within the United States. 

(45) Comment: The 2016 interim rule 

should be amended to allow accredited 

veterinary medical diagnostic 

laboratories to exchange, receive, and 

accept live or dead specimens, 

including parts of the 201 listed species, 

without the requirement of first 

obtaining a Federal permit. The first 

step in any Bsal response is to obtain an 

accurate and confirmed diagnosis of 

Bsal. Requiring accredited labs to first 

obtain a permit is an unnecessary 

burden that slows the diagnostic process 

and any confirming diagnostic testing at 

different labs. 

Response: We agree that the first step 

to any Bsal response is obtaining 

accurate diagnosis of Bsal. However, a 

permit is no longer necessary for 

shipment between States within the 

continental United States, as explained 

below in the preamble to this final rule. 

(46) Comment: The doublecontainment requirements for transport 

and storage and uncertainties therein 

are concerning. More explicit guidance 

is requested regarding the doublecontainment requirements for transport, 

housing, or storage, or handling of 

animals, tissues, or other samples. 

Specifically, how does this requirement 

apply to species repatriation projects or 

State-approved releases of injurious 

listed salamanders back into the wild? 

Many States conduct health testing (in 

collaboration with diagnostic lab 

partners) and have established 

standards that must be met before 

repatriation is conducted. Such Statesponsored activities should be exempt. 

Another solution is to permit 

exemptions for double containment of 

fixed tissues, where the threat of Bsal 

transmission is removed by virtue of the 

fixative agent. 

Response: The Service posted 

additional guidance on our website that 

includes further discussion about the 

‘‘double escape-proof’’ containment for 

live animals and samples (https://

www.fws.gov/node/266100). It is 

possible, however, in situations where 

live animals have been permitted and 

for which the ‘‘double escape-proof’’ 

containment requirements would apply, 

that repatriation would run counter to 

that requirement. This is the first time 

that native species that might be part of 

repatriation or recovery efforts have 

been listed as injurious. Because 

injurious wildlife must be carefully 

handled, all of the containment 

requirements must be met when 

salamanders are in captivity. However, 

the Service will work with people or 

institutions that are involved in Stateapproved repatriation efforts to facilitate 

these efforts. Finally, we clarify in A. 

Background in II. Final Rule to the 2016 

Interim Rule that preserved tissues are 

not considered injurious. 

Other Impacts 

(47) Comment: Collection of fishes for 

shipment cannot totally ensure that 

other species of ‘‘free riders,’’ such as 

non-marketed amphibians, are not 

unintentionally included in the 

shipment process. Unintentionally 

including a single regulated amphibian, 

regardless of whether it is infected with 

Bsal, would subject the transporting 

farmer to severe civil and even criminal 

penalties. Notably, actual interstate 

transport of Bsal by some means not 

including a listed amphibian would not 

violate the rule. 

Response: As explained in the final 

rule, interstate transport prohibitions 

have been clarified. We encourage 

anyone who transports live fishes to use 

best management practices that include 

transporting only the traded species and 

their uncontaminated media. 

Unintentional importation or transport 

between the enumerated jurisdictions in 

the shipment clause of 18 U.S.C. 42 (the 

continental United States, the District of 

Columbia, Hawaii, the Commonwealth 

of Puerto Rico, and any territory or 

possession of the United States), which 

are also set forth at 50 CFR 16.3, through 

nursery stock or other pathways would 

also be a violation of the prohibitions 

from listing. 

Inaccurate or Incomplete Science 

(48) Comment: The rule does not 

account for pressures that amphibians 

are already facing, such as habitat loss, 

rising temperatures, pesticide use, and 

siltation from agriculture. The Service’s 

focus should be on the systematic 

degradation of the ecosystems in which 

the amphibians live and the capacity of 

the salamanders to fight the fungus. 

Response: The Service noted in the 

rule that salamanders may demonstrate 

more severe clinical disease when 

infection is combined with additional 

stressors in the wild. The comment does 

not provide any evidence of how habitat 

loss, rising temperatures, pesticide use, 

and siltation from agriculture 

diminishes the need for or benefits of 

the rule that may prevent salamander 


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mortality. Native salamander species 

known to be negatively affected by Bsal 

infection under experimental conditions 

may demonstrate more severe clinical 

disease when infection is combined 

with additional stressors in the wild, as 

has been found for other diseases. 

Besides this rule, the Service is engaged 

in many other conservation measures 

designed to help improve and protect 

salamander habitats across the United 

States. 

(49) Comment: Using a method of 

infecting a salamander from one genus 

with Bsal in a laboratory setting and 

then extrapolating results to all species 

within that genus is not in conformance 

with the framework of the World 

Organisation for Animal Health code or 

the National Aquatic Animal Health 

Plan (National Aquaculture Health Plan 

and Standards) for the United States and 

is contrary to the credible scientific 

findings of Martel et al. (2014). 

Therefore, the 2016 interim rule is 

arbitrary and capricious in violation of 

the APA. In addition, other aquatic 

diseases have shown laboratory 

infection, but the affected fish species 

are not included in regulatory lists. 

Response: The WOAH has a different 

purpose than the injurious wildlife 

listing provisions of the Lacey Act, and 

the standards the WOAH uses are 

appropriate for their purposes. We 

followed the standards in 18 U.S.C. 42 

and the APA. The issue of fishes that 

may carry diseases is beyond the scope 

of this rulemaking. Surrogate species are 

used elsewhere in the 2016 interim rule, 

such as for Bd for where information is 

lacking for Bsal and is common in 

scientific literature. 

(50) Comment: The spread dynamics 

of Bd and Bsal are considerably 

different. Given that Bd is endemic to 

the United States, the estimated 

potential for Bsal distribution has been 

overestimated. This overestimation is 

confirmed by salamander import data, 

the lack of presence of Bsal in animals 

entering the United States, and its lack 

of presence in wild populations. 

Response: The commenter states that 

Bd originated in the United States and 

is therefore endemic. We agree that Bd 

has occurred in the United States for 

many years and is currently ubiquitous 

throughout North America; however, we 

do not consider Bd endemic to the 

United States. For reasons identified in 

the 2016 interim rule and this second 

interim rule, we conclude that Bsal does 

pose a risk to native salamander 

populations. We have updated the 

research cited and still conclude that 

there is a risk of Bsal entering the 

country with salamanders, and that risk 

is greatly reduced by listing the genera 

in this rule. 

(51) Comment: The 2016 interim rule 

reports that there is no accurate way to 

test for or eliminate Bsal in captivity. 

PCR-based testing has been well 

established for many years for the 

related Bd and has been effectively 

demonstrated for Bsal. Effective 

measures for clearing salamanders using 

heat alone or heat in conjunction with 

anti-fungal medications have also been 

published. The authors of both of those 

studies have reiterated in personal 

communication that the stated Service 

position in the rule justification is 

contradictory to the published data. 

Another comment noted that combined 

experience from members of 

Caudata.org in the captive maintenance 

and breeding of the species subject to 

this rule has shown that the 

temperatures required by these 

treatments are safe and will not harm 

the majority of salamanders of the 

Salamandridae, the family containing 

the bulk of the regulated species. 

Response: While the comments do not 

provide any information on how the 

Service’s finding is contradictory to the 

published data, the second interim rule 

clarifies these issues. We have revised 

the rule to note that testing and 

prophylactic treatments of imports of 

salamanders to manage Bsal are 

available but have uncertain 

effectiveness when applied as a 

nationwide regulatory tool by the 

Service. 

(52) Comment: There is likely no Bsal 

in the United States, even with the huge 

numbers of salamanders that have 

recently been imported, because it gets 

too hot in the summer and too cold in 

the winter. 

Response: As discussed under Bsal 

Risk Assessment, we found that there is 

a significant risk that Bsal can establish 

and spread in the United States. Some 

areas, such as south Florida, are likely 

to have low consequences from Bsal 

introduction, in part due to 

temperatures found in the region. The 

areas most likely to have consequences 

from Bsal introduction are the Pacific 

Coast and Appalachian Mountains 

(Richgels et al. 2016). Based on 

environmental suitability, areas of the 

United States most suited to Bsal growth 

(Blooi et al. 2015a), including the 

Southwest, Southeast (except south 

Florida as just noted), and Pacific 

regions, are also the areas of highest 

salamander diversity. The large land 

mass of the United States has a broad 

range of climates, many of which are 

similar to the other continents where 

Bsal is currently found. 

(53) Comment: The Service did not 

publish the text of articles or the risk 

assessment it used for the 2016 interim 

rule. 

Response: The file for the references 

used, as well as other supporting 

information used to develop the 2016 

interim rule, was posted under 

‘‘Supporting & Related Material’’ in 

https://www.regulations.gov (Docket No. 

FWS–HQ–FAC–2015–0005), and is 

available for public inspection as noted 

under ADDRESSES in the 2016 interim 

rule. Comments and materials we 

received, as well as citations for 

supporting documentation we used in 

preparing the interim rule, were 

available for public inspection. The 

texts of publications are often covered 

by copyright laws and those are 

therefore not posted. 

(54) Comment: Species from the 

genera Ambystoma and Gyrinophilus 

were not listed because they were tested 

and proved resistant to Bsal. Why then 

were all Plethodon listed, since the one 

species tested (P. glutinosus) was also 

demonstrated to be resistant? 

Response: We did not believe that 

there was enough evidence to list 

Ambystoma or Gyrinophilus at the time 

of the 2016 interim rule, but we found 

evidence to support listing Plethodon. 

Three native salamander species 

identified as resistant to Bsal infection 

included the spring salamander 

(Gyrinophilus porphyriticus), marbled 

salamander (Ambystoma opacum), and 

spotted salamander (A. maculatum) 

(Martel et al. 2014). At the time the 

public-comment period closed, there 

was no evidence that any species within 

these genera are carriers of Bsal. We 

discuss our reasoning for listing all 

Plethodon species in the second interim 

rule under Vulnerability and Carrier 

Status. In short, however, further 

histological analysis of the slimy 

salamander revealed that Bsal could 

invade the skin long enough to move or 

transmit the pathogen to other 

salamanders. No such evidence existed 

then or now for any species in the genus 

Gyrinophilus; therefore, we are not 

listing species from that genus. As 

explained in the second interim rule 

under Vulnerability and Carrier Status 

of Native Species, we have evidence 

now of carrier capability for Ambystoma 

maculatum, A. mexicanum, and A. 

opacum and are listing the genus. 

(55) Comment: In Europe, where Bsal 

is believed to have been introduced by 

Asian imports, Bsal was found in 

populations of Salamandra and Alpine 

newts (Ichthyosaura alpestris) in the 

Netherlands and in Belgium. It has also 

been found in captive Salamandra in 

the United Kingdom and Germany, and 


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possibly in wild German populations. 

Martel et al. (2014) shows that most 

lethally vulnerable species exposed to 

Bsal in the lab showed signs of infection 

within 8 days and were dead within 3 

to 4 weeks. This means that nonresistant infected species in captive 

collections would have died during the 

comment period on the 2016 interim 

rule. 

Response: The comment suggests that 

there are no specimens carrying Bsal at 

this time that might enter the United 

States and allow Bsal to be introduced, 

establish, and spread, but does not 

provide evidence that we can use in our 

analysis of the rule. Lethally vulnerable 

specimens can still appear if the 

pathogen spreads, or if Bsal persists in 

tolerant or susceptible populations or 

carcasses. 

(56) Comment: The 2016 interim rule 

states that surveys of anglers have 

indicated that they routinely release 

salamanders into the areas where they 

fish, which includes areas that are not 

part of the salamanders’ native U.S. 

habitats, suggesting that animals are 

routinely moved long distances. No 

similar survey data exists for pet 

owners, so assuming the pet trade is the 

problem for releases is unfounded and 

targeting the pet trade simply because it 

is an easy target is unjust. 

Response: Our statement relating to 

anglers was used to note that this 

invasion expansion pathway has been 

attributed to the use and subsequent 

release of salamanders used as fishing 

bait. Along with the other evidence we 

documented, we found that there is the 

potential for salamanders carrying Bsal 

to escape or be released into the wild 

where they can transmit the pathogen to 

native species. We provided evidence in 

the rule that we used to conclude that 

international trade is the main pathway 

for the global spread of Bsal. 

Additional Science Needed 

(57) Comment: Several areas would 

benefit from further investigation. For 

example, the origins of Bsal in wild 

salamanders needs to be better 

understood. It is important to continue 

and expand testing of salamanders in 

the wild and in trade in various 

locations. Additional testing of species 

within the same genus would be 

beneficial to guide field and collection 

surveillance. 

Response: Since the 2016 interim rule 

was published, many studies have been 

published that address the commenter’s 

concerns and are applicable to the rule, 

including a major surveillance of 

salamanders in the wild by the USGS. 

We have reviewed the studies, and they 

support our final and second interim 

rules. We agree that additional science 

will help address issues related to better 

understanding of this pathogen and 

preventing its introduction into the 

United States, but we understand the 

need to take action now to list the 

species in the genera in this rule to 

prevent the introduction, spread, and 

establishment of Bsal. 

Economic and Trade Data 

(58) Comment: If the salamanders are 

already here, and Bsal is not, then that 

means that any salamanders traveling 

across State lines pose no risk. This law 

estimates that it will cause $3.8 million 

in damage to the U.S. economy, mostly 

in the small business sector 

(‘‘Regulatory Flexibility Act,’’ paragraph 

nine). Those numbers could be greatly 

lessened if interstate travel were 

allowed. 

Response: As explained above in III. 

Final Rule to the 2016 Interim Rule in 

A. Background and D. Required 

Determinations, the current prohibition 

on interstate transport in 18 U.S.C. 42(a) 

has been clarified and does not apply to 

interstate transport between States 

within the continental United States. 

Thus, the costs incurred are expected to 

be less than originally estimated in the 

2016 draft economic analysis. 

(59) Comment: Caudata.org conducted 

an online public survey from February 

1 to March 12, 2016, to gather additional 

data of U.S. domestically bred animals. 

A total of 797 respondents to the survey 

reported shipping 25,649 domestically 

bred caudates across State lines in 1 

year. Due to the low response rate 

relative to the number of U.S. registered 

members on Caudata.org (8 percent) and 

the short duration of the survey, this 

number likely represents a small 

fraction of the actual trade. It can be 

safely extrapolated that the Service has 

underestimated the trade in captivebred newts and salamanders by at least 

two orders of magnitude. Caudata.org is 

uniquely situated at the interface of 

hobbyists, entrepreneurs, researchers, 

zoos, and aquariums. A summary of that 

data and some important numbers are 

presented here. Respondents to the 

survey possessed a total of 28,228 

domestically bred salamanders or 

newts, the majority of which are subject 

to the rule. Respondents shipped on 

average 25,649 salamanders or newts 

over State lines per year. This number 

is nearly two orders of magnitude 

greater than the ‘‘338’’ cited by the rule 

and represents just a small fraction of 

our members. The total yearly 

salamander- and newt-related revenue 

reported by our respondents was 

$207,528 for 2015. 

The commenter further stated that 

Caudata.org has more than 10,000 

unique registered U.S. members who 

have accessed their website in the past 

5 years. Their total number of unique 

U.S. visitors (people who did not 

register for an account) in that time is 

orders of magnitude greater than this 

number. The commenter stated that, 

apparently, the Service did not perform 

due diligence in ascertaining the 

number of private U.S. citizens affected 

by this rule. 

Response: While we did obtain data 

from the Pet Industry Joint Advisory 

Council (PIJAC; currently known as the 

Pet Advocacy Network) for the 2016 

interim rule, we appreciate Caudata.org 

for supplying additional data. The 

survey does not indicate: (1) whether 

the salamanders are domestically bred; 

(2) the net importation for each State; or 

(3) what species they are. Since the 

survey data did not include information 

on species or whether they are 

transported between the listed 

jurisdictions, it is unknown if any of the 

revenue discussed would be lost due to 

prohibitions under the rule. 

Consequently, the data are not used in 

the final economic analysis for the 2016 

interim rule. Furthermore, unlike the 

2016 interim rule, the final rule clarifies 

that the current prohibition on interstate 

transport in 18 U.S.C. 42(a) does not 

apply to interstate transport between 

States within the continental United 

States. 

(60) Comment: The economic figures 

provided by the Service are a gross 

understatement. Caudata.org has 

submitted the results of a survey on the 

numbers of animals sold across State 

lines, and just from their members, 

reported roughly $207,528 in income. 

Actual figures are probably much 

higher, given that this amount likely 

represents just a portion of the trade in 

the entire United States, and 

Caudata.org pertains only to captivebred animals. In addition to the money 

spent purchasing animals, there’s also 

food, lighting, enclosures, plants, 

decorations, filters, shipping and 

packaging fees, and other costs 

associated with keeping salamanders. 

To house a pair of salamanders can cost 

$100 to $200 or more, with ongoing 

feeding costs. Overall, the U.S. 

salamander hobby probably represents 

well over $5 million to $10 million in 

economic activity each year. 

Response: Regarding the Caudata.org 

data, see also response to Comment 59. 

The economic analysis addresses 

primary support services (such as food 

and shipping) and secondary economic 

impacts in Sections 2.3.2 and 3.1.3, 

respectively. 


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(61) Comment: The 2016 interim rule 

states that ‘‘a minimum of 338 

domestically bred salamanders may be 

affected due to the interstate 

transportation prohibition.’’ As an 

individual, the commenter has legally 

shipped 150-plus live specimens (eggs, 

larvae, and adults) in a single year and 

knows that many more people legally 

ship more specimens than that amount 

in the same period. The commenter has 

also received dozens of animals in a 

single year and knows that this 

occurrence is not unique. Many 

individuals will be affected by the 

listing. 

Response: The minimum is based on 

available data from PIJAC and is stated 

as a minimum due to the expectation of 

the actual number being potentially 

larger. For this salamander breeding 

data, it is unclear which species are 

shipped and whether these specimens 

are shipped between listed jurisdictions. 

Unlike the 2016 interim rule, the final 

rule clarifies that the current prohibition 

on interstate transport in 18 U.S.C. 42(a) 

does not apply to interstate transport 

between States within the continental 

United States. Therefore, it is not 

incorporated into the final economic 

analysis. 

(62) Comment: Many small businesses 

have commented that the prohibition on 

interstate transport will have a greater 

impact than the Service anticipates. In 

the 2016 interim regulatory flexibility 

analysis, the Service stated that it does 

not believe that the impact of 

prohibiting interstate transport will be 

significant. However, several small 

breeders and hobbyists involved in 

selling salamanders in the United States 

have indicated a substantial domestic 

trade in salamanders. The U.S. Small 

Business Administration Office of 

Advocacy commented that a small 

business representative indicated that 

this number could be as high as 1,500 

specimens shipped in a year for certain 

businesses. The difference between the 

limited information in the analysis and 

the information provided by 

commenters indicates that the analysis 

underestimates the effect of the 

prohibition of interstate transport. 

Response: As explained above in III. 

Final Rule to the 2016 Interim Rule in 

A. Background, the current prohibition 

on interstate transport in 18 U.S.C. 42(a) 

has been clarified and does not apply to 

interstate transport between States 

within the continental United States. 

Therefore, the interstate data provided 

are not incorporated into the final 

economic analysis and final regulatory 

flexibility analysis. As discussed in 

Comment 61, salamander breeding data 

that are submitted without specific 

details regarding species type are too 

general to be incorporated into the final 

economic analysis. It is possible that the 

domestic market is more robust than 

estimated. However, it is unclear 

whether any additional sales are related 

to species that are listed or not listed 

under the rule. 

(63) Comment: Many breeders who 

produce these animals as their main 

source of income will lose significant 

income or go out of business without 

the ability to sell across State lines. For 

example, last year, a business owner 

produced more than 100 neotenic 

Ichthyosaura alpestris, and this year 

[2016] they will have to cull those eggs 

in light of the prohibition. 

Response: The comment is incorrect 

that they will have to cull eggs, because 

eggs of listed salamander species are not 

considered injurious because they do 

not have the potential to serve as 

carriers of Bsal. Furthermore, as 

explained above in III. Final Rule to the 

2016 Interim Rule in A. Background, the 

current prohibition on interstate 

transport in 18 U.S.C. 42(a) has been 

clarified and does not apply to interstate 

transport between States within the 

continental United States after April 7, 

2017. Therefore, the interstate data 

provided are not incorporated into the 

final economic analysis and final 

regulatory flexibility analysis. 

(64) Comment: Many States also 

prohibit or limit sale by biological 

supply companies of certain native 

species, and the authority to regulate 

nonnative species may be either with 

the State fish and wildlife agency or the 

State’s department of agriculture or 

shared in some instances. For example, 

commercial production of native 

salamanders is currently not legal in 

California, and the State’s department of 

agriculture does not regulate or track 

production or sale of nonnative 

salamanders in the State. The only way 

to legally sell native salamanders in 

California is as a biological supply 

house with a permit to collect wild 

specimens for sale to scientific and 

educational facilities. Only one business 

is currently in possession of this permit, 

and it has not collected or sold 

salamanders. 

Response: We appreciate the 

information. 

(65) Comment: One commenter’s 

company produces about 1,000 

Neurergus kaiseri, 100 N. crocatus, 100 

N. strauchii, and 200 Ichthyosaura 

alpestris a year. 

Response: We appreciate the 

commenter supplying domestic 

breeding data. It has been incorporated 

as appropriate into the appendix to the 

final economic analysis. 

(66) Comment: An international 

prohibition on trading gives small-time 

breeders within the United States an 

economic boost to supply the demand 

for these pets. 

Response: We acknowledge that an 

international prohibition can have an 

indirect effect of reducing competition 

for domestic breeders in some markets. 

The rule was not implemented to 

provide an advantage to domestic 

breeders but rather to prevent Bsal 

introduction, establishment, and spread 

in the United States by salamander 

species that are carriers of the pathogen. 

(67) Comment: The Service estimates 

that, without the 2016 interim rule, 

217,000 salamanders would be imported 

each year. These imports will be 

prohibited if the 20 genera are listed 

under the Lacey Act as set forth in the 

2016 interim rule. The Service further 

estimates that 338 domestically bred 

salamanders would be affected by the 

interstate transportation prohibition per 

year, resulting in impacts to domestic 

breeders of up to $23,000. These 

domestic production numbers do not 

pass a straight-face test; for the estimate 

to be accurate, each salamander would 

need to be worth an average of $68. In 

reality, salamanders typically sell for 

between $10 and $50, depending on the 

species. As several USARK members 

and others in the herpetoculture 

industry have reported to the Service in 

written comments, including trade 

numbers provided by Caudata.org, the 

actual number of domestically bred 

salamanders shipped across State lines 

is far higher than 338. The species listed 

in the 2016 interim rule comprise the 

overwhelming majority of those in the 

pet trade, so the economic effect of the 

listing will amount to nearly the full 

total of the industry’s value. 

Response: The minimum number for 

domestic production (338) and the 

corresponding prices for those 

salamanders were provided by PIJAC. 

The detailed data they provided is in 

table A1–2. Furthermore, after the 2016 

interim rule was issued, as explained 

above in A. Background in III. Final 

Rule to the 2016 Interim Rule, the 

current prohibition on interstate 

transport in 18 U.S.C. 42(a) has been 

clarified and does not apply to interstate 

transport between States within the 

continental United States. 

Use of Categorical Exclusion 

(68) Comment: The interim rule is not 

possible without the recently 

implemented categorical exclusion that 

bypasses the requirement to consider 

economic and social impacts under the 

National Environmental Policy Act 

(NEPA). The decision to use the 


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categorical exclusion for the 2016 

interim rule is flawed. 

Response: We determined the 

categorical exclusion for injurious 

wildlife listing, located in the 

Department of the Interior Manual at 

516 DM 8.5 C(9), applies to the action 

in accordance with the requirements of 

NEPA. The categorical exclusion does 

not bypass NEPA. We reviewed the rule 

under NEPA requirements and prepared 

an environmental action statement for 

the 2016 interim rule that was available 

for review (see ‘‘Supporting & Related 

Material’’ at https://

www.regulations.gov under Docket 

Number FWS–HQ–FAC–2015–0005). 

Under NEPA, the human environment is 

interpreted comprehensively to include 

the natural and physical environment 

and the relationship of people with that 

environment (40 CFR 1508.14), and the 

economic or social effects are not 

intended by themselves to require 

preparation of an EIS (40 CFR 1508.14). 

We prepared a draft economic analysis 

and regulatory flexibility analysis 

separately as part of the required 

determinations under the APA for the 

2016 interim rule and made them 

available for public comment. We 

determined that the regulations in that 

rule will not individually or 

cumulatively have a significant effect on 

the human environment. 

Inaccurate Use of 18 U.S.C. 42(a)(1) 

(69) Comment: The use of the Lacey 

Act in this manner opens the door for 

similar regulations of other animals, 

such as dogs, cats, fishes, horses, and 

chickens. The list of species is infinite, 

as would be the economic impact they 

could have. 

Response: Under the authorities 

provided under the injurious wildlife 

provisions of the authorizing statute (18 

U.S.C. 42), the Service can list only wild 

mammals, wild birds, fishes, mollusks, 

crustaceans, amphibians, and reptiles as 

injurious wildlife, meaning the Service 

cannot list domesticated species, thus 

eliminating the possibility to list 

domesticated dogs, domesticated cats, 

domesticated horses, and domesticated 

chickens. 

(70) Comment: Congress has never 

interpreted the Lacey Act to apply to 

shipment between States within the 

continental United States. 

Response: After the 2016 interim rule 

was issued, as explained above in III. 

Final Rule to the 2016 Interim Rule, A. 

Background, the current prohibition on 

interstate transport in 18 U.S.C. 42(a) 

has been clarified and does not apply to 

interstate transport between States 

within the continental United States. 

The final rule has been modified 

consistent with the prohibition in the 

shipment clause of 18 U.S.C. 42, which 

has been codified in Federal regulations 

at 50 CFR 16.3, for transport between 

the enumerated jurisdictions (the 

continental United States, the District of 

Columbia, Hawaii, the Commonwealth 

of Puerto Rico, and any territory or 

possession of the United States). 

(71) Comment: Although the pet trade 

is primarily regulated by USDA 

agencies, including the Animal and 

Plant Health Inspection Service, 

requirements for the movement of pets 

across State lines are generally reserved 

to individual States. Furthermore, 

courts have often found individual 

animals to be exempt from livestock 

regulations that would otherwise apply 

when those animals are characterized as 

household pets. Salamanders in the pet 

trade should similarly not be considered 

‘‘wildlife’’ for purposes of Federal 

regulation of interstate transport. Other 

commenters also stated that the Service 

should defer this issue to another 

agency with additional resources for 

controlling importation (such as the 

USDA) because the Service has stated 

that an injurious listing under the Lacey 

Act is their only means of attempting to 

control Bsal. 

Response: The provisions of 18 U.S.C. 

42 make no distinction between pet 

salamanders and other salamanders. 

The purpose of listing these salamander 

species as injurious wildlife is to 

prevent the introduction, establishment, 

and spread of Bsal in the wild in the 

United States to protect wildlife and 

wildlife resources. The authority to take 

action to list species as injurious 

wildlife under 18 U.S.C. 42 lies solely 

with the U.S. Department of the Interior. 

(72) Comment: The regulations 

promulgated in the 2016 interim rule 

restrict not only international and 

interstate transport but any movement 

whatsoever of the listed genera. The 

regulatory language prohibits the 

importation, transportation, or 

acquisition of any live or dead 

specimen, including parts, but not eggs 

or gametes, of the genera. There is 

simply no authority in the Lacey Act to 

prohibit acquisition. Because the Lacey 

Act does not forbid acquisition of a 

listed animal, the interim regulation is 

beyond the law to the extent it purports 

to prohibit the same. The Service must 

amend the 2016 interim rule to clarify 

that the prohibitions do not apply to 

intrastate activities. 

Response: Under the Lacey Act 

Amendments of 1981, 16 U.S.C. 

3372(a)(1), it is unlawful among other 

things for any person to sell, receive, 

acquire, or purchase any wildlife 

transported in violation of any law of 

the United States. This includes 

acquiring any injurious wildlife 

imported into the United States or 

transported between the enumerated 

jurisdictions in violation of the 

shipment clause of 18 U.S.C. 42 (the 

continental United States, the District of 

Columbia, Hawaii, the Commonwealth 

of Puerto Rico, and any territory or 

possession of the United States), also set 

forth at 50 CFR 16.3. The conditions of 

injurious wildlife permits may also 

place limitations on subsequent sale or 

transfer of injurious wildlife under the 

permit without prior authorization from 

the Service. Those activities are in 

connection with transport of injurious 

wildlife between the listed jurisdictions 

in the shipment clause or import into 

the United States under 18 U.S.C. 42. It 

is the responsibility of a person who 

may be engaged in salamander 

acquisition to be aware of any Federal, 

State, Tribal, or territorial law or 

regulation that applies to that activity. 

The rule adding injurious 

salamanders to the lists of species does 

not change the scope of the prohibitions 

in 18 U.S.C. 42, 16 U.S.C. 3372, 50 CFR 

16.3, or otherwise found in 50 CFR part 

16. The regulatory language referenced 

by the commenters (50 CFR 16.14) is 

identical to longstanding, existing 

language that appears at 50 CFR 16.11, 

16.12, 16.13, and 16.15. Revision of the 

general regulations found at 50 CFR part 

16 is beyond the scope of this 

rulemaking. 

(73) Comment: The Lacey Act defines 

‘‘transport’’ as ‘‘to move, convey, carry, 

or ship by any means, or to deliver or 

receive for the purpose of movement, 

conveyance, carriage, or shipment.’’ 

Even if the law did authorize the 

Service to bar personal transport, which 

it does not, this definition would reach 

solely intrastate activities. Although the 

Service has agreed that States, not the 

Service, have the power to regulate 

ownership and sales within their 

borders, the commenter is concerned 

that the Service is laying the 

groundwork to involve itself in Federal 

regulation of wholly intrastate activities. 

Response: The definition quoted by 

the commenter applies to the law 

codified at 16 U.S.C. 3371(j), also 

known as the Lacey Act Amendments of 

1981. Consistent with this definition, 

Service regulations also provide a 

definition of transport found in 50 CFR 

10.12. It is the responsibility of a person 

who may be engaged in salamander 

transportation to be aware of any 

Federal, State, Tribal, or territorial law 

or regulation that applies to such 

activity. For further information see also 

response to Comment 72. 


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Interim Rule Is a Regulatory Taking 

(74) Comment: The Service evaluated 

the 2016 interim rule and determined 

that it does not constitute taking. This 

conclusion is facially false—a restriction 

on interstate travel with a family pet not 

only is impermissible under the law, but 

most certainly denies the pet owner 

enjoyment and companionship 

(amounting to use) of that pet. 

Response: Import and transport of 

injurious wildlife between the 

enumerated jurisdictions in the 

shipment clause of 18 U.S.C. 42 (and at 

50 CFR 16.3) of any of the listed species 

is prohibited. The provisions of 18 

U.S.C. 42(a) do not prohibit any person 

who owns one of the listed species at 

time of listing from continuing to 

possess the species (such as listed 

salamanders) or engaging in transport 

and other activities within the 

enumerated jurisdictions of the 

shipment clause, as allowed under 

State, Tribal, or territorial law. 

Therefore, we concluded that the 2016 

interim rule and this final rule do not 

constitute a regulatory taking. This 

action is consistent with all previous 

injurious wildlife listings that have 

affected listed species that members of 

the public might have owned at the time 

of listing. It is the responsibility of a 

person who may be engaged in 

salamander transportation to be aware 

of any Federal, State, Tribal, or 

territorial law or regulation that applies 

to that activity. 

Federalism Assessment Under 

Executive Order 13132 

(75) Comment: Under Executive Order 

13132, the 2016 interim rule requires a 

federalism assessment as the rule’s 

provisions have significant federalism 

effects and will have several direct 

effects on States, which have primary 

jurisdiction over native wild animals 

not in captivity. The regulation of the 

movement of pets across State lines is 

reserved to individual States. Under 

Executive Order 13132, this interim rule 

does have sufficient federalism 

implications to warrant the preparation 

of a federalism assessment. 

Response: A federalism assessment is 

not required. Executive Order 13132 

says that policies that have federalism 

implications refer to regulations, 

legislative comments, or proposed 

legislation, and other policy statements 

or actions that have substantial direct 

effects on the States, on the relationship 

between the national government and 

the States, or on the distribution of 

power and responsibilities among the 

various levels of government. This rule 

does not limit the policymaking 

discretion of the States or preempt State 

law. The States are not restricted from 

also regulating the transport of listed 

salamanders or other activities related to 

such species within their State 

boundaries, such as sale or possession. 

The commenter did not provide 

evidence showing how the rule would 

be a substantial direct action impacting 

the States. 

Law Enforcement Issues 

(76) Comment: Salamanders will be 

smuggled into the country or sold 

through the black market once they are 

prohibited, as all contraband inevitably 

is, with no regard for fungal safety or 

often the health of the animals. One 

commenter noted that they have 

received messages from overseas asking 

them to illegally ship animals, with 

instructions for how to package and 

ship animals to demonstrate how easy it 

is to do so. The rule will not prevent 

Bsal from entering the United States. 

Response: The injurious wildlife 

provisions of 18 U.S.C. 42 serve an 

important role in protecting humans, 

the interests of agriculture, horticulture, 

and forestry, and the wildlife or wildlife 

resources of the United States from 

injurious wildlife. The rule is intended 

to reduce opportunities for Bsal to 

spread disease to native species in the 

wild. As previously explained, the 

listing of salamander species that may 

be carriers of Bsal results in prohibitions 

on import and transport between the 

enumerated jurisdictions in the 

shipment clause, and violations of these 

prohibitions are subject to strict 

liability, 18 U.S.C. 42(b) (Whoever 

violates this section, or any regulation 

issued pursuant thereto, shall be fined 

under this title or imprisoned not more 

than six months, or both.). Additionally, 

pursuant to 18 U.S.C. 42(a)(1), all 

prohibited injurious wildlife imported 

or transported in violation of the Lacey 

Act ‘‘shall be promptly exported or 

destroyed at the expense of the importer 

or consignee.’’ Where applicable, 

penalties may also be assessed under 

the Lacey Act Amendments of 1981, 16 

U.S.C. 3371 et seq. Although we 

acknowledge that some unscrupulous 

dealers may take advantage of people or 

engage in illegal trade, the regulatory 

provisions we are promulgating play an 

important role in deterring and, as 

necessary, penalizing and remedying 

unlawful activity, in order to protect the 

interests under the Act. We strongly 

encourage compliance with the law, and 

we may take appropriate enforcement 

action against violations that may occur. 

However, our experience is that pet 

owners prefer to be responsible, lawabiding citizens and would make 

informed decisions not to engage in 

import or transport contrary to the 

Lacey Act and thereby reduce the risk 

of spreading Bsal. 

(77) Comment: The final rule will 

subject exhibitors, nature centers, 

wildlife rehabilitators, private citizen 

hobbyists, and commercial breeders to 

Federal prosecution and penalties from 

felonies under the Lacey Act. 

Response: The interstate transport 

language in 18 U.S.C. 42 is clarified in 

A. Background in III. Final Rule to the 

2016 Interim Rule. Prohibitions remain 

for importation and transportation 

between the enumerated jurisdictions of 

18 U.S.C. 42, also listed in 50 CFR 16.3. 

Violations of the injurious wildlife 

listing prohibitions are a misdemeanor, 

not a felony. 

(78) Comment: For salamander 

species not listed as injurious, the final 

rule should incorporate authority for the 

Service to collect Bsal samples from any 

shipment where dead animals are 

present upon importation. This is a 

noninvasive procedure, and these data 

are needed to help modify this rule in 

the future if additional Bsal carrier 

species are discovered. The presence of 

dead salamanders upon importation can 

be a smoking gun for the presence of 

Bsal (and other harmful pathogens). 

Response: The recommended action is 

outside the scope of this final rule 

relating to the listing of injurious 

wildlife under 18 U.S.C. 42(a). 

(79) Comment: A Federal ban on 

interstate movement of salamanders is 

unenforceable given the Service’s 

resource limitations. Many of the exotic 

caudate species listed as injurious are 

already widely distributed in private 

collections in virtually every State. 

There is no system in existence (or 

resources to create a system) to register 

or effectively monitor their numbers or 

locations. 

Response: The Federal ban was lifted 

in 2017 as the result of a Federal court 

decision regarding the interpretation of 

the statute 18 U.S.C. 42(a). This final 

rule has been amended to address 

interstate transport as explained above 

in III. Final Rule to the 2016 Interim 

Rule in A. Background and D. Required 

Determinations. Whether a Federal ban 

on interstate movement is enforceable is 

beyond the scope of this rule. 

(80) Comment: Regulation alone will 

not put a halt to the international and 

interstate traffic in species listed as 

injurious under the Lacey Act or under 

various State regulations. Accordingly, 

adequate law enforcement, especially at 

ports of entry, is critical to manage the 

ongoing, and possibly increased, 

volume of underground traffic in 

regulated wildlife. The commenter 


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encourages building cooperative 

partnerships between State and Federal 

enforcement agencies to increase 

capacity and to capitalize on the 

specific expertise that the respective 

programs can bring to bear on this 

problem. 

Response: We agree. The Service’s 

Law Enforcement Office has a 

longstanding relationship with Customs 

and Border Protection and USDA 

inspectors regarding cooperation for 

enforcement on the borders. The 

Service’s law enforcement officers at 

ports have and will continue to 

maintain strong working relationships 

with their State counterparts. 

(81) Comment: The trade in species 

not listed by the rule needs to be 

monitored. The rule’s prohibitions of 

the vast majority of commonly traded 

species may inadvertently create a new 

legal market for species not previously 

in demand by the salamander trade 

community. Those newly traded species 

could be carriers of Bsal. 

Response: The Service collects 

information on all imported 

salamanders, listed or otherwise. This 

situation will not change with the 

listing of the species in this rule as 

injurious wildlife. This rule does not 

preclude the ability to take additional 

regulatory actions if new information 

emerges. 

(82) Comment: There was a push to 

acquire species before the prohibition 

could go into effect after it was 

announced. Prior to the ban, some 

people would have only purchased 

captive-bred or long-time-in-captivity 

amphibians. Due to the prohibition, 

they stepped out of their comfort zone 

and purchased wild-caught 

salamanders. 

Response: The comment supports the 

Service’s decision to implement an 

effective date of 15 days after the date 

of publication for the 2016 interim rule 

and again for the second interim rule. 

We wanted to give shipments in transit 

or pending transit the time needed to 

complete the travel for the welfare of the 

live animals, but we did not want to 

encourage a rush to import over a longer 

period. Purchasing a wild-caught 

salamander listed under this rule is not 

prohibited under 18 U.S.C. 42, provided 

transport of the specimen occurs only 

within an enumerated jurisdiction of 18 

U.S.C. 42 (also listed at 50 CFR 16.3) 

and complies with any permit condition 

for a specimen traded under an 

injurious wildlife permit. It is the 

responsibility of a person who may be 

engaged in salamander acquisition to be 

aware of any Federal, State, Tribal, or 

territorial law or regulation that applies 

to that activity. 

Alternatives to the 2016 Interim Rule 

(83) Comment: Numerous 

commenters recommended health 

certification as an alternative to the 

injurious wildlife listing. For example, a 

commenter urged the Service to 

reconsider the listing of the 201 

salamander species, and instead to 

employ models proven effective by 

USDA’s Animal and Plant Health 

Inspection Service, such as utilization of 

certificates of veterinary inspection, preor post-import quarantine and 

treatment, import permits, and import 

restrictions based on risk assessments 

for given countries of origin. Another 

commenter urged the Service and other 

relevant agencies to work with its 

members to develop immediate 

measures to allow for preventive 

treatment and certification, without 

causing undue personal impacts. 

Response: While we do work 

collaboratively with the USDA and 

nongovernmental organizations on 

many invasive-species issues, the 

authority to list species as injurious 

wildlife under 18 U.S.C. 42 lies with the 

U.S. Department of the Interior. 

Although some countries may have the 

necessary skills to prepare a health 

certificate that salamanders are free of 

Bsal, not all exporting nations may have 

the necessary skills or resources. 

Scientists and diagnostic laboratories 

are also working to standardize 

laboratory protocols. Please see heading 

in the second interim rule on Ability To 

Prevent or Control the Spread of 

Pathogens or Parasites for more 

explanation. The Service will continue 

to seek opportunities to work with 

partners to ensure salamander 

conservation consistent with its mission 

but cannot commit to specific actions 

that do not fall under the scope of this 

rulemaking. 

(84) Comment: Related to other 

comments about establishing a 

certification system, the Service should 

consider establishing a permit from 

which the proceeds would help manage 

certification, testing, and conservation 

efforts and, therefore, could both help 

fund the program and make it more 

scientifically accurate. 

Response: While we are not 

establishing a certification system at this 

time under this rule, we acknowledge 

that the general statutory authority to 

charge fees for processing applications 

for permits and certificates is found in 

31 U.S.C. 9701, which states that 

services provided by Federal agencies 

are to be ‘‘self-sustaining to the extent 

possible.’’ Federal user-fee policy, as 

stated in Office of Management and 

Budget (OMB) Circular No. A–25 

Revised, requires Federal agencies to 

recoup the costs of Federal activities 

that provide ‘‘special benefits’’ to 

identifiable recipients. Permits are 

special services, authorizing identifiable 

recipients to engage in activities not 

otherwise authorized for the general 

public. Please also see our response to 

Comment PR31. 

(85) Comment: Chain pet stores 

should be prohibited from selling 

salamanders because it is too hard to 

regulate them. Only specialty pet places 

and breeders that have a permit should 

be allowed to sell salamanders. 

Response: Regardless of the business 

size or type, as explained in the final 

rule, all import and transport of 

injurious wildlife between the 

enumerated jurisdictions in the 

shipment clause of 18 U.S.C. 42, 

codified in Federal regulations at 50 

CFR 16.3, are prohibited for salamander 

species listed by this rule except by 

permit for authorized purposes. 

Otherwise regulating the sale of 

salamanders is not within the scope of 

this rulemaking. It is the responsibility 

of a person who may be engaged in 

salamander sales to be aware of any 

Federal, State, Tribal, or territorial law 

or regulation that applies to that 

activity. 

(86) Comment: A mandatory holding 

period for salamanders should be 

considered for transport across State 

lines. Studies show that if an animal is 

infected, it will die within a very short 

period. Only animals that test negative 

for Bsal should be allowed to be 

shipped. All animals in quarantine 

should also be treated to prevent 

infection and spread of Bsal, once 

reliable protocols are developed, as they 

have been for Bd (Pessier and 

Mendelson 2010). Quarantine efforts 

would facilitate both prevention of 

introduction and compliance. 

Response: Only lethally vulnerable 

species are expected to die in response 

to Bsal infection. Other species listed by 

this rule are also capable of carrying 

Bsal without lethal consequences and 

transmitting the fungus to native 

species. For these and other reasons 

discussed in this rule regarding 

certification and testing options, while 

research is ongoing, it is currently not 

feasible to establish such a system. The 

interstate prohibition has also been 

clarified as discussed in the second 

interim rule. 

(87) Comment: Consider instead a 

CITES import ban of all species of 

salamanders and newts, under the 

notion that CITES exists to protect 

endangered species. There are 35 

species of amphibians that would be at 

risk of being wiped out entirely if Bsal 


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becomes introduced into the United 

States. 

Response: CITES exists for a different 

purpose from the injurious wildlife 

provisions of the Lacey Act, and the 

purpose and use of CITES is beyond the 

scope of this rulemaking. 

(88) Comment: The Service has not 

acknowledged nonregulatory 

approaches. The pet industry has taken 

voluntary action to halt the import of 

known carriers. For example, PIJAC 

called for an immediate, temporary 

moratorium of Oriental fire-bellied newt 

and paddle-tailed newts on November 

20, 2015. 

Response: One of our alternatives 

(Alternative 1) involved taking no action 

on the Service’s part. This is our status 

quo. We would not list any species of 

salamanders as injurious under this 

alternative. We did not select this 

option because of the significant risk 

that Bsal poses to native species and 

other wildlife resources in the United 

States. We expect that significantly 

greater financial and natural resources 

losses will be incurred by us and our 

partners in having to manage and 

respond to Bsal if the fungus establishes 

and spreads in the United States than by 

taking action now to prevent its 

introduction. 

While we appreciate and support 

voluntary conservation efforts, we 

concluded that regulatory action was 

necessary to ensure compliance and 

protect native species. For example, the 

voluntary moratorium called for by 

PIJAC affected only two not-listed 

species, even though more have been 

identified as carriers (Martel et al. 2014). 

The species with the highest number of 

imports into the United States from 

2004 to 2014 was the Oriental firebellied newt. This species comprised 54 

percent of the total number of imported 

salamanders (USFWS OLE 2015). A 

review of LEMIS (Law Enforcement 

Management Information System) data 

in August 2016 shows that there were 

no shipments after November 20, 2015, 

for Oriental fire-bellied newts declared 

to the Service, except for 6 shipments 

totaling 539 live animals that occurred 

since the inception of the voluntary 

moratorium (all but 39 were before the 

rule took effect). This situation suggests 

that the rule is necessary because some 

importers, even if only a few, did not 

follow the voluntary moratorium and 

imported hundreds of specimens. 

However, since the 2016 interim rule 

took effect and as of the end of 2020, no 

Pachytriton spp. salamanders (not 

listed) have been imported, and we 

recognize and appreciate the role that 

the PIJAC moratorium likely played. 

(89) Comment: Although Alternative 3 

of the draft economic analysis, which 

declares 201 salamander species as 

injurious, is deemed most effective, the 

commenter suggests that Alternative 4, 

which declares all species of 

salamander as injurious, is necessary to 

fully prevent the spread of Bsal in the 

United States. Furthermore, the 

economic loss associated with 

Alternative 3 is estimated to be $10 

million, while the economic loss 

associated with Alternative 4 is $10.7 

million. The benefit of almost certainly 

preventing the spread of the fungus into 

the United States as provided by 

Alternative 4 far outweighs the marginal 

cost as compared to Alternative 3. 

Response: The expected increase in 

cost from Alternative 3 to Alternative 4 

was not considered in our 

determination about the injuriousness of 

the species. The Service determined that 

there was unknown risk from genera 

where no species have yet been tested 

for Bsal and, therefore, could not list 

those genera at this time. 

(90) Comment: We need more citizen 

scientists to help with salamander 

conservation. Many knowledgeable 

hobbyists are available to assist if asked. 

Response: We recognize that the 

public can play a critical role in 

conservation; however, this comment is 

outside the scope of this rulemaking. 

(91) Comment: Put more funding into 

Bsal research to find a cure, treatments, 

and other ways of reducing the risk. 

Response: We recognize the important 

contributions made by Bsal research; 

however, this comment is outside the 

scope of this rulemaking. 

(92) Comment: As new evidence 

becomes available and while Bsal 

remains undetected in the United 

States, the commenter would like to see 

a proposed rule with a comment period 

for native U.S. species, rather than an 

interim final rule, before these new 

listings go into effect. For nonnative 

species, however, we would support 

other interim final rules to further 

reduce the chances of introduction via 

the importation pathway. 

Response: This second interim rule is 

adding new nonnative and native 

species to the injurious list. See above 

in III. Final Rule to the 2016 Interim 

Rule in A. Background. Also, several 

native species are raised outside the 

United States and then imported into 

the country; this supports the Service’s 

decision to implement a nearly 

immediate effective date of 15 days for 

all species listed under the rule. See 

also response to Comment 24. 

(93) Comment: The costs to State fish 

and wildlife agencies to deal with pet 

salamanders that cannot be transported 

across State lines when the owner 

moves do not appear to have been 

evaluated and could place a significant 

burden on State agency staff that would 

be tasked with informing the public 

about the rules, working with rescues 

and zoos to provide rehoming 

opportunities, and law enforcement. 

The commenter would like to see a 

Service-administered education and 

outreach program that provides explicit 

instructions, and assistance, for pet 

owners to properly rehome or dispose of 

their salamanders. One commenter 

mentioned that the State of Florida has 

an Exotic Pet Amnesty Program in place 

that allows the public to surrender their 

regulated or unwanted exotic pets 

without penalty or cost. The commenter 

encourages continued Federal support 

of this program as an integral part of 

managing risks of nonnative 

introductions. 

Response: The interstate prohibition 

was clarified by a court decision in 2017 

as explained in the final rule, so the 

costs for transporting across State lines 

between States within the continental 

United States is not an issue now, 

unless regulated by other State or 

Federal laws. We share concerns about 

the irresponsible re-homing and 

disposal of pet salamanders into the 

wild and are working with partners, 

including the industry, to help ensure 

that release does not occur. The Service 

has been a partner with the State of 

Florida’s Exotic Pet Amnesty Program 

and will continue to work with other 

partners to help encourage the public 

not to release animals that they own 

into the wild. The Service does not have 

the funds necessary to implement a 

national amnesty and rehoming 

program. 

(94) Comment: In the past, increased 

restrictions on species already in 

widespread possession (personal and 

commercial) have been accompanied by 

additional releases (such as walking 

catfishes, snakeheads). The commenter 

recommends consideration of regulatory 

approaches with the flexibility to 

accommodate existing ownership. 

Further Federal restrictions, without 

this ‘‘grandfathering’’ approach for 

current pet owners, may lead to an 

increase in the rates of release. 

Response: The commenter offers no 

proof that releases have been caused by 

the new Federal regulation. The 

injurious wildlife provisions of the 

Lacey Act do not prohibit continued 

ownership of injurious wildlife that 

members of the public own at the time 

of listing. Under the injurious wildlife 

provisions of the Lacey Act, the Service 

is not authorized to grandfather in 

existing salamander owners as exempt 


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from subsequent activities that are 

prohibited with injurious wildlife, 

including import or transport between 

the enumerated jurisdictions. 

(95) Comment: Prohibit the use of 

amphibians as fishing bait. It has been 

shown that using animals, such as tiger 

salamanders, as fishing bait has led to 

species introductions (posing a major 

threat to California tiger salamanders) 

and the spread of disease, particularly 

Bd and ranaviruses. If Bsal ever enters 

the United States, it is far more likely to 

be spread through bait shops and 

fishermen than from hobbyists shipping 

to one another. Even if studies have 

shown tiger salamanders are unlikely to 

carry Bsal, the practice has already been 

shown to have spread other diseases, 

and other, more susceptible species may 

be used. 

Response: The request is beyond the 

scope of this rulemaking. The Service, 

under the injurious wildlife provisions 

of 18 U.S.C. 42, is not authorized to 

prohibit amphibians for use as fishing 

bait, unless they are imported, 

transported between the enumerated 

jurisdictions, or subject to injurious 

wildlife permits. We also note that the 

Service’s State partners regulate fishing 

activities within their States and can, 

and often do, regulate use of amphibians 

for fishing bait. 

(96) Comment: Include a clause that if 

a North American species is determined 

to be a carrier or lethally infected, it will 

immediately be included in the 

prohibition, and any species screened 

and determined to be insensitive and 

not capable of carrying Bsal will be 

removed from the list in a timely 

manner. 

Response: The Service does not have 

authority to include or remove species 

on the injurious wildlife list without 

evidence regarding whether the wildlife 

is injurious to the interests protected 

under the Lacey Act. The determination 

of injuriousness is based on defensible 

scientific evidence. This rule does not 

preclude the ability to take additional 

regulatory actions if new information 

emerges. If a species is found to be 

incapable of carrying Bsal under all 

conditions, we may consider its removal 

from the injurious list by conducting an 

evaluation and promulgating a rule. 

Likewise, if a species is found to be a 

carrier of Bsal, we may consider its 

addition or the addition of its genus to 

the injurious list through this same 

regulatory process. 

(97) Comment: Continue exploring a 

clean-trade program for future emerging 

infectious diseases. As indicated in 

previous correspondence with the 

Service, the commenter has consistently 

supported the concept of a clean-trade 

program for salamanders imported into 

the United States rather than restricting 

interstate movement of salamanders at 

this point. The commenter appreciates 

that the current situation makes 

executing such a program difficult, if 

not impossible; however, the 

commenter hopes that the Service will 

continue exploring options for such a 

program in the future for other emerging 

infectious diseases that are likely to 

impact U.S. wildlife species. 

Response: The interstate prohibition 

has been clarified as explained in the 

final rule. Please also see heading in the 

second interim rule on Ability To 

Prevent or Control the Spread of 

Pathogens or Parasites. The Service will 

consider other options as opportunities 

arise. 

(98) Comment: The Service cites 

inadequate agency resources to conduct 

inspections and expenses associated 

with testing as additional reasons 

supporting its finding that there are not 

less restrictive means to prevent Bsal 

than those selected for the 2016 interim 

rule. To the extent that those expenses 

and hardships fall upon the owners of 

salamanders, a commenter would like to 

work with the Federal Government in 

developing safe, practical procedures. 

To the extent that those burdens fall 

upon the agency, the Service must not 

discriminate between regulation of 

salamanders in the pet trade and other 

species for which it has dedicated 

resources to developing satisfactory 

testing protocols. 

Response: The Service welcomes and 

encourages engagement by a myriad of 

entities that can develop the science and 

help better manage wildlife pathogens 

entering, becoming established in, and 

spreading in the United States. No safe, 

effective alternatives have yet been 

presented to us. The fungus that affects 

the salamanders was discovered in 

2013, much more recently than the 

pathogens infecting salmonids for 

which the Service has testing protocols. 

Much research needs to be done on the 

tremendous diversity of salamanders 

and their in situ environmental 

conditions to find an equitable, reliable, 

economical test as well as testing 

facilities in other countries. 

Other Issues 

(99) Comment: The Wildlife Society 

recommends the development of new 

comprehensive legislation to address 

the complexities of emerging wildlife 

diseases that encourages investment, 

increases professional capacity, focuses 

on collaborative prevention, and uses a 

multidisciplinary approach to better 

understand the interaction and 

transmission mechanisms of wildlife 

pathogens. 

Response: The comment is outside the 

scope of this rulemaking. 

(100) Comment: Since no method of 

pathogen control is likely to reduce risk 

of invasion by 100 percent, it is equally 

important to invest in proactive 

monitoring for Bsal emergence within 

the United States. In August 2015, the 

commenter launched a citizen science 

project on iNaturalist for people to 

report sightings of dead or diseased 

salamanders. The commenter would be 

happy to work together with the Service 

more quickly to identify and respond to 

potential sites of Bsal emergence. More 

information about the project ‘‘Saving 

Salamanders with Citizen Science’’ can 

be found at: http://www.inaturalist.org/ 

projects/saving-salamanders-withcitizen-science. 

Response: We shared this feedback 

with our National Wildlife Refuge 

System’s Inventory and Monitoring 

Program. The Service is also helping the 

National Bsal Task Force and PARC to 

develop the protocols to monitor for 

Bsal’s introduction and to allow for 

rapid response if it is identified in the 

United States. 

(101) Comment: The commenter 

requests increased communication and 

education efforts around the Bsal rule. 

There is still a significant amount of 

confusion around the reasoning behind 

the scope of the action taken in the rule, 

including, but not limited to, why 

certain species were chosen and why 

interstate commerce was included. 

Addressing these concerns through a 

coordinated education and 

communication initiative would likely 

help garner further support for the 

implementation of the rule. Many 

groups, such as caudata.org, the 

National Bsal Task Force, and Partners 

in Amphibian and Reptile Conservation 

(PARC) would likely be able to play a 

role in helping to disseminate this 

information. 

Response: The interstate prohibition 

has been clarified as explained in the 

final rule. The Service is a partner in 

PARC and a member of the Bsal Task 

Force and appreciates the need to better 

address the communication needs 

associated with the rule. We have also 

been providing additional information 

through our website to assist 

stakeholders in understanding the need 

for the rule and clarifying the permit 

process. 

(102) Comment: The rule is being 

exploited by animal rights organizations 

who do not represent the majority of 

views of U.S. citizens. This rule was 

formulated in part due to a petition by 

the ‘‘Save the Frogs’’ organization. 


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Response: The Service received the 

petition from the Center for Biological 

Diversity and Save the Frogs! in midMay 2015. However, we began 

discussions on what action to take in 

October 2014 and had already begun the 

regulatory process several months 

before we received the petition. Letters 

to the Service Director from such 

agencies as AFWA urged the Service to 

take action to prevent the fungus from 

entering the United States, and we took 

action as soon as we were able to make 

a determination based on defensible 

scientific evidence and comply with 

applicable rulemaking requirements to 

promulgate injurious wildlife listings 

under the Lacey Act within our 

available resources. 

C. Affirmation of the 2016 Interim Rule 

After careful consideration of the 

comments received and information 

presented, we are affirming our 2016 

listing of the 20 genera of salamanders 

that the 2016 interim rule added to the 

lists of injurious wildlife in 50 CFR part 

16 (81 FR 1534, January 13, 2016). All 

species in the 20 genera continue to be 

listed as injurious wildlife. The 

defensible scientific evidence continues 

to indicate that the importation of these 

genera poses significant risks of 

introducing Bsal into the United States, 

and none of the inputs received in 

response to the 2016 interim rule have 

changed this determination. Therefore, 

with this document, we affirm the 

addition of the following genera to 50 

CFR 16.14: Chioglossa, Cynops, 

Euproctus, Hydromantes, Hynobius, 

Ichthyosaura, Lissotriton, Neurergus, 

Notophthalmus, Onychodactylus, 

Paramesotriton, Plethodon, Pleurodeles, 

Salamandra, Salamandrella, 

Salamandrina, Siren, Taricha, Triturus, 

and Tylototriton. Because we consider 

rulemaking on the 2016 interim rule to 

end with the publication of this 

document affirming the 2016 interim 

rule, we are not soliciting comments 

regarding the genera listed in this final 

rule. 

D. Required Determinations 

We hereby affirm our responses to the 

following determinations required of the 

Federal rulemaking process as 

published in the January 13, 2016, 

interim rule (81 FR 1534): 

• Executive Orders 12630, 12866, 

12988, 13132, 13175, 13211, and 13563; 

• Regulatory Flexibility Act and 

Congressional Review Act (5 U.S.C. 601 

et seq. and 804(2)) (except a decrease in 

the economic effect on U.S. industries 

has occurred due to the clarification of 

the interstate transport prohibition); 

• Unfunded Mandates Reform Act (2 

U.S.C. 1501 et seq.); 

• Paperwork Reduction Act of 1995 

(44 U.S.C. 3501 et seq.); 

• National Environmental Policy Act 

(42 U.S.C. 4321 et seq.); and 

• Government-to-Government 

Relations with Native American Tribal 

Governments (59 FR 22951 and 512 DM 

2). 

IV. Second Interim Rule 

Summary of Changes to the 2016 

Interim Rule 

For the injurious wildlife evaluation 

in this second interim rule, in addition 

to information used for the 2016 interim 

rule, we considered: (1) comments and 

new information from the public 

comment period for the 2016 interim 

rule, (2) comments on the 2016 interim 

rule from three peer reviewers, and (3) 

new information acquired by the 

Service after the 2016 interim rule 

published. This new information was 

used to update the science about Bsal 

and determine if any additional genera 

of salamanders should be added to the 

list according to the criteria laid out in 

the 2016 interim rule. 

This second interim rule incorporates 

into 50 CFR 16.14 the clarifications and 

changes to the 2016 interim rule based 

on comments we received that are 

discussed above in the final rule under 

B. Summary of Comments Received on 

the 2016 Interim Rule. This is because 

only one revision of 50 CFR 16.14 will 

be made from both rules and will 

include the clarifications from the final 

rule and the new genera and 

clarifications from this second interim 

rule. 

We are clarifying, in response to 

public and peer-review comments, what 

is and is not injurious on a cellular or 

molecular level based upon chemical 

preservation or other methods that will 

kill the fungus. Unpreserved swabs are 

injurious; however, preserving swabs, 

such as by using 70 percent (or higher) 

ethanol for at least 1 minute (Van Rooij 

et al. 2017), renders the fungus unviable 

and, therefore, preserved swabs are not 

injurious and are excluded from the 

prohibitions. In addition, purified 

extracted genetic material of 

salamanders (salamander DNA or RNA) 

is unable to cause pathogen 

transmission; therefore, it is not 

injurious. Swabs collected for molecular 

biology applications should be 

preserved by using a higher ethanol 

concentration (95–99 percent), which is 

adequate for both the molecular 

preservation of DNA for testing and 

denaturing the proteins on the surface of 

fungi, rendering them unviable and thus 

not injurious (Marquina et al. 2021). 

We provide evidence here that 

specimens that are chemically preserved 

to deactivate any live Bsal and purified 

extracted genetic material are not 

considered injurious. The Service has 

concluded that there is a low risk of 

transmission of Bsal to native species 

from eggs and gametes, preserved 

specimens, and purified extracted 

genetic material, which is consistent 

with the intent of what is not injurious 

in the 2016 interim rule. However, all 

other parts, such as unpreserved 

salamander tissues, fluids, and cells 

carried on swabs and on or in other 

media, will continue to be regulated 

under the listing. Specimens that are 

frozen are also included in the listing. 

The Service reviewed research that 

has published since the 2016 interim 

rule took effect and is adding 16 genera 

of salamanders to the 20 already listed 

in 50 CFR 16.14. This action adds 

approximately 164 species in this 

second interim rule to the previously 

listed 201 species. The genera are added 

under the same criteria that were used 

for the original 20 genera in the 2016 

interim rule. However, the scientific 

community has made changes to 

salamander taxonomy within the 20 

genera in the 2016 interim rule. Thus, 

the number of species that we identified 

in those genera increased from 201 to 

262 species as of May 1, 2023. The 

combined total is 36 genera with 

approximately 426 species. We note that 

taxonomic changes within each genus 

may occur at any time for such reasons 

as new species discovered, subspecies 

elevated to full species, species split 

into two species, and other 

modifications resulting from genomic 

testing. All species subsequently 

scientifically added to the genera that 

we are listing are also heretofore 

considered listed species. For this 

reason, we are not enumerating all of 

the known salamander species in each 

of the 36 listed genera in 50 CFR 16.14. 

The statute (18 U.S.C. 42(a)(1)) refers 

to ‘‘the offspring or eggs of any of the 

foregoing’’ as being injurious. Therefore, 

we are clarifying that hybrids of species 

in any listed genus, including offspring 

from a listed and a nonlisted parent, are 

injurious. 

In response to the D.C. Circuit Court 

of Appeals Decision in United States 

Association of Reptile Keepers, Inc. v. 

Zinke, 852 F.3d 1131 (D.C. Cir. 2017), 

the prohibition on interstate transport in 

the 2016 interim rule has been 

modified. The D.C. Circuit Court of 

Appeals held that 18 U.S.C. 42(a) does 

not prohibit transport of injurious 

wildlife between States within the 


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continental United States. Therefore, 

this interim rule clarifies that 50 CFR 

16.3 does not prohibit interstate 

transport between States within the 

continental United States. This means 

that transportation of injurious wildlife 

between the 49 States within the 

continental United States (the 

contiguous 48 States and Alaska, 

provided no international borders are 

crossed) is not prohibited by the statute 

or injurious wildlife regulations, unless 

that movement of the wildlife is 

restricted due to conditions associated 

with issued injurious wildlife permits. 

This change took effect as of April 7, 

2017. However, import of injurious 

wildlife into the United States remains 

prohibited. In addition, transport of 

injurious wildlife between the listed 

jurisdictions in the shipment clause (the 

continental United States, the District of 

Columbia, Hawaii, the Commonwealth 

of Puerto Rico, and any territory or 

possession of the United States) remains 

prohibited. As before, injurious species 

may not transit into or out of the District 

of Columbia. Also, injurious wildlife 

permits under 50 CFR 16.22 continue to 

be required to import injurious wildlife 

and to transport injurious wildlife 

between the listed jurisdictions for 

zoological, educational, medical, and 

scientific purposes; movements within 

the continental United States may be 

subject to conditions from an injurious 

wildlife permit. 

Additionally, injurious wildlife 

unlawfully imported into the United 

States or transported between the 

enumerated jurisdictions is still 

unlawful to transport, including within 

the continental United States. Under the 

Lacey Act Amendments of 1981, 16 

U.S.C. 3372(a)(1), it is unlawful for any 

person to import, export, transport, sell, 

receive, acquire, or purchase any 

wildlife transported in violation of any 

law of the United States. This includes 

transport of any injurious wildlife 

imported into the United States or 

transported between the enumerated 

jurisdictions in violation of 18 U.S.C. 

42(a). 

Need for an Interim Rule 

Rulemaking under 18 U.S.C. 42 is 

governed by the Administrative 

Procedure Act (APA) (5 U.S.C. 551 et 

seq.). The process of issuing a proposed 

rule, providing the opportunity for 

public comment, and completing a final 

rule can take a significant amount of 

time to complete. During that time, the 

species proposed for listing are still 

allowed to be imported and transported, 

offering increased opportunities for 

introduction, establishment, and harm. 

Under section 553(b)(3)(B) of the APA, 

however, a proposed rule is not required 

when the agency for good cause finds 

(and incorporates the finding and a brief 

statement of reasons therefor in the 

rules issued) that notice and public 

procedure thereon are impracticable, 

unnecessary, or contrary to the public 

interest. There is good cause to forgo 

notice and public comment on a 

proposed rule in this case and instead 

take immediate action in the form of an 

interim rule to help prevent the Bsal 

fungus from being introduced, 

established, or spread in the United 

States. Providing notice and public 

comment prior to implementing the 

injurious wildlife prohibitions would be 

contrary to the public interest because 

of the need to take immediate action 

due to the significant risk from Bsal. Not 

only could the fungus cause the 

devastation of some populations of 

native salamanders critical to ecosystem 

health, but it could also cause mortality 

if it spreads in the salamander pet trade. 

For these reasons, we find good cause in 

accordance with 5 U.S.C. 553(d)(3) to 

make the second interim rule effective 

15 days after the date of publication in 

the Federal Register. 

This second interim rule is the result 

of peer-reviewed, scientific information 

published since the publication of the 

2016 interim rule. At the time the 2016 

interim rule published, there was very 

little information on the newly 

described chytrid fungus species 

affecting salamanders (discovered in 

2013). We used defensible scientific 

evidence to quickly stop the importation 

of the host species (salamanders) of the 

fungus. However, after the 2016 interim 

rule published, many research 

institutions realized the need for more 

research, both on the novel fungus and 

on the effect on and variety of host 

amphibians, to assess the validity of 

previous studies and determine other 

potential carriers. That body of research, 

done primarily in the United States, 

Europe, and Asia, has taken years to 

develop and put through the peerreview and journal-publication 

processes. 

We have now compiled a more 

comprehensive picture of the fungus, 

including 16 more genera that we 

determined are injurious (using the 

same criteria as in the 2016 interim 

rule). We still have the opportunity to 

prevent the contagious lethal fungus 

from being introduced into the United 

States on salamanders in trade, hence 

the need for the second interim rule 

with the new high-risk species. Some of 

the new species are in the pet trade, and 

a proposed rule would give the public 

the counterproductive opportunity to 

rush to import the proposed species 

prior to the regulation. We note that a 

shipment of 24 live fire salamanders 

(Salamandra salamandra) was imported 

into Los Angeles on January 26, 2016, 

which was 2 days before the listing took 

effect (USFWS OLE 2021), although we 

do not know that the shipment was 

intentionally shipped to preemptively 

avoid the 2016 interim rule’s effective 

date. Fire salamanders are the species 

that brought this lethal fungus to the 

attention of scientists, and the shipment 

was exported from Germany, where the 

fungus had been detected in 2015 

(Schultz et al. 2020), making the 

potential for Bsal introduction from this 

shipment a genuine threat. Fortunately, 

there is no evidence that those imported 

salamanders carried Bsal. 

Purpose of Listing as Injurious 

The purpose of listing the live 

specimens, dead specimens, hybrids, 

and parts (but not eggs, gametes, 

preserved specimens or parts (including 

tissue), and purified extracted genetic 

material) of 16 genera of salamanders as 

injurious wildlife is to prevent the 

accidental or intentional introduction of 

salamanders that are expected to serve 

as carriers of Batrachochytrium 

salamandrivorans (hereafter, Bsal), a 

fungus that poses a risk to native species 

of salamanders, into the United States. 

The genera are all from the order 

Caudata and are commonly referred to 

as salamanders and newts (hereafter, 

salamanders). If Bsal is introduced into 

wild populations of native salamanders, 

we expect it to cause significant harm to 

wildlife and the wildlife resources of 

the United States. 

Under 18 U.S.C. 42(a), the Service, 

through the Secretary of the Interior, 

may prescribe by regulation any wild 

mammals, wild birds, fishes, mollusks, 

crustaceans, amphibians, reptiles, or the 

offspring or eggs of any of the foregoing 

found to be injurious to human beings, 

to the interests of agriculture, 

horticulture, forestry, or to wildlife or 

the wildlife resources of the United 

States. Salamanders are amphibians, 

and the Service has the authority to list 

amphibians when we find that they are 

injurious to one or more of the statutory 

interests. We may list species before 

they are introduced into the United 

States and have the opportunity to harm 

interests of the United States as 

enumerated under 18 U.S.C. 42. 

We have determined that salamanders 

that potentially carry Bsal are injurious 

to wildlife and wildlife resources of the 

United States. With this second interim 

rule, we are attempting to prevent the 

introduction and subsequent 

establishment and spread of the 

salamander chytrid fungus, Bsal, which 


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is a pathogen capable of causing 

significant harm to native salamander 

species and their ecosystems. As 

described below under Role of 

Salamanders in the Ecosystem, the 

benefits that these native salamander 

species provide to ecosystems in 

ensuring ecosystem health and stability, 

and, in turn, the ecosystem services that 

benefit people, are significant. 

As of the publication of the 2016 

interim rule, Martel et al. (2014) and 

Cunningham et al. (2015) identified 

some of the salamander species that can 

carry Bsal and are at risk from infection. 

The researchers tested a limited number 

of the approximately 689 (currently 804) 

known species of salamanders that exist 

worldwide and found that not every 

species was negatively affected by the 

fungus, as determined by standard 

detection methods. However, the results 

clearly indicated a severe threat for 

many species of salamanders that will 

be negatively affected by this pathogen 

and others that could carry the fungus 

to the vulnerable species. Research 

showed that some tested species that are 

native to the United States were found 

to be lethally vulnerable to the fungus. 

Such an emerging infectious disease of 

fungal origin can cause a significant loss 

in biodiversity and ecosystem services 

(Fisher et al. 2012). Bsal research results 

and concerns about emerging infectious 

disease, as described by Spitzen-van der 

Sluijs et al. (2013), Martel et al. (2013), 

and Martel et al. (2014), generated a 

strong response from academia, industry 

groups, and conservation and other 

organizations who wrote to the Service 

seeking quick and decisive action to 

ensure that Bsal does not have a similar 

impact on salamander populations that 

Batrachochytrium dendrobatidis (Bd) 

has had on frogs (see the 2016 interim 

rule). In early November 2014, the 

Service initiated a review to determine 

whether salamanders capable of 

carrying Bsal should be listed as 

injurious. 

Martel et al. (2014, and others later) 

used several methods to determine 

vulnerability to Bsal-caused disease of 

some salamander species but do not 

have a category for the status as a 

carrier. While the vulnerability of native 

species is of great concern to the 

Service, the 2016 interim rule was 

primarily concerned with the ability of 

viable Bsal spores to remain on 

salamander species or their parts, thus 

introducing and spreading the fungus to 

the United States, causing 

chytridiomycosis disease outbreaks in 

native salamander populations. We 

reviewed the literature and based our 

criteria for determining carrier status of 

genera on whether a species was found, 

as determined by microscopic analysis 

of preserved tissue specimens 

(histology), qPCR (quantitative 

polymerase chain reaction), or other 

confirmatory approach to harbor viable 

spores. 

We also looked at challenge studies, 

where a salamander that is free of Bsal 

as determined by initial pathogenspecific qPCR, is then inoculated with 

Bsal spores. A follow-up swab for qPCR 

is done at a specified period of days 

later to see if the spores caused disease 

according to field observations or 

histology or did not cause disease but 

was able to invade the skin of that 

species long enough to move or transmit 

the fungus to other salamanders, as 

confirmed by histology. We also looked 

at surveillance studies of swabs of wild 

and captive salamanders, where 

presence or absence of Bsal was 

determined by qPCR; negative results 

were not evidence for being classified as 

noncarrier whereas positive results were 

classified as carriers. 

Regardless of the vulnerability of a 

species or the ability to manifest 

disease, if the species is a carrier, we 

consider that genus to be listable as 

injurious. However, if there is 

conclusive countervailing evidence that 

at least one species in that genus is not 

a carrier, as shown by histology, then 

we do not list the genus. Case definition 

and diagnostic criteria are described in 

White et al. (2016). 

The 2016 interim rule effectively 

reduced import volume of targeted 

species, but new research on species 

susceptibility suggests the list of 

regulated species was incomplete 

regarding Bsal reservoir species (Grear 

et al. 2021). Since the publication of the 

2016 interim rule, additional research 

has provided additional evidence of the 

diversity of species and genera affected 

by Bsal or determined to be carriers (for 

example, Yuan et al. 2018, Carter et al. 

2020, Barnhart et al. 2020, Gray et al. 

2023). Based on the Service’s genuslevel carrier extrapolation from data 

obtained from the aforementioned 

publications, and because Bsal has not 

been found in natural environments in 

the United States (Waddle et al. 2020), 

the opportunity still exists to prevent 

the introduction of Bsal by adding new 

genera of salamanders to the injurious 

list. In 2017, following the 2016 interim 

rule and Canada’s temporary import ban 

of all living or dead salamanders, eggs, 

sperm, tissue cultures, and embryos 

(made permanent in 2018; Environment 

and Climate Change (ECCC) 2018), we 

received a letter from the American 

Society of Ichthyologists and 

Herpetologists (ASIH 2017) requesting 

the Service prohibit all salamander 

imports into the United States. For 

reasons explained herein, we are not 

listing all salamanders, but we are 

adding more genera, which we 

suggested in the 2016 interim rule was 

a possibility. We specifically solicited 

comment on whether there is there any 

evidence suggesting that additional 

species are carriers of Bsal and should 

be listed by this rule, and if so, what 

species. 

We reviewed Bsal risk in conjunction 

with those salamander species known or 

suspected to carry the fungus utilizing 

injurious wildlife evaluation criteria, 

described in more detail as part of this 

interim rule in G. Factors That 

Contribute to Injuriousness of 

Salamanders. These criteria were 

previously developed by the Service to 

evaluate whether a species qualifies as 

injurious under 18 U.S.C. 42. The 

resulting analyses form the basis for the 

Service’s regulatory decisions regarding 

injurious wildlife-species listings. This 

rule finds that Bsal is a significant threat 

to the wildlife and wildlife resources of 

the United States and lists 16 genera of 

salamanders that we have determined to 

be injurious because they are likely 

carriers of Bsal. 

Multiple factors confirm that Bsal can 

be introduced, become established, and 

spread, thereby causing substantial 

damage and harm in the United States 

(Spitzen-van der Sluijs et al. 2013; 

Martel et al. 2014; Cunningham et al. 

2015; Chytridcrisis 2015b). Specifically, 

these factors include: (1) the discovery 

of the newly emerging fungus Bsal in 

the Netherlands and the associated 

deleterious effects to native salamanders 

(ibid.); (2) its subsequent spread in the 

wild to Germany and Belgium (Spitzenvan der Sluijs et al. 2016) and Spain 

(Lastra Gonza ́laz et al. 2019; Martel et al. 

2020); (3) the appearance in captive 

collections in the United Kingdom, 

Germany, and Spain (Spitzen-van der 

Sluijs et al. 2016; Thumsova et al. 2021); 

and (4) laboratory research (numerous 

papers cited in this rule). The United 

States leads all other countries in the 

number of native salamander species; 9 

of the 10 families of salamanders 

worldwide are found in the United 

States (AmphibiaWeb 2023a). Based on 

scientific evidence as of publication of 

the 2016 interim rule, we knew that the 

fungus is lethal to at least two 

salamander species native to the United 

States (eastern newt Notophthalmus 

viridescens in the Eastern States and 

rough-skinned newt Taricha granulosa 

along the Pacific coast). 

Of the 221 native U.S. species known 

as of the preparation of this second 

interim rule (AmphibiaWeb 2023a), and 

including both rules in this document, 


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we have determined that 13 genera with 

164 species may be carriers, and 9 

species are lethally vulnerable. Most of 

the remaining 10 genera (with 57 

species) have not been scientifically 

tested, with a few that have had testing 

that was not conclusive; these may also 

be found to be carriers eventually. 

While the Service’s greatest concerns 

are for species that are likely to die from 

Bsal, salamander species known to be 

tolerant of, or susceptible to, Bsal 

infection under experimental conditions 

may also develop clinical disease or 

experience increased severity of disease 

in the wild. These species may be Bsal 

carriers and are concerning because 

their long lifespans increase their 

likelihood of spreading the fungal 

spores and serving as Bsal reservoirs 

(Gray et al. 2023). A disease reservoir 

may be defined as ‘‘a passive host or 

carrier that harbors pathogenic 

organisms without injury to itself and 

serves as a source from which other 

individuals can be infected’’ (in Laking 

et al. 2017). Nonlethal infection in 

salamanders may have other negative 

effects, such as slowing their growth 

(Barnhart et al. 2020). Bsal infections 

have been found to increase in severity 

as animals are exposed to additional 

stressors in the wild, including other 

amphibian diseases (Wobeser 2007; 

Kerby et al. 2011; Kiesecker 2011; Longo 

et al. 2019; McDonald et al. 2020). 

Experience with the introduction of 

Bsal into the Netherlands and associated 

deleterious effects to native 

salamanders, along with laboratory 

research, confirm that Bsal can be 

introduced, become established, spread, 

and cause substantial and immediate 

harm in the United States (Spitzen-van 

der Sluijs et al. 2013; Martel et al. 2014; 

Cunningham et al. 2015; Chytridcrisis 

2015b). The United States leads all other 

countries in salamander diversity 

(PARC 2014). Based on scientific 

evidence, we know that the fungus is 

lethal to at least nine salamander 

species native to the United States. 

While the Service’s greatest concern 

will be for species that are lethally 

vulnerable to Bsal, salamander species 

known to be tolerant of or susceptible to 

Bsal infection under experimental 

conditions may also develop clinical 

disease or increased severity of disease, 

respectively, when infection is 

combined with additional stressors in 

the wild, as has been found for other 

diseases, including those in amphibians 

(Wobeser 2007; Kerby et al. 2011; 

Kiesecker 2011). 

In the United States, Bsal has either 

not been introduced, has been 

introduced but has failed to become 

established, or is present but has not 

been positively detected. Although we 

do not have any conclusive evidence 

showing that introductions have 

occurred, history from other pathogens 

similar to Bsal, such as Bd, suggests that 

the fungus is likely to spread quickly 

throughout the United States if it is not 

prevented from being introduced. 

Moreover, efforts to control or eradicate 

introduced or established invasive 

species and manage the costs they incur 

to society are generally less effective 

and more expensive and difficult than 

efforts that prevent establishment 

(Leung et al. 2002; Finnoff et al. 2007). 

Prevention of invasive species is 

typically the most cost-effective 

measure to avoid the damage that such 

species cause (Leung et al. 2002; Lodge 

et al. 2006; Keller and Springborn 2014). 

As noted in the 2016–2018 National 

Invasive Species Management Plan, 

preventing the introduction of 

potentially harmful organisms is not 

only the first line of defense for 

minimizing the spread and impact of 

invasive species, it is also the most costeffective strategy; science-based risk 

analyses are used to inform regulations 

that can prohibit the entry of certain 

nonnative organisms at jurisdictional 

borders (National Invasive Species 

Council 2016). Invasive species 

prevention is a priority of the 

Department of the Interior (2021). 

If Bsal has unknowingly been 

introduced but failed to establish in the 

United States for unknown reasons, it is 

still important to act now because 

additional introductions increase the 

likelihood of establishment and harm. 

As more salamanders that can carry Bsal 

are imported into the United States, the 

probability increases that one or more of 

those salamanders, through a 

phenomenon called propagule pressure 

or ‘‘introduction effort,’’ described in 

Lockwood et al. (2005) as a measure of 

the number of nonnative individuals 

released into a region, will give Bsal the 

opportunity to establish in the United 

States and spread. The 2016 interim rule 

significantly reduced the number of 

salamanders being imported by about 95 

percent (average per year for the period 

2016–2020) from the 6 years before 

publication of the 2016 interim rule. 

The salamander species listed by this 

second interim rule follow the same 

criteria as for the 20 genera in the final 

rule to the 2016 interim rule and are 

those found within genera for which we 

have evidence that at least one species 

in that genus is a carrier of Bsal with no 

countervailing conclusive evidence that 

other species in that genus are not 

carriers. We describe our rationale for 

this course of action below under 

Vulnerability and Carrier Status. Our 

decision making for the final rule to the 

2016 interim rule and the second 

interim rule included the following 

considerations: All 20 genera of 

salamanders in the final rule to the 2016 

interim rule, plus any new species 

identified within the genera listed by 

this second interim rule, are found to be 

injurious because suitable climate exists 

in parts of the United States to support 

Bsal; even if a salamander listed by this 

second interim rule could not establish 

a population in the wild, an infected 

captive salamander (or the water and 

soil in which it came into contact) 

released into the environment can 

transmit Bsal to native populations; Bsal 

is capable of causing extensive damage 

to wildlife and wildlife resources, 

including federally endangered and 

threatened species; eradicating Bsal 

would be extremely difficult once 

introduced and established; and 

controlling Bsal in wild salamanders is 

not practical. 

Listing the salamanders as injurious 

will help keep Bsal out of the United 

States by preventing the importation of 

salamanders capable of carrying the 

fungus and serving as the vector of 

introduction into U.S. ecosystems, 

thereby causing injurious effects 

consistent with 18 U.S.C. 42. Bsal is not 

known to be present in U.S. ecosystems 

(Waddle et al. 2020). Given the expected 

consequences that the introduction of 

Bsal would have to wildlife and wildlife 

resources of the United States, we are 

listing species that we have determined 

to be injurious. This second interim rule 

lists some species that are currently in 

U.S. trade as well as some that are not. 

We have the authority to list species as 

injurious even if they are not currently 

in trade or known to exist in the United 

States. This regulation is not a ban on 

possessing or selling any of the species. 

The import and transport between the 

enumerated jurisdictions in the 

shipment clause in 18 U.S.C. 42 (the 

continental United States, the District of 

Columbia, Hawaii, the Commonwealth 

of Puerto Rico, and any territory or 

possession of the United States), 

codified in Federal regulations at 50 

CFR 16.3, of any of the listed species is 

prohibited. The provisions of 18 U.S.C. 

42(a) do not prohibit any person who 

owns one of the listed species at the 

time of listing from continuing to 

possess the salamander or engaging in 

transport and other activities within the 

enumerated jurisdictions, unless such 

movement of the wildlife is restricted 

due to conditions associated with issued 

injurious wildlife permits. Those 

activities may be regulated by other 

Federal, State, Tribal, or territorial law. 


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It is the responsibility of a person who 

may be engaged in transport or use of 

injurious wildlife to be aware of any 

Federal, State, Tribal, or territorial law 

or regulation that applies to that 

activity. 

The salamander species listed by this 

second interim rule are those, in 

addition to the species listed in the final 

rule to the 2016 interim rule, that are 

found within genera for which we have 

evidence that at least one species in that 

genus is a carrier of Bsal with no 

conclusive countervailing evidence that 

other species in that genus are not 

carriers. We describe our rationale for 

this course of action below under 

Vulnerability and Carrier Status. Our 

decision making also included the 

following considerations: 

• All 16 genera of salamanders, plus 

any new species identified within the 

genera listed by this second interim 

rule, are found to be injurious because 

suitable climate exists in parts of the 

United States to support Bsal; 

• Even if a salamander listed by this 

second interim rule could not establish 

a population in the wild, a carrier 

salamander that was released from 

captivity (or the water and soil in which 

it came into contact) can transmit Bsal 

spores to native populations; 

• Bsal is capable of causing extensive 

injury to wildlife, including federally 

endangered and threatened salamander 

species; 

• No method is known to eradicate 

Bsal in the environment once it is 

introduced and established; and 

• Controlling Bsal is not practical in 

free-ranging salamanders carrying the 

fungus. 

We clarify what is considered a 

salamander part that is not injurious 

and that would not need an injurious 

wildlife permit (partially adapted from 

WOAH 2021a): 

• Heat-sterilized hermetically sealed 

amphibian products, that is, a heat 

treatment at 121 °C for at least 3.6 

minutes (or any time or temperature 

equivalent that has been demonstrated 

to inactivate Bsal) (WOAH 2021a); 

• cooked amphibian products that 

have been subjected to heat treatment at 

100 °C for at least 1 minute (or any time 

or temperature equivalent that has been 

demonstrated to inactivate Bsal) 

(WOAH 2021a); 

• pasteurized amphibian products 

that have been subjected to heat 

treatment at 90 °C for at least 10 minutes 

(or any time or temperature equivalent 

that has been demonstrated to inactivate 

Bsal) (WOAH 2021a); 

• mechanically dried amphibian 

products and skin leather (that is, a heat 

treatment at 100 °C for at least 30 

minutes or any time or temperature 

equivalent that has been demonstrated 

to inactivate Bsal) (WOAH 2021a); and 

• chemical treatment of amphibian 

skin leather that inactivates Bsal (Van 

Rooij et al. 2017). 

The above conditions apply to all 

salamanders listed as injurious in 50 

CFR 16.14 due to the risk of carrying 

Bsal. Also not considered injurious, and 

therefore exempt, are eggs, gametes, 

chemically preserved specimens or 

parts (including tissues), and molecular 

specimens consisting of only the nucleic 

acids (DNA or RNA) from organisms. 

The appropriate concentration and 

minimum exposure time for a given 

chemical preservative or fixative to 

render any Bsal organisms non-viable 

varies with the precise chemical 

formulation and should be utilized as 

described in association with such 

actions in the peer-reviewed literature. 

For example, Bsal is killed when 

exposed to 70 percent ethanol for at 

least 60 seconds (Van Rooij et al. 2017). 

However, parts that are otherwise 

preserved by air-drying or at a 

temperature and time that does not meet 

the above criteria or at a cold 

temperature (such as freezing) are 

considered injurious because Bsal is not 

inactivated by those methods. Purchase, 

sale, and other activities with the listed 

salamanders strictly within the 

boundaries of the enumerated 

jurisdictions within the shipment clause 

are not regulated under 18 U.S.C. 42. 

This second interim rule takes effect 

on the date specified above in DATES, 

but we are providing the public with a 

period of time to comment on the listing 

and associated documents. The 

resulting final rule will contain 

responses to comments received on the 

second interim rule, state the final 

decision, and provide the justification 

for that decision. 

Listing Species That Carry Pathogens 

Pathogens are such agents as viruses, 

bacteria, and fungi that cause disease in 

animals and plants. The Service does 

not have the direct authority under 18 

U.S.C. 42(a)(1) to list pathogens as 

injurious. We also cannot list or regulate 

fomites (materials, such as water, that 

can act as passive carriers and transfer 

pathogens). However, we can list wild 

mammals, wild birds, fishes, mollusks, 

crustaceans, amphibians, or reptiles that 

are hosts to or carriers of pathogens and 

that can be injurious if the likelihood, 

scope, and severity of effects 

significantly affect one or more of the 

interests listed in the statute. Even if the 

host species cannot establish 

populations in the wild, the host can 

present significant risk if the pathogen 

the host is carrying can infect wildlife 

or wildlife resources or affect human 

beings or the interests of agriculture, 

horticulture, or forestry in the United 

States. Among other impacts, diseases 

caused by introduced pathogens reduce 

biodiversity (the variety of different 

types of life in a region) and have been 

implicated in the local extinction of 

many animal taxa (Daszak et al. 2000). 

Listing and Evaluation Process 

The regulations contained in part 16 

of title 50 of the Code of Federal 

Regulations (CFR) implement 18 U.S.C. 

42(a)(1) and include the names of 

species determined by the Service or by 

Congress to be injurious. Under the 

terms of the statute, the Secretary of the 

Interior may prescribe by regulation 

those wild mammals, wild birds, fishes, 

mollusks, crustaceans, amphibians, 

reptiles, and the offspring or eggs of any 

of the foregoing that are injurious to 

humans, to the interests of agriculture, 

horticulture, or forestry, or to the 

wildlife or wildlife resources of the 

United States. The lists of injurious 

wildlife species are found at 50 CFR 

16.11–16.15, with § 16.14 being for 

amphibians. Under these regulations, 

species are added to the lists of 

injurious wildlife to protect statutorily 

enumerated interests from potential and 

known negative effects. Most species 

listed have the capacity to establish 

populations in the wild, spread, and 

cause harm. However, a species can be 

listed based solely on its capacity to 

cause harm. For example, uneviscerated 

dead salmonids without a health 

certificate are not capable of 

establishing in the United States, but 

they are injurious because the pathogens 

they may carry are harmful. 

Under 18 U.S.C. 42, the Service can 

list species that are nonnative and those 

that are indigenous to the United States. 

In the case of an indigenous species, for 

example, the Service may find that it is 

injurious because its transport and 

release outside the species’ range may 

cause harm to human beings, 

agricultural or forestry interests, or 

natural systems. Furthermore, a species 

does not have to be currently imported 

or present in the wild in the United 

States for the Service to list it as 

injurious. For species not yet imported 

into the United States, the objective of 

listing is to prevent that species’ 

importation and likely introduction and 

possible establishment and spread in 

the wild, thereby preventing injurious 

effects consistent with the purposes of 

18 U.S.C. 42. 

In response to the D.C. Circuit Court 

of Appeals Decision in United States 

Association of Reptile Keepers, Inc. v. 


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Zinke, 852 F.3d 1131 (D.C. Cir. 2017), 

the prohibition on interstate transport in 

the 2016 interim rule has been 

modified. Because of the court’s 

decision, injurious wildlife listings, 

including those listed by Congress 

through statutes (fruit bats (genus 

Pteropus), mongoose, zebra mussel, 

brown tree snake, bighead carp, quagga 

mussel), no longer result in a statutory 

prohibition on interstate transport of 

injurious wildlife between States within 

the continental United States. This 

means that transportation of injurious 

wildlife between the 49 States within 

the continental United States (the 

contiguous 48 States and Alaska) is not 

prohibited by 18 U.S.C. 42(a) (codified 

in Federal regulations at 50 CFR 16.3), 

unless that movement of the wildlife is 

restricted due to conditions associated 

with issued injurious wildlife permits. 

Thus, an injurious wildlife permit is 

generally not required to transport 

injurious species across State lines of 

any of the 49 continental States. 

However, a permit is still required for 

the movement of an injurious animal 

that was previously permitted for 

import or for the progeny of an 

individual that was permitted for 

import. 

Import of injurious wildlife into the 

United States remains prohibited. In 

addition, transport of injurious wildlife 

between the enumerated jurisdictions in 

the shipment clause of 18 U.S.C. 42 (the 

continental United States, the District of 

Columbia, Hawaii, the Commonwealth 

of Puerto Rico, and any territory or 

possession of the United States), 

codified in Federal regulations at 50 

CFR 16.3, remains prohibited. These 

prohibited activities may be undertaken 

by permit for zoological, educational, 

medical, or scientific purposes (in 

accordance with permit regulations at 

50 CFR 16.22), or by Federal agencies 

without a permit solely for their own 

use, upon filing a written declaration 

with the District Director of Customs 

and the U.S. Fish and Wildlife Service 

inspector at the port of entry. Interstate 

transport between States within the 

continental United States is not 

prohibited under the current 

prohibitions of 18 U.S.C. 42(a), and 18 

U.S.C. 42(a) does not prohibit intrastate 

transport (transport within a State or 

territory) or possession of injurious 

species. However, injurious wildlife 

unlawfully imported into the United 

States or transported between the 

enumerated jurisdictions is unlawful to 

transport within the continental United 

States, or to transport within a State or 

Territory, under the Lacey Act 

Amendments of 1981, 16 U.S.C. 

3372(a)(1). It is the responsibility of a 

person who may be engaged in transport 

or use of injurious wildlife to be aware 

of any Federal, State, Tribal, or 

Territorial law or regulation that applies 

to that activity. 

The Service prepares a listing rule by 

first assessing the relevant aspects of the 

biology of the species, such as its 

habitat, diet, reproductive capacity, 

climate, predatory capacity, and threats 

to its survival. This assessment is used 

to develop the next step, which is to 

evaluate whether any of these aspects 

contribute to the species being invasive 

or otherwise harmful. 

The Service uses one or more of the 

injurious wildlife listing criteria 

identified below to evaluate whether a 

species qualifies as injurious under 18 

U.S.C. 42. The results of the analysis 

using these criteria serve as a general 

basis for the Service’s regulatory 

decisions regarding injurious wildlife 

species listings. Biologists and risk 

managers within the Service who are 

knowledgeable about a species that is 

being evaluated assess both the factors 

that contribute to and the factors and 

measures that reduce or remove the 

likelihood of injuriousness. 

(1) Factors that contribute to 

injuriousness: 

• The likelihood of release or escape; 

• Potential to survive, become 

established, and spread; 

• Impacts on wildlife resources or 

ecosystems through hybridization and 

competition for food and habitats, 

habitat degradation and destruction, 

predation, and pathogen transfer; 

• Impacts to threatened and 

endangered species and their habitats; 

• Impacts to human beings, forestry, 

horticulture, and agriculture; and 

• Wildlife or habitat damages that 

may occur from control measures. 

(2) Measures that reduce the 

likelihood of the species being 

considered as injurious: 

• Ability to prevent escape and 

establishment; 

• Potential to eradicate or manage 

established populations (for example, 

making organisms sterile); 

• Ability to rehabilitate disturbed 

ecosystems; 

• Ability to prevent or control the 

spread of pathogens or parasites; and 

• Any potential ecological benefits to 

introduction. 

For this second interim rule, we 

provide a general summary of the 

biology of salamanders and of the 

fungus, followed by the evaluation for 

both as injurious. For injuriousness of 

the salamanders, we focused on the 

third bullet above ‘‘Impacts on wildlife 

resources or ecosystems through * * * 

pathogen transfer.’’ The issue in this 

rule is not about a given salamander 

species or genus being invasive but 

rather the role of salamanders in 

introducing the Bsal fungus into the 

United States and the scope and severity 

of effects caused by salamanders that are 

carriers of Bsal on the wildlife or 

wildlife resources of the United States. 

The Service obtains an extensive 

amount of amphibian import data from 

our Office of Law Enforcement’s (OLE) 

Law Enforcement Management 

Information System (LEMIS). LEMIS is 

an electronic database utilized by all 

Service law enforcement officers, 

including the Service’s conservation 

officers, wildlife inspectors, refuge 

officers, and special agents. LEMIS 

serves as the portal in which all Service 

wildlife violations are documented and 

intelligence is gathered and shared 

between law enforcement offices across 

the country. LEMIS also serves as the 

conduit for all declared imports and 

exports of wildlife and wildlife products 

and the database of all such wildlife 

trade data in the United States, both 

legal and illegal. The database provided 

us with information for this rule on 

what species were imported; quantity; 

countries of origin; ports of import; 

whether imported as live, dead, eggs, 

parts, or other; purpose for importing; 

and other relevant variables for the 

years 2010 to 2020 (USFWS OLE 2021). 

We evaluated Bsal and the 

salamander species that carry this 

fungus using the injurious wildlife 

evaluation criteria, described in more 

detail as part of this second interim rule 

in G. Factors That Contribute to 

Injuriousness of Salamanders, which 

the Service developed to evaluate 

whether a species qualifies as injurious 

under the Act. The resulting analysis 

serves as a basis for the Service’s 

regulatory decision regarding injurious 

wildlife species listings. This second 

interim rule finds that Bsal is a 

significant threat to the wildlife and 

wildlife resources of the United States 

and lists 16 genera of salamanders that 

we have determined to be injurious 

because they are likely carriers of Bsal 

and may introduce the fungus into the 

United States. 

A. Species Information for Salamanders 

Salamander Nomenclature and 

Taxonomy 

The Service does not have a uniform 

policy for taxonomically identifying 

amphibians. In this interim rule, we use 

taxonomic nomenclature as described 

by AmphibiaWeb (http://

amphibiaweb.org) with some 

comparison to the Integrated Taxonomic 


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Information System (ITIS) (http://

www.itis.gov). The system used by 

AmphibiaWeb represents one of the 

most widely accepted salamander 

taxonomic systems in the scientific 

community because it relies on criteria 

including, but not limited to, 

monophyly (common descent from a 

single ancestor), stability, expertise of 

scientists, and general acceptance by the 

amphibian conservation community and 

is frequently updated. As a Federal 

resource for taxonomic information, the 

Service also uses ITIS as an agency 

resource. As of May 1, 2023, 

AmphibiaWeb (2023b) reported 804 

species of all salamanders in 68 genera 

and 10 families, and ITIS reported 738 

species in 70 genera and 9 families. 

In this rule, when we refer to 

salamanders, we include animals from 

the order Caudata commonly referred to 

as salamanders and newts. The 

nomenclature and taxonomy of 

salamander species that we are 

regulating should be provided as 

accurately as possible to the public so 

that the public and law enforcement 

officers know what is being regulated. 

However, the science is evolving, 

making consistency even from the 2016 

interim rule difficult. The classification 

remained relatively unchanged from the 

1960s until the 1990s, when advances in 

DNA sequencing enabled researchers to 

examine species relationships more 

closely (Petranka 1998). Furthermore, 

dozens of amphibian species from 

remote regions of the world are 

discovered every year (AmphibiaWeb 

2021). This is generally why the number 

of species listed increased within the 20 

genera in the final rule to the 2016 

interim rule. For these reasons, we are 

not including the names of the species 

within each listed genus in 50 CFR 

16.14. As long as the species is within 

a listed genus, it is covered as an 

injurious species, as in the final rule. 

Salamander Biology 

Salamanders belong to the class 

Amphibia, a group of cold-blooded 

vertebrate animals comprising frogs and 

toads (order Anura), salamanders and 

newts (order Caudata), and caecilians 

(order Gymnophiona). The word 

‘‘amphibian’’ is derived from the fact 

that most of the species spend part of 

their lives in water and part on land. 

Frogs and toads have legs but no tails 

as adults, and caecilians have tails but 

no legs. Morphologically, salamanders 

are generally characterized by their 

relatively large, vertically flattened tails, 

two front and two hind legs that are 

approximately the same size (Petranka 

1998), and skin with glands that can be 

either rough or smooth (Stebbins and 

Cohen 1997). Exceptions include 

Sirenidae, which have two small 

forelimbs and no hindlimbs, and 

Amphiumidae, which have four 

vestigial limbs. Adult salamanders range 

in length from around 4 centimeters (1.5 

inches) to over 1.5 meters (5 feet) 

(Stebbins and Cohen 1997). Another 

distinction between anurans (the frogs 

and toads) and the caudata (salamanders 

and newts) is that the anurans have 

internal gills as larvae and salamanders 

have external gills as larvae. 

Salamanders can live for many years, 

but documented lifespans vary. Larger 

salamanders tend to live longer than 

smaller ones, and, with proper care, 

salamanders in captivity frequently live 

longer than those in the wild (Duellman 

and Trueb 1986). Records for captive 

animals range from 5 years for most 

plethodontids (lungless salamanders) to 

55 years for the Japanese giant 

salamander (Andrias japonicus) 

(Duellman and Trueb 1986). The Olm or 

blind cave salamander (Proteus 

anguinus), which lives in caves in 

southern Europe, has been documented 

living for at least 48 years in the wild, 

with an estimated lifespan of more than 

100 years (Live Science 2015). 

Salamanders are carnivorous and eat 

a wide variety of prey, depending on 

habitat and the stage of their life cycle. 

Terrestrial adult salamanders eat 

earthworms, insect eggs, and other small 

invertebrates, while aquatic 

salamanders eat all of these in addition 

to small fish, aquatic insects, and other 

amphibians. Some salamander larvae 

can also be omnivorous and eat plants 

and animals. 

Many salamanders have unique 

structural features, including costal 

grooves (grooves on the sides of the 

body that increase skin surface area for 

water absorption and transport) and 

nasolabial grooves (vertical slits 

between the nostril and upper lip used 

for sensing chemical stimuli in the 

environment) that can be used to 

differentiate between salamander 

species (Petranka 1998). Important 

features for identifying salamanders 

include head shape and size, fin shape 

and color, gill morphology, color 

patterns, number of toes, size, body 

shape, tooth patterns, and number of 

costal grooves. Some species appear 

similar to each other, and similarity of 

appearance within some families, such 

as Salamandridae, can make it difficult 

to differentiate between species, 

requiring close inspection of small 

physical characteristics. 

Salamanders occupy a wide range of 

habitats, including streams, trees, land 

(including forests, grasslands, and rocky 

slopes), underground, and caves. 

Salamanders are cryptic (difficult to 

find) partly because they occupy moist, 

cool places, such as underneath logs 

and between rock crevices on land or 

under rocks and logs in the water. 

Salamander courtship between males 

and females is regulated by chemicals 

that are released from specialized glands 

in the skin. Most salamanders reproduce 

by laying eggs in water with two 

exceptions: most members of family 

Plethodontidae lay their eggs on land, 

and the European species known as the 

alpine salamander (Salamandra atra) 

gives birth to live young (Stebbins and 

Cohen 1997). Eggs are surrounded by a 

protective jelly or membrane that keeps 

them from drying out. Almost all 

species of salamanders breed during 

specific seasons, and the length of time 

between mating and egg-laying varies 

considerably between species (Petranka 

1998). Species that lay aquatic eggs 

place them in either streams or ponds, 

and species that lay their eggs on land 

choose hidden places, such as 

underground burrows, decaying logs, 

and moist rock crevices (Petranka 1998), 

where the young typically undergo 

direct development, whereby 

metamorphosis occurs in the egg and 

fully formed salamanders emerge from 

the eggs. 

The majority of the species in 

Ambystomatidae (mole salamanders) 

spend most of the year underground in 

rodent burrows and emerge only on 

rainy nights to mate and feed. 

Ambystomatid salamanders are famous 

for the migration of large numbers of 

individuals to breeding ponds. One 

example of a species that spends most 

of its life on land but that moves to 

aquatic areas to breed is the California 

tiger salamander (Ambystoma 

californiense). During winter rains, this 

species migrates across land to aquatic 

pools, such as cattle tanks and 

ephemeral pools, to breed. At the 

breeding pools, individuals come in 

contact with each other, even though 

they may not come in contact with each 

other during most of the rest of their 

lives on land (Barry and Shaffer 1994). 

However, the related axolotl (A. 

mexicanum) is unlike other 

salamanders by being neotenic (they do 

not undergo metamorphosis). 

Furthermore, some ambystomatids 

retain their larval morphology as 

reproductive adults until certain 

environmental cues trigger 

metamorphosis into terrestrial adult 

morphology. 

Habitat Conditions and Native Range of 

U.S. Salamanders 

With more native salamander species 

than any other country in the world, the 


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United States is a salamander diversity 

hotspot (PARC 2015). Salamanders are 

widespread in the United States 

(Caudata Culture 2015a; U.S. National 

Park Service 2015). Areas of particularly 

high salamander diversity include the 

Eastern United States, with large 

numbers of plethodontid salamanders in 

the southern Appalachian Mountains 

(Richgels et al. 2016). 

Salamanders in the United States 

occupy a wide range of habitats, 

including streams, trees, land (including 

forests, grasslands, and rocky slopes), 

underground, and caves. These 

locations are most conducive to the 

relatively cool, moist conditions under 

which both salamanders and Bsal thrive 

(Duellman and Trueb 1986; Piotrowski 

et al. 2004; Blooi et al. 2015a). Central 

and North American salamanders as a 

group are active at average temperatures 

of 11 °C (52 °F) to 20 °C (68 °F) 

(Duellman and Trueb 1986), fully 

encompassing the optimum temperature 

for Bsal growth as described below 

under Climate Tolerance. Salamanders 

require some amount of constant 

moisture for physiological function, 

such as osmoregulation (controlling 

body fluid, water, and salt balance) or 

for cutaneous respiration, as in the 

lungless family Plethodontidae, or for 

temperature regulation (Duellman and 

Trueb 1986). 

Twenty species of U.S. salamanders 

from seven genera (Ambystoma, 

Batrachoseps, Cryptobranchus, Eurycea, 

Necturus, Phaeognathus, Plethodon) are 

currently listed as endangered or 

threatened under the Endangered 

Species Act of 1973, as amended (16 

U.S.C. 1531 et seq.) (ESA). The specific 

vulnerability and carrier status of these 

species to Bsal is described below in 

Vulnerability and Carrier Status of 

Threatened and Endangered Species. 

Of the 221 salamander species in 23 

genera in 9 families native to the United 

States, we expect that at least 164 

species in 13 genera and in 5 families 

are capable of being carriers of Bsal: 

Ambystomatidae, Cryptobranchidae, 

Salamandridae, Sirenidae, and 

Plethodontidae. In North America, 

species in the family Salamandridae 

occur on the west coast of the United 

States and Canada, from southern 

California to southeastern Alaska, as 

well as much of the eastern half of the 

United States and extreme southeastern 

Canada (Caudata Culture 2015a). 

Members of the family Sirenidae occur 

throughout the southeastern Atlantic 

and Gulf of Mexico coastal plains and 

the Mississippi River Valley (Leja 2005) 

(lesser siren, Siren intermedia) and in 

the Atlantic coastal plains from south 

Florida to Virginia (greater siren, Siren 

lacertina) (Hendricks 2005). The 

distribution of salamanders of the 

family Plethodontidae in the western 

hemisphere is from southern Canada to 

Bolivia and Brazil, except for members 

of the genus Hydromantes, which occur 

in California (AmphibiaWeb 2023a, 

Caudata Culture 2015a). 

Ambystomatidae has only one genus, 

and the 32 species are widely 

distributed in North America from 

southern Canada south to Mexico City. 

Cryptobranchidae is represented by one 

species in North America (eastern 

hellbender Cryptobranchus 

alleganienesis). 

Role of Salamanders in the Ecosystem 

Salamanders play important roles in 

ecosystem function and as indicators of 

ecosystem health and stability (Davic 

and Welsh 2004). For example, 

salamanders of family Plethodontidae 

have life-history characteristics that 

make them highly effective at 

controlling invertebrates that would 

otherwise consume the leaf litter, thus 

releasing carbon to the environment 

(Best and Welsh 2014). 

In forests, salamanders are also among 

the most abundant vertebrates. Despite 

the relatively small size of most 

salamanders compared to most other 

native vertebrates, this sheer abundance 

contributes to a significant amount of 

biomass in the ecosystem, and, 

therefore, salamanders make significant 

contributions to nutrient cycling and 

transport (Burton and Likens 1975). For 

example, Ambystomatid salamanders 

can make significant contributions to 

energy and nutrient transport in forest 

ecosystems (Regester et al. 2006) and in 

pond ecosystems (Holomuzki et al. 

1994). Many terrestrial salamanders 

consume arthropods (insects and related 

invertebrates) that feed on leaf litter, 

and the invertebrates’ feeding process 

causes the release of carbon dioxide 

from the ground into the atmosphere. 

With fewer salamanders and more litterconsumers, more carbon is released 

from the soil, contributing to an excess 

of carbon dioxide in the atmosphere. 

Salamander populations help reduce 

carbon emissions from leaf litter 

decomposition, which has implications 

for the global carbon cycle (Wyman 

1998; Best and Welsh 2014; North 

American Bsal Task Force 2020, Laking 

et al. 2021). This process is known as 

carbon sequestration, which is the 

storage of carbon dioxide to slow or 

reverse atmospheric carbon dioxide 

pollution and to mitigate or reverse 

climate change. Salamanders that live 

underground also contribute to soil 

dynamics by creating, modifying, and 

otherwise regulating the systems of 

underground burrows in which they 

live (Davic and Welsh 2004). Finally, 

salamanders are important prey species 

themselves and provide energy sources 

for higher predators (Davic and Welsh 

2004), including fishes, reptiles, birds, 

and mammals. 

In vernal pond communities, 

Ambystoma species are the top 

predators and, therefore, control the 

abundance of aquatic invertebrates and 

other amphibians (Petranka 1998). The 

high numbers of many amphibians, 

including salamanders, in some 

ecosystems also provide a substantial 

source of prey for other vertebrates in 

the ecosystem (Harper et al. 2008; Davic 

and Welsh 2004); therefore, other native 

species that prey on salamanders can 

also be affected by disease-related 

declines. 

B. Species Information for 

Batrachochytrium Salamandrivorans 

General Information About Bsal 

Bsal is a fungus in the phylum 

Chytridiomycota and the order 

Rhizophydiales. It was identified in 

2013 after reports of a dramatic 

mortality event of fire salamanders in 

Europe (Martel et al. 2013). In drawing 

some of our conclusions about the 

effects of Bsal on U.S. wildlife and 

wildlife resources for the 2016 interim 

rule, the Service used Batrachochytrium 

dendrobatidis (Bd) as a surrogate 

(similar substitute) species because little 

was known about the emerging disease 

caused by Bsal. Considerably more was 

known about Bd due to its discovery 

and description more than 15 years 

earlier (Berger et al. 1998, Longcore et 

al. 1999). Bd is found on every 

continent that supports amphibians, 

while Bsal is known to be only in 

Europe and Asia. Bd has resulted in the 

serious decline and extinction of more 

than 200 species of amphibians 

worldwide and has posed the greatest 

threat to biodiversity of any known 

pathogen (Martel et al. 2013). The severe 

effects that Bd, also a fungal pathogen 

species closely related to Bsal, has had 

on amphibian populations raised 

additional alarm about the expected 

consequences of a Bsal introduction and 

the need to take immediate action under 

an interim rule. 

Two scientific risk assessments of 

Bsal used Bd in determining the risk of 

Bsal based on transmission, spread, and 

population-level effects (Stephen et al. 

2015; Richgels et al. 2016). Gray et al. 

(2015) found that both fungi infect the 

epidermal cells of the amphibian skin, 

and the clinical signs for both include 

excessive skin shedding, lethargy, 

anorexia, abnormal posture, and death; 


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however, the lesions produced by Bd 

are mainly epidermal hyperplasia and 

hyperkeratosis (rarely ulcerations), and 

those produced by Bsal are mainly skin 

ulcerations and destruction of the 

epidermis. Similarities and differences 

between the two fungal pathogens and 

the diseases they cause are discussed in 

Farrar et al. (2017), Longo et al. (2019), 

Rebollar et al. (2020), and Rollins and 

Le Sage (2021). 

Since the 2016 interim rule was 

published, scientists have been studying 

many aspects of Bsal, and this rule 

reflects new research. Relevant studies 

confirm or expand on our previous 

information, with many adding new 

insight to the fungus and disease, and 

some providing documentation to 

support adding new genera as carriers. 

Until Bsal was discovered, the fungal 

disease chytridiomycosis was thought to 

be caused by a single species of 

pathogenic fungus, Bd, which was the 

only species from that phylum known to 

parasitize vertebrate hosts (Longcore 

1999; Johnson and Speare 2003). Bd has 

been implicated in the decline and 

extinction of amphibian species at the 

global scale (Berger et al. 1998; Daszak 

et al. 2003; Lips et al. 2006; Walker et 

al. 2008; Vredenburg et al. 2010; Cheng 

et al. 2011). Bd has been found on every 

continent except Antarctica, and it is 

known to have affected more than 500 

species of amphibians, including all 

orders of amphibians (frogs, 

salamanders, and caecilians) worldwide 

(Chytridcrisis 2015a; Fisher et al. 2009; 

Olson et al. 2013). 

Bsal came to the attention of the 

scientific community in 2013 when 

Spitzen van der Sluijs et al. (2013) 

observed a 96 percent decline in fire 

salamanders in the Netherlands but was 

‘‘unable to attribute this to any known 

cause of amphibian decline, such as Bd, 

ranavirus or habitat degradation.’’ 

Martel et al. (2013) subsequently 

identified the cause of the salamander 

decline in the Netherlands as a newly 

described species of fungus now known 

as Bsal. Their work confirmed that Bsal 

is closely related to Bd and is also 

capable of causing chytridiomycosis; 

both are in the genus Batrachochytrium. 

Analysis of a broad range of 

representative chytrid fungi show that 

Bsal represents a previously 

undescribed species that shares early 

evolutionary origins with the 

pathogenic fungus Bd (Martel et al. 

2013). 

The natural amphibian hosts of Bsal 

remain unknown, but as of publication 

of the 2016 interim rule, Bsal had been 

found only in salamanders and 

appeared capable of causing lethal 

chytridiomycosis only in salamanders 

(Martel et al. 2014). Subsequently, 

several species of anurans have been 

found to be carriers, such as the 

midwife toad (a frog) Alytes obstetricans 

(Stegen et al. 2017) and small-webbed 

firebelly toad Bombina 

microdeladigitora (Nguyen et al. 2017). 

How the Fungus Affects Salamanders 

The ‘‘salamandrivorans’’ in 

Batrachochytrium salamandrivorans 

means ‘‘salamander eating’’ and 

figuratively describes the effects of the 

fungus on salamanders. Bsal infects 

primarily the skin of salamanders, but 

deeper tissues or internal organs may be 

affected (McDonald et al. 2020). The 

skin of post-embryonic salamanders has 

a layer of keratin (Seifert et al. 2019) 

covered by a mucosal microbiome of 

beneficial biota that normally protect 

them from harmful microbes entering 

the body (Bletz et al. 2018; Rebollar et 

al. 2020), such as a fungus. 

The cells of the fungus (thalli) embed 

themselves in the skin cells of the 

salamander, thereby causing erosive 

lesions. Lesions consist of sores on the 

skin that erode and ulcerate, with 

secondary bacterial infection likely 

occurring after the sores appear (Martel 

et al. 2013; Gray et al. 2015; Bletz et al. 

2018), although many of the 

salamanders reported at the beginning 

of the European Bsal outbreak seemed to 

lack obvious external lesions (Spitzenvan der Sluijs et al. 2013). The loss of 

epidermal integrity from the lesions 

impairs the skin’s ability to maintain 

fluid and electrolyte balance and also 

perforates the barrier that protects the 

animal from pathogens and 

compromises their line of defense 

against disease (Gray et al. 2015; 

Rebollar et al. 2020). Experimental 

infections of fire salamanders in the 

laboratory caused death 12 to 18 days 

after exposure, with the same clinical 

signs and pathological lesions found in 

the European outbreak (Martel et al. 

2013) and in another experimental 

infection around 3 weeks after a high 

dose exposure (Stegen et al. 2017). 

Martel et al. (2013) found that infected 

fire salamanders developed shallow 

skin lesions and deep ulcerations all 

over the body, became anorexic and 

apathetic, and suffered from 

neurological signs including a loss of 

voluntary movement and muscle 

coordination. Death occurred within 7 

days of clinical signs first appearing in 

species with lethal vulnerability. Death 

is generally preceded by a brief episode 

of abnormal body posture and behavior 

(Gray et al. 2015). 

Some species succumb quickly to Bsal 

after infection, while others seem to 

tolerate it and eventually clear the 

infection (Martel et al. 2014). However, 

long-term effects on tolerant species are 

not known. Several studies suggest 

negative long-term effects. For example, 

the long-term proliferation of the fungus 

within the keratinized limb tissue of the 

olm (Proteus anguinus) may coincide 

with a more subtle cost associated with 

increased energy expenditure, impaired 

locomotion, or increased vulnerability 

of the limbs to secondary infection (Li 

et al. 2020). Another study detected 

potential sublethal reductions of growth 

caused by Bsal exposure in juvenile 

spotted salamanders soon after 

metamorphosis, although not in older 

juveniles (Barnhart et al. 2020). The 

initial exposure to Bsal may have 

created a stress response that helped 

activate the immune system; this 

activation dissipated after the threat 

dissipated but may come at a cost 

because juvenile salamanders with 

higher corticosterone release rates 

immediately after exposure to Bsal had 

lower growth compared to control group 

salamanders 30 days post-exposure. 

The outcomes of coinfections by Bd 

and Bsal on salamanders have been 

studied because they both affect the 

skin. Bd and Bsal are the only known 

Chytridiomycota to have adapted to 

colonize vertebrates, yet Bd infects all 

three orders of Amphibia (especially 

Anura), while Bsal is currently known 

to infect primarily the order Caudata 

(Farrar et al. 2017). If Bsal enters the 

Western Hemisphere where Bd is 

already widespread, coinfections could 

occur, and some research suggests the 

results could be more serious to the 

Bsal-naı ̈ve salamanders than Bd 

infection alone (Longo et al. 2019; 

McDonald et al. 2020). Longo et al. 

(2019) and McDonald et al. (2020) 

studied coinfection in eastern newts 

(Notophthalmus viridescens), a 

widespread native salamander. Longo et 

al. (2019) tested newts to see if prior 

exposure to Bd provided immune 

protection from Bsal or instead reduced 

the protection. They found that newts 

can clear Bd alone; resistance is specific 

to Bd and does not prevent Bsal 

infection; simultaneous coinfections 

were the most lethal, even at reduced 

dosages; host mortality from Bsal can be 

much slower than previously found; 

and some wild newts may have innate 

Bsal immunity from prior exposure to 

Bd, but other factors may be involved. 

McDonald et al. (2020) found that Bd 

and Bsal coinfection reduced the host’s 

immune response more than with Bsal 

alone. Thus, if Bsal enters a newt 

population where Bd already exists, the 

Bsal infection may be compounded by 

the Bd infection. 


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The Service has no direct evidence 

that Bsal affects reproductive tissue, 

such as eggs or gametes. Since Bsal 

attaches to and utilizes keratincontaining substrate for growth, and 

eggs and gametes do not contain keratin, 

we have no evidence that eggs and 

gametes will carry Bsal (L. Sprague, 

USFWS, pers. comm. 2021). Thus, we 

do not believe that salamander 

reproductive material can serve as a 

vector for Bsal introduction into the 

United States. 

Thermal Tolerance 

Temperature has a significant effect 

on the growth and disease development 

of Bsal in salamanders (Martel et al. 

2014; Carter et al. 2021). Bsal appears to 

prefer an in vitro temperature range for 

growth and infection of 10–15 °C (50– 

59 °F) (Martel et al. 2013; Blooi et al. 

2015a; Stephen et al. 2015; Thomas et 

al. 2019). Bsal has shown some spore 

growth in temperatures as low as 5 °C 

(41 °F) and dies at 25 °C (77 °F) and 

above (Martel et al. 2013). However, the 

majority of Bsal-infected salamanders in 

natural Vietnamese ponds were in water 

temperatures of 20–25 °C (68–77 °F) and 

as high as 26.43 °C (79.6 °F) (Laking et 

al. 2017). In a laboratory study, 

salamanders were most easily infected 

by Bsal at temperatures of 15 °C (59 °F) 

and 20 °C (68 °F), while Bsal growth was 

inhibited at 25 °C (77 °F) (Blooi et al. 

2015a). The same temperature response 

was also observed for Bsal raised in 

culture (Blooi et al. 2015a). Grear et al. 

2021 used 15 °C (59 °F) as the thermal 

optimum for Bsal growth and evaluated 

the impact of a 27 °C (81 °F) thermal 

maximum to the resulting risk. Carter et 

al. (2021) found that adult and juvenile 

Notophthalmus viridescens died faster 

due to Bsal chytridiomycosis at 14 °C 

(57 °F) than at 6 °C (43 °F) and 22 °C 

(72 °F). 

These experimental data suggest that 

salamanders living at cooler 

temperatures are more at risk to 

infection by Bsal. Animals that survive 

at temperatures above the optimal range 

for fungal growth are likely to be at 

reduced risk to infection. However, the 

average temperature range of North and 

Central American salamander species is 

from 11 °C (52 °F) to 20 °C (68 °F) 

(Duellman and Trueb 1986; the citation 

does not separate North and Central 

American data), so salamanders 

regularly reaching 25 °C (77 °F) in the 

natural environment is uncommon. 

Bales et al. (2015) noted that the native 

salamander species, and by extension 

ecosystems, most at risk from a Bsal 

introduction would likely be those that 

occupy similar thermal ranges as the 

European fire salamander (Bales et al. 

2015). Richgels et al. (2016) also cited 

research that Bd is capable of infecting 

amphibians along a larger temperature 

profile than originally predicted, though 

it is unknown whether this is the case 

for Bsal. 

Ecology and Habitat Preferences 

The chytrid fungus Bd can live 

outside of a host and requires water to 

disperse because it reproduces asexually 

by forming motile zoospores; 

preliminary studies of Bsal indicate that 

similar modes of survival and 

transmission are highly likely (Longcore 

1999; Martel et al. 2013). As the threat 

assessment by Stephen et al. (2015) 

noted, ‘‘Bd is known to remain viable 

for several days to weeks in water 

(Johnson and Speare 2013) and moist 

organic matter (Johnson and Speare 

2003), even in the absence of nutrients. 

It is likely that Bsal can also survive in 

moist environments, independent of an 

amphibian host.’’ Stegen et al. (2017) 

states that Bsal adopts a dual 

transmission strategy, with 

environmentally resistant nonmotile 

spores in addition to the motile spores 

identified in Bd. Bsal retains its 

virulence in water and soil as well as in 

anurans and less susceptible salamander 

species that function as a reservoir of 

infectious pathogens. The combined 

characteristics of the disease ecology 

suggest that Bsal is able to rapidly 

extirpate highly susceptible salamander 

populations across Europe. Stegen et al. 

(2017) also found that infected fire 

salamanders were shown to contaminate 

the forest soil and Bsal DNA could be 

detected even after 200 days. Actual 

transmission through contaminated 

forest soil was demonstrated up to 48 

hours after the soil had been in contact 

with an infected animal. Encysted Bsal 

spores were shown to remain infectious 

in pond water for at least 31 days. 

Together, the presence of a resistant 

spore with the ability to persist 

environmentally and to transmit 

through contaminated water and soil, 

combined with the occurrence of longterm-infected and pathogen-shedding 

amphibian hosts, creates the potential 

for extensive environmental reservoirs 

and hampers any effort to eradicate Bsal 

from an infected ecosystem. 

Environmental Conditions Needed To 

Survive 

The transmission and ecology of Bsal 

in the wild is likely to be similar to Bd 

based on the close taxonomic 

relationship between the species, their 

structural similarities, and their 

comparable pathophysiology (Martel et 

al. 2013; Stephen et al. (2015). Disease 

transmission is the means by which 

communicable pathogenic 

microorganisms, such as fungi, are 

spread from one organism to another. 

Johnson and Speare (2003) reported that 

Bd can survive in tap water and 

deionized water for up to 3 and 4 weeks, 

respectively, and up to 7 weeks in lake 

water. Bsal is also likely to survive in 

moist environments independent of an 

amphibian host; for example, Stegen et 

al. (2017) found that encysted spores 

can survive and remain infectious for at 

least 31 days in filtered pond water. 

While we do not have information on 

the response of Bsal to desiccation, Bd 

is highly impacted by drying and can 

survive desiccation for no more than 1 

hour in the laboratory (Garmyn et al. 

2012); Bsal would likely respond in a 

similar way. Bsal appears to be adapted 

to lower preferential temperatures 

compared to Bd, with optimal growth 

between 10 ßC and 15 ßC, and Bsal spore 

death occurring at temperatures greater 

than 25 ßC (Martel et al. 2013). These 

findings support the hypothesis that the 

pathogen coevolved with salamanders 

in the part of the world from which it 

is endemic, most likely in Asia (Martel 

et al. 2014; Laking et al. 2017). 

C. Population-Level and EcosystemLevel Effects of Bsal 

Population-Level Effects 

Several pathogens, including Bsal, Bd, 

ranaviruses, and aquatic oomycetes 

(water molds), have caused significant 

population-level declines in a range of 

amphibian species, and disease is 

thought to be a major driver of global 

amphibian decline (Bosch et al. 2001; 

Daszak et al. 2003; Martel et al. 2013). 

Disease poses a greater risk to small, 

isolated populations as well as those 

with decreased genetic diversity (Smith 

et al. 2008). Within the United States, 

diseases have been cited as contributing 

factors in the listing or need for recovery 

of several native amphibian species 

under the ESA. Examples include Bd in 

the Ozark hellbender (Cryptobranchus 

alleganiensis bishopi) (76 FR 61956, 

October 6, 2011), an undiagnosed 

disease in Sonora tiger salamanders 

(Ambystoma tigrinum stebbinsi) (62 FR 

665, January 6, 1997), and Bd in the 

mountain yellow-legged frog (Rana 

muscosa) (82 FR 24256, April 29, 2014; 

Vredenburg et al. 2010). 

As noted above in General 

Information About Bsal, Bsal is the most 

recently discovered pathogen associated 

with population-level amphibian 

declines, including a 96 percent 

reduction in Dutch populations of the 

European fire salamander in the period 

2010–2013 (Spitzen-van der Sluijs et al. 

2013; Martel et al. 2013). Due to the 


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overall sensitivity of amphibian 

populations to disease; a history of 

adverse, population-level effects in 

native amphibians; a direct association 

between Bsal and the decline of at least 

one European salamander population; 

and the adverse effects of some native 

salamanders to Bsal under experimental 

conditions, we conclude that the 

introduction of Bsal into the United 

States would cause significant, adverse, 

population-level effects in a number of 

native species. 

Ecosystem-Level Effects 

The preferred temperature range of 

Bsal can help predict those ecosystems 

that are at greatest risk should Bsal be 

introduced into the United States 

(Stephen et al. 2015). The native 

salamander species, and by extension 

ecosystems, most at risk from a Bsal 

introduction would likely be those that 

occupy similar thermal ranges as the 

European fire salamander (Bales et al. 

2015). 

Salamanders are important parts of 

the ecosystems in which they occur. 

Salamanders are often the most 

abundant vertebrates in terrestrial forest 

and riparian (the banks of watercourses) 

ecosystems, where they may compose a 

total biomass greater than or equal to 

birds or small mammals (Davic and 

Welsh 2004). This means that, despite 

their small size, the total weight of all 

salamanders in a given area may be 

more than the combined total weight of 

all birds or all small mammals. Because 

of their abundance under normal 

circumstances, salamanders are 

important prey species themselves and 

are energy sources for higher predators 

(Davic and Welsh 2004), including 

fishes, reptiles, birds, and mammals. 

Salamanders may be the dominant 

predator in headwater streams and 

ephemeral waterbodies where fish are 

absent (Davic and Welsh 2004). Within 

some food webs, salamanders are 

considered keystone predators due to 

their control of invertebrate prey 

populations and their resulting 

regulation of detritus decomposition 

and nutrient cycling (Davic and Welsh 

2004). By definition, keystone species 

are those that occupy niches that affect 

ecosystems and have little functional 

overlap with other species (Davic and 

Welsh 2004). Therefore, loss of these 

keystone species would result in 

significant ecosystem-level change. 

In addition to their roles in food webs 

and nutrient cycling, salamanders 

participate in a number of interspecific 

(between species) ecological 

relationships. Salamander species 

interact with one another through 

competition and predation to control 

the composition of their assemblages 

(taxonomically related species that 

occur within the same geographic 

community) (Davic and Welsh 2004; 

Fauth et al. 1996). Frequently, a single 

species is dominant within a given 

assemblage, particularly in terrestrial 

habitats, but which species dominates 

varies by location and ecosystem (Davic 

and Welsh 2004). We expect that 

ecosystems where the dominant 

salamander species is susceptible to 

lethal or sublethal Bsal infection would 

be at risk from an introduction of this 

pathogen. 

Salamanders also interact with 

invertebrate species in other 

ecologically important ways. Semiaquatic salamander species can move 

mollusks and shrimp eggs between 

waterbodies during their migrations, 

allowing these invertebrates to inhabit 

new areas (Davic and Welsh 2004). For 

example, a native species of salamander, 

the mudpuppy (Necturus maculosus), is 

a required host for developing stages of 

the salamander mussel (Simpsonaias 

ambigua), a native, freshwater mollusk 

(Davic and Welsh 2004; Gangloff and 

Folkerts 2006). We conclude that 

invertebrate species that depend on 

salamanders for aspects of their life 

cycle or ecology are likely to be 

adversely affected if their host species 

declines in response to a Bsal 

introduction. 

D. Invasiveness of Salamanders and 

Bsal 

Invasiveness of Salamanders 

Executive Order 13751 defines an 

‘‘invasive species’’ as a nonnative 

organism with regard to a particular 

ecosystem whose introduction causes or 

is likely to cause economic or 

environmental harm, or harm to human, 

animal, or plant health. Two key 

components of invasiveness are 

introduction into nonnative ecosystems 

and causing harm. 

Globally, 90 percent of salamander 

introductions have occurred through 

intentional releases (Tingley et al. 2010). 

As of 2010, salamanders comprised 22 

percent of all recorded amphibian 

introductions, with the highest number 

of salamander introductions (15) from 

the family Salamandridae, followed by 

salamanders from the families 

Ambystomatidae (4), Cryptobranchidae 

(2), and Proteidae (2) (Tingley et al. 

2010). Nonnative salamander 

introductions have been documented in 

the United States. The USGS’s 

Nonindigenous Aquatic Species 

database has U.S. records for 17 

salamander species that have been 

observed outside their native range 

(USGS 2023). Of those, 14 are native to 

the United States but were discovered 

outside of their native ranges, and 3 

(Japanese fire-bellied newt Cynops 

pyrrhogaster, Oriental fire-bellied newt 

Cynops orientalis, and paddle-tailed 

newt Paramesotriton (Pachytriton) 

labiatus) are species native to the 

eastern hemisphere. In Florida, the 

Oriental fire-bellied newt and paddletailed newt (family Salamandridae), 

which are native to China, have been 

found in the wild near an animal 

importer’s facility, either as the result of 

intentional releases or escapes from 

enclosures (Krysko et al. 2011), although 

none have been reported since 2010 

(USGS NAS 2021 [CYOR, PALA]). 

While these two species apparently did 

not establish invasive populations, their 

presence in the wild demonstrates a 

possibility that escaped individuals can 

persist long enough in the wild to 

transmit Bsal to native populations. 

Other invasions have been attributed 

to the use and subsequent release of 

salamanders used as fishing bait. 

Surveys of anglers have indicated that 

they routinely release salamanders into 

the areas where they fish, which 

includes areas that are not part of the 

salamander’s native habitats, suggesting 

that animals are routinely moved long 

distances (Picco and Collins 2008). 

Furthermore, Picco and Collins (2008) 

found that salamanders sold as bait 

were highly infected with both 

ranavirus and Bd, thereby increasing the 

likelihood of disease transmission into 

new areas of the United States through 

the act of fishing. 

Invasiveness and Transmission of Bsal 

As noted above under General 

Information About Bsal, Europe has 

been experiencing a severe decline in 

wild fire salamander populations in the 

Netherlands (Spitzen-van der Sluijs et 

al. 2013). This decline is so significant 

that fire salamander populations are 

facing local extinction in the 

Netherlands. A sharp decline in 

numbers has been observed since 2010, 

despite the species being listed as 

endangered on the Netherlands Red 

List, and at population levels that were 

thought to be stable. This enigmatic 

decline was not attributed to any known 

cause of amphibian decline, such as 

chytridiomycosis due to Bd, ranavirus, 

or habitat degradation. In late 2013, Bsal 

was isolated from infected fire 

salamanders in the Netherlands (Martel 

et al. 2013). 

Martel et al. (2014) later established 

the highly pathogenic nature of this new 

chytrid fungus. Molecular testing found 

Bsal in specimens collected from the 

wild (though none from North America) 


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and even in an archival (museum) 

sample that was 150 years old (Martel 

et al. 2014). Alpine newts (Ichthyosaura 

alpestris) and smooth newts (Lissotriton 

vulgaris) in the wild are also now 

known to be infected (Spitzen-van der 

Sluijs et al. 2016), as are palmate newts 

(L. helveticus; Lastra Gonza ́laz et al. 

2019). 

A wide variety of salamanders are 

negatively affected by the pathogen, but 

10 species of frogs and toads and the 1 

caecilian species did not appear to be 

(Martel et al. 2014). More recently, two 

anuran species have been shown to be 

carriers of Bsal. Small-webbed firebellied toads from wild populations in 

Vietnam and from individuals from that 

region imported into Germany tested 

positive for Bsal by qPCR swabs 

(Nguyen et al. 2017), and the midwife 

toad was capable of infecting fire 

salamanders for several weeks after 

experimental inoculation (Stegen et al. 

2017). However, we are listing only 

caudate genera with this rule, and 

anurans would need to be considered in 

separate rulemaking. The pathogenic 

nature of the fungus and its ability to 

infect a wide variety of salamanders, as 

described below in Vulnerability and 

Carrier Status, definitively demonstrate 

an invasive threat to salamanders in the 

United States. 

Emerging infectious diseases that can 

cause mass mortality are especially 

worrisome because they can cause 

extinction and subsequent loss of 

biodiversity relatively rapidly. The only 

in situ example of the spread of Bsal is 

with fire salamanders in Europe. Using 

this example, Schmidt et al. (2017) 

developed a model to explore the effects 

of the recently emerged Bsal. They 

showed that disease outbreaks can occur 

at very low host densities (one female 

per hectare (2.5 acres)) in the wild and 

that this is much lower than host 

densities in the wild. Therefore, all 

naturally occurring populations are at 

risk, and the model predicts a rapid 

collapse of the host population. 

Experiments have shown that Bd can be 

transferred from one species to another 

when an uninfected species comes into 

contact with an infected species (active 

carrier) or infected fomites (Carey et al. 

2006) (passive carrier). Bsal can 

similarly be transmitted from one 

species to another (Martel et al. 2014; 

Stegen et al. 2017; Homan et al. 2018). 

Bd has contributed to the decline of at 

least 501 amphibian species worldwide 

(6.5 percent of described amphibian 

species), which is the largest 

documented loss of global biodiversity 

attributable to a pathogen (Scheele et al. 

2019). Bsal is expected to have a similar 

effect, although perhaps not as much on 

anurans. 

Salamanders that breed in ponds and 

temporary wetlands are often explosive 

breeders, meaning that hundreds to 

multiple thousands of individuals will 

congregate at the same time (Gill 1978), 

creating dense numbers of individuals 

and increasing opportunities for the 

pathogen to spread. After breeding, the 

adults then return by land to their 

original habitats, potentially carrying 

the pathogen to new locations. 

Pathogens are also likely to be 

transmitted by salamander species that 

travel long distances for breeding and 

dispersal migrations, such as those that 

exhibit a metapopulation structure 

(Bancroft et al. 2011). A metapopulation 

is a group of discrete breeding 

populations of the same species (Gill 

1978). For example, within salamander 

metapopulations, California tiger 

salamanders have been documented 

traveling up to 1.2 miles (1.9 kilometers) 

from upland habitat to aquatic breeding 

sites (USFWS 2000), and newts travel 

many kilometers to breeding sites (Gill 

1978). 

Salamander species that have 

abundant populations with widespread 

distributions can also contribute to the 

spread of Bsal because of the increased 

likelihood that they will come in close 

contact with other salamanders that 

could then become infected. 

Salamanders that can carry Bsal from 

one place to another are more likely to 

do so if they have a broad range where 

they will come in contact with other 

members of the same species (for 

abundant distributions) or other species 

(for widespread distributions). Species 

with broad distributions are adapted to 

a wide range of environmental 

conditions that are more likely to 

overlap with habitat suitable for Bsal as 

well as habitat suitable for that species, 

providing increased opportunities for 

Bsal to spread. 

For example, the rough-skinned newt 

has a wide range along the West Coast 

from Alaska to California, and the 

eastern newt is found widely across the 

Eastern United States and Canada, 

occurring in 34 States (AmphibiaWeb 

2023a). Both species have had lethal 

responses with laboratory infections of 

Bsal (Martel et al. 2014; Carter et al. 

2021; Gray et al. 2023), and both are 

capable of carrying Bsal. In addition to 

its broad range, the eastern newt also 

migrates long distances; this species 

will frequently travel many kilometers 

to migrate to new ponds (Gill 1978), 

further increasing the risk of this species 

spreading Bsal. The eastern newt’s 

widespread distribution, high dispersal 

ability, high susceptibility, and 

juxtaposition with a high diversity of 

other salamander species position the 

species to significantly contribute to the 

spread of Bsal and the decline of 

salamander populations in the Eastern 

United States (Malagon et al. 2020). 

E. Pathway Analysis 

Introduction Pathways 

The main pathway for the global 

spread of Bsal is the international trade 

in salamanders (Martel et al. 2014; Yuan 

et al. 2018). The introduction of Bsal 

into mainland Europe is linked with the 

commercial trade of Asian salamanders 

(Cynops spp.) from East Asia, 

particularly Thailand, Vietnam, Japan 

(Martel et al. 2014), and China (Yuan et 

al. 2018). Combined, species from the 

genus Cynops were by far the most 

commonly imported into the United 

States from 2004 to 2014 (USFWS OLE 

2015), with Cynops orientalis alone 

comprising 54 percent of the 

salamander imports. Since the 2016 

interim rule went into effect, Cynops 

spp. imports have been restricted to 

those with approved permits from the 

Service. As described above in How the 

Fungus Affects Salamanders, there is no 

evidence that eggs and gametes are 

vectors. However, salamanders that 

have been identified as carriers, whether 

live or dead (except if chemically or 

heat preserved), appear capable of 

transmitting Bsal through contact with 

their skin (Gray et al. 2015; Van Rooij 

et al. 2015; Carter et al. 2020), which 

contains keratin (Seifert et al. 2019). We 

are also concerned that any infected and 

lethally vulnerable salamanders may die 

in transport and continue post-mortem 

to carry Bsal into the United States, or 

that salamanders may knowingly be 

imported dead. Bsal can remain viable 

inside dead host tissue (Martel et al. 

2013), although it is unclear how long 

a dead host remains infectious (WOAH 

2021b). One study found that viable 

Bsal loads on carcasses of eastern newts 

did not decline in 72 hours after 

euthanasia, and carcasses were capable 

of transmitting Bsal to susceptible hosts 

for at least 72 hours after death; the 

infections that developed in the 

susceptible animals caused nearly 100 

percent mortality in cohousing 

treatments that allowed for contact 

(Carter et al. in review). Therefore, we 

expect unpreserved, dead salamanders 

and body parts, except for purified 

extracted genetic material, to be a 

pathway for introduction. 

Individual amphibians in trade are 

often transported in containers with 

many other individuals of the same 

species or with many other species that 

can all be from different sources. These 


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conditions are highly conducive to 

pathogen transmission and dispersal. 

Pathogens can transfer from host to host 

in crowded conditions, and crowded 

conditions create stress on animals that 

can reduce amphibian hosts’ natural 

ability to resist infections (Rowley et al. 

2007, Rachowicz et al. 2005, RollinsSmith et al. 2011). Bsal can also be 

introduced into the environment 

through the improper disposal of 

contaminated water or other materials 

used to transport salamanders. As 

described above under Environmental 

Conditions Needed To Survive, the 

fungus can likely persist in those 

materials independent of the presence 

of a salamander. Water and fomites have 

served as a transmission medium and 

passive vector, respectively, to 

introduce other similar pathogens into 

the environment. For example, Bd has 

been found in water used to transport 

amphibians that were traded in Hong 

Kong (Kolby et al. 2014). As the authors 

noted, ‘‘[T]he abundance of aquatic 

amphibian species traded by Hong Kong 

* * *, prolonged environmental 

persistence of infectious * * * Bd 

particles, and employment of trade 

activities that neither disinfect water 

nor safely dispose of deceased animals 

creates an ideal pathway for disease 

transmission to native Hong Kong 

amphibians.’’ While experiments of 

fomite transmission for Bsal have not 

been conducted yet to our knowledge, 

Bd has been shown capable of infecting 

boreal toads (Bufo boreas) if an 

uninfected individual comes in contact 

with water where an infected individual 

has shed spores, even if the infected 

individual is no longer present (Carey et 

al. 2006). Similarly, encysted Bsal 

spores have been shown to remain 

infectious in pond water for at least 31 

days and capable of adhering to 

salamander skin and the feet of 

waterfowl (Stegen et al. 2017). 

Disinfecting of containers and 

substrate is always advisable when 

transporting amphibians. However, Van 

Rooij et al. (2017) found that the cell 

wall of the zoosporangium and the 

encysted spores within it provide a 

double barrier against the action of the 

disinfectants. This may explain 

(partially) why higher disinfectant 

concentrations or a longer contact time 

are necessary to achieve full fungal 

killing of Bsal, compared to those 

necessary for inhibition of Bd. Also, the 

clustering of multiple Bsal zoosporangia 

may protect centrally located sporangia 

from the full impact of a given 

disinfectant. 

Drawing on this evidence, the primary 

pathway for the entry of salamanders 

that are carriers of Bsal into the United 

States is through the international 

commercial wildlife trade. Overall, 99.9 

percent of live salamander importation 

into the United States is for commercial 

purposes (USFWS OLE 2021). From 

2010 to 2015, live salamanders were 

imported through 20 ports of entry into 

the United States; the 3 ports of entry 

with the largest numbers of imported 

salamanders were Los Angeles (54.6 

percent), Tampa (34.8 percent), and 

New York (6.8 percent). From 2016 to 

2020, live salamanders were imported 

through 8 ports of entry, with the top 4 

from Miami (63.3 percent), Boston (11 

percent), Newark (10.5 percent), and 

New York (9.2 percent). After import, 

many of the salamanders are transported 

to animal wholesalers, who then 

transport the salamanders to pet 

retailers. 

The most likely pathway of a 

salamander that is a host to Bsal into the 

United States would be through captive 

salamander commercial trade. We make 

that conclusion based on Martel et al. 

(2014), who noted that, given the 

discontinuity of the global distribution 

of Bsal, introduction from Asia into 

Europe must have been human 

mediated. Cunningham (2015), SabinoPinto et al. (2015), and Grear et al. 

(2021) also implicated trade as a key 

factor in the spread of Bsal. The United 

States is more isolated than European 

countries from other countries where 

salamanders could migrate 

independently, but there is an active 

trade in salamanders into the United 

States (USFWS OLE 2021), as discussed 

more in the following section (and 

would be more so without the second 

interim rule and final rules). People can 

purchase salamanders from pet stores or 

online retailers and keep them in 

captivity. Amphibians and reptiles kept 

as pets may eventually be released by 

their owners into the wild either 

intentionally or accidentally (Kraus 

2009, Krysko et al. 2011). For example, 

owners may no longer be able to care for 

their captives or an animal may escape 

its enclosure. Sick captive salamanders 

are often released instead of being 

euthanized; for example, around 200 

Japanese fire-bellied newts were 

released in the Miami, Florida, area in 

1964 when they became sick and 

unsuitable for sale (USGS NAS 2021 

[CYPY]). They died, but fortunately, this 

was before highly contagious amphibian 

pathogens were known to be in the 

United States. 

In addition to the risk from a release 

of an infected pet salamander into the 

wild, the water that is used to house an 

infected pet in captivity could feasibly 

contain Bsal zoospores. As a result, the 

discharge of untreated water used to 

house infected, captive animals could 

be a pathway for releasing infective 

zoospores into the environment and 

exposing native salamanders to Bsal 

(Stephen et al. 2015). A reduction in the 

transport of salamanders in trade would 

reduce the potential for contaminated 

water to carry spores to other areas. 

International Trade in Salamanders 

Trade in wildlife occurs on a global 

scale, and amphibians are some of the 

most commonly traded animals (Smith 

et al. 2009). More than 52,149,000 

documented amphibians were imported 

into the United States from 2004 to 

2014, based on the Service’s LEMIS data 

(USFWS OLE 2015), and 37,344,000 

were imported from 2010 to 2020 

(USFWS OLE 2021). Salamanders 

comprised 2,504,590 (4.8 percent) of the 

total imports of amphibians (USFWS 

OLE 2015) and 892,190 (2.4 percent) 

from 2010 to 2020 (USFWS OLE 2021). 

The 2004 to 2014 LEMIS dataset should 

be considered as a conservative estimate 

because many import records identified 

the animal being imported only as a 

member of the Class Amphibia (rather 

than identifying it to species or genus 

level). In addition, incorrect salamander 

identifications to genus and species 

level appear to have commonly 

occurred in reporting to LEMIS (USFWS 

OLE 2015). LEMIS data for 2004 to 2014 

shows that 65 percent of imported 

salamanders came from captive sources, 

and 35 percent were from wild sources 

(USFWS OLE 2015); for 2016 to 2020, 

46 percent came from captive sources, 

and 54 percent were wild caught 

(USFWS OLE 2021). The LEMIS data 

recorded only 83 percent of declared 

salamander imports at the species level, 

whereas 17 percent were recorded only 

to the genus level (USFWS OLE 2015); 

for 2010 to 2020, it was 95.5 percent to 

species and 6.5 percent to genus 

respectively (USFWS OLE 2021). 

The four salamander genera most 

commonly imported into the United 

States from 2004 to 2014 were Cynops, 

Paramesotriton, Triturus, and 

Pachytriton (USFWS OLE 2015). 

Cynops, Triturus, and Paramesotriton 

are three genera that are known to serve 

as carriers for Bsal (Martel et al. 2014). 

Of the 20 genera listed by the 2016 

interim rule, 18 had been traded (live or 

parts) over the 11 pre-interim rule years, 

and they comprised 95 percent of 

imported salamanders. From 2016 

through 2020, live imports of the top 

four salamander genera were 

Paramesotriton (524), Cynops (500), 

Bolitoglossa (191), and Pleurodeles 

(179); all but the Bolitoglossa were 

under injurious wildlife permits. 


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The species with the highest number 

of imports into the United States from 

2004 to 2014 was the Oriental firebellied newt. This species comprised 54 

percent of the total number of imported 

salamanders (USFWS OLE 2015). 

Twelve species of salamanders that are 

native to the United States were also 

imported into the United States from 

other countries from 2004 through 2014 

(USFWS OLE 2015). From 2016 through 

2020, 16 species of native salamanders 

were imported. Live eggs from the 

spotted salamander (Ambystoma 

maculatum), axolotl, and Japanese firebellied newt were imported between 

2016 and 2020. 

F. Risk Assessments of Bsal 

Bsal Risk Assessments 

Three Bsal risk assessments were used 

to help determine the risk associated 

with Bsal introduction into North 

America for the 2016 interim rule. 

Richgels et al. 2016 and Yap et al. (2015) 

conducted risk assessments for the 

United States that helped determine the 

level of risk associated with Bsal 

introduction. Stephen et al. (2015) also 

conducted a Bsal risk assessment for 

Canada that showed Canada is also at 

risk. 

Richgels et al. (2016) concluded that 

the potential for Bsal introduction into 

the United States is high, the United 

States has suitable conditions for Bsal 

survival, and the consequences of 

introduction into the United States are 

expected to be severe and occur across 

a wide range of the United States. To 

evaluate the potential for Bsal 

introduction, the assessment combined 

information on the number of 

individual salamanders imported at 

each port of entry and the number of 

pet-supply establishments by county. 

Based on this evaluation, Bsal 

introduction potential was highest in 

central and southern Florida, southern 

California, and near New York City, 

New York (Richgels et al. 2016). 

As noted in Richgels et al. (2016), the 

areas of highest potential for Bsal 

introduction are not necessarily the 

same as the areas of greatest risk for 

impacts to wildlife and wildlife 

resources. To determine the 

consequences of Bsal introduction into 

the United States, including from the 

areas where the introduction potential 

was highest, Richgels et al. (2016) 

evaluated environmental suitability, 

spatial data on imports and pet trade 

activity, species richness, and predicted 

species susceptibility. Overall, the total 

risk of Bsal to native salamanders is 

high. While not all areas of the United 

States are at risk from Bsal, based on 

both likely introduction and resultant 

consequences, the risk of Bsal is highest 

for the Pacific coast, southern 

Appalachian Mountains, and midAtlantic regions (Richgels et al. 2016). 

Some areas, such as south Florida and 

parts of the West, are likely to have low 

consequences from Bsal introduction. 

The areas most likely to have 

consequences from Bsal introduction 

are the Pacific Coast and Appalachian 

Mountains (Richgels et al. 2016). Based 

on environmental suitability, areas of 

the United States most suited to Bsal 

growth (Blooi et al. 2015a), including 

the Southwest, Southeast, and Pacific 

regions, are also the areas of highest 

salamander diversity (Richgels et al. 

2016). 

In the United States, Yap et al. (2015) 

identified the Southeastern (southern 

end of the Appalachian Mountains and 

neighboring southeast region) and 

Western United States (Pacific 

Northwest and the Sierra Nevada) as 

zones of high risk. Yap et al. (2015) 

identified a narrower total risk in the 

United States over a smaller area for 

Bsal compared to Richgels et al. (2016). 

For example, Yap et al. (2015) identified 

south Florida as low risk of Bsal 

vulnerability, while Richgels et al. 

(2016) found that there was some risk, 

if not the highest, of Bsal to native 

salamanders in Florida. Richgels et al. 

(2016) noted differences in the methods 

used between the two papers as the 

reason. For example, Richgels et al. 

(2016) uses the thermal range of Bsal 

rather than the native Asian host 

distribution applied in Yap et al. (2015), 

which Richgels et al. (2016) noted may 

not be vulnerable to infection across the 

entire range of those species. The model 

in Richgels et al. (2016) took 

environmental suitability into account 

but also used species diversity, 

proximity to ports of entry, and areas of 

high pet-trade activity to predict total 

risk. This approach may over- or 

underestimate risk for some areas. 

Some salamander species may be 

protected from Bsal by temperatures in 

their regions that are outside of the Bsal 

optimal growth range (Richgels et al. 

2016) (see Climate Tolerance section). 

However, the average temperature 

preferences of salamanders from Central 

and North America (Duellman and 

Trueb 1986), which range from ¥2.0 °C 

(28.4 °F) to 30.0 °C (86.0 °F), suggest that 

most salamander species, including 

those within the United States, are 

active near the thermal growth optimum 

for Bsal (Blooi et al. 2015a). As a result, 

most salamander species in the United 

States are not protected from Bsal by 

living outside of the Bsal optimal 

growth range or in areas beyond the 

threshold where Bsal can survive. 

Most U.S. salamander species are also 

dependent upon forests, a habitat type 

dominated by relatively cool, moist 

conditions, for the majority of their life 

cycle (Davic and Welsh 2004). It is 

possible that cool seasons or 

microclimate selection by salamanders 

could facilitate disease outbreaks in 

areas where the average temperatures 

are outside the preferred range of the 

fungus. 

A fourth risk assessment was added 

for this second interim rule, led by the 

USGS (Grear et al. 2021). They used 

post-interim rule outcomes and new 

information to update the Richgels et al. 

(2016) assessment. Grear et al. (2021) 

evaluated the effects of the 2016 interim 

rule on the introduction of Bsal into the 

United States and reviewed new 

information on species susceptibility to 

reevaluate the risk of Bsal to the United 

States. Since no comprehensive 

surveillance for Bsal was available prior 

to the 2016 interim rule, they also used 

the results from Waddle et al. (2020) of 

their large-scale surveillance for Bsal 

across 594 counties in 35 States and 1 

site in Mexico, with 11,189 swab 

samples of wild salamanders and some 

anurans. The surveillance sites were 

strategically chosen for highest risk 

based on species susceptibility and 

geography. The surveillance did not 

detect any Bsal, which was as hoped, 

but the surveillance plan they 

developed can continue to be used for 

early detection of Bsal at high-risk 

locations. 

Grear et al. (2021) also found that the 

2016 interim rule reduced the 

importation of listed genera by several 

orders of magnitude, which concurs 

with our results. They noted several 

additional genera of salamanders and 

several of anurans (particularly 

Bombina spp.) that could be carriers of 

Bsal but were not listed in 2016; the 

additional salamander genera are 

included for listing in this interim rule, 

while the anurans are not (see Impacts 

on Wildlife Resources or Ecosystems for 

explanation why). Among other 

variables, Grear et al. (2021) used 

updated information about the thermal 

tolerance of Bsal that included water 

temperatures associated with detection 

of Bsal in its presumed native 

geographic and host ranges. They noted 

that the change and spatial variation in 

risk scores from considering a higher 

maximum-temperature threshold had no 

discernible net effect on the 

consequence score. Therefore, they 

chose not to use the temperature 

threshold in recalculating the 

consequence and total-risk estimation; 


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instead, they focused their risk 

comparisons on import regulations, 

host-species range, and surveillance. 

The study looked at the change in ports 

of import between pre- and post-interim 

rule and calculated the risk change for 

those regions. They concluded that, 

while the import regulations mitigated 

some of the import risk, overall risk is 

still driven by the potential of release 

from undetected Bsal circulating in 

captive amphibians in the United States 

and the consequences of that release in 

high salamander-biodiversity areas; 

however, little information is known on 

movement of captive host species and 

possible undetected Bsal in U.S. 

captives. 

Vulnerability and Carrier Status 

The urgent need to prevent Bsal 

introduction risks with the 2016 interim 

rule was raised by evidence presented 

by Martel et al. (2014), who tested Bsal 

on 35 species from all 3 orders of 

amphibians: frogs, salamanders, and 

caecilians. Martel et al. (2014) further 

screened 5,391 specimens collected 

from 4 continents for evidence of Bsal 

infection. Martel et al. (2014) defined a 

‘‘resistant’’ salamander as one that 

either was not infected or developed a 

short-term infection without clinical 

signs following exposure to Bsal; a 

‘‘tolerant’’ salamander is one that 

maintains a more prolonged infection 

with no signs of disease; a ‘‘susceptible’’ 

salamander becomes infected and has 

clinical signs of disease with the 

possibility of subsequent recovery; and 

a salamander that responds in a ‘‘lethal’’ 

manner to Bsal dies as a result of 

infection. According to Martel et al. 

(2014), resistant salamanders are not a 

risk for transmitting Bsal. However, 

based on the available scientific data, 

we concluded that resistant species with 

evidence of short-term or transient 

infection, as well as those reported to 

have tolerant, susceptible, or lethal 

responses to Bsal, are carriers capable of 

transmitting Bsal to other salamanders 

and introducing the fungus into the 

United States. 

The Service considered a species to be 

a ‘‘noncarrier’’ when Martel et al. (2014) 

classified the species as ‘‘resistant’’ and 

no histologic or field surveillance data 

was found to suggest that short-term 

Bsal infection could occur; 

‘‘noncarriers’’ were considered 

incapable of transmitting Bsal to other 

salamanders or introducing the fungus 

into the United States. If Martel et al. 

(2014) classified the species as resistant 

with no histology (or qPCR) to verify, its 

carrier status was inconclusive. We use 

this same definition in this second 

interim rule. 

We also found, and still maintain, that 

the likelihood of species within the 

same genus as being carriers can be 

drawn from a comparison to Bd, which, 

as described above under General 

Information About Bsal, is a close 

relative of Bsal. As noted earlier, the 

two risk assessments of Bsal used Bd in 

determining the risk of Bsal based on 

transmission, spread, and populationlevel effects (Richgels et al. 2016; 

Stephen et al. 2015). Considerably more 

was known about Bd than Bsal due to 

its discovery and description more than 

15 years earlier (Berger et al. 1998; 

Longcore et al. 1999), while Bsal was 

discovered in 2013 (Martel et al. 2013). 

Bd has caused amphibian declines and 

extinctions worldwide (Skerratt et al. 

2007). Bd affects species in patterns 

(Skerratt et al. 2007), and more closely 

related species have similar outcomes 

for Bd at the family level (Smith et al. 

2009; Bancroft et al. 2011). 

Amphibians experiencing the most 

severe declines are grouped by 

relatedness, which is likely due to the 

shared evolutionary histories of closely 

related species with a similar response 

to chytridiomycosis (Corey and Waite 

2008). The USDA uses a similar 

approach. Closely related species are 

considered more likely to have similar 

traits and are used in risk assessments 

to determine threats from a target 

species of interest; a potential pest is 

regarded as a threat when other species 

in a genus pose a similar threat 

(Wapshere 1974; Gilbert et al. 2012). 

The European Union’s study on the 

feasibility of a movement ban of traded 

salamanders concluded that, due to the 

complexity of the taxonomy as well as 

the lack of evidence related to all the 

species, a movement ban at the level of 

taxonomic order is likely to be more 

effective and more feasible than a 

species-specific ban (EFSA et al. 2017). 

Many salamanders exhibited a strong, 

adverse response to experimental Bsal 

infection; many species from outside of 

the native range of the fungus (Asia) 

exhibited lethal vulnerability. For the 

2016 interim rule, our review of Martel 

et al. (2014) and follow-up 

communication (A. Martel, University 

of Maryland, pers. comm. 2015) 

categorized 25 species from 19 genera as 

carriers of Bsal. Additional 

communications (Chytridcrisis 2015b; 

Cunningham et al. 2015; P. Nanjappa, 

Association of Fish and Wildlife 

Agencies, pers. comm. 2015) identified 

another two species from two separate 

genera as carriers: the pygmy marbled 

newt (Triturus pygmaeus) and the 

golden striped salamander (Chioglossa 

lusitanica). Because Martel et al. (2014) 

had previously identified members of 

the Triturus genus as carriers, it was 

already accounted for within the 19 

genera. The addition of this species 

brought the total number of known 

carrier species to 26. In addition to 

Triturus, Chioglossa was identified as 

another genus capable of serving as a 

Bsal carrier by Chytridcrisis (2015b), 

Cunningham et al. (2015), and P. 

Nanjappa (Association of Fish and 

Wildlife Agencies, pers. comm. 2015). 

As a result, the total number of genera 

known to serve as carriers of Bsal was 

20 genera, and these were Chioglossa, 

Cynops, Euproctus, Hydromantes, 

Hynobius, Ichthyosaura, Lissotriton, 

Neurergus, Notophthalmus, 

Onychodactylus, Paramesotriton, 

Plethodon, Pleurodeles, Salamandra, 

Salamandrella, Salamandrina, Siren, 

Taricha, Triturus, and Tylototriton. 

Further studies since the 2016 interim 

rule have included species from some of 

the same genera listed in 2016, which 

we reviewed to see if they were 

consistent with earlier conclusions. 

Carrier status is further supported for 

Chioglossa (C. longipes and C. 

lusitanica in Bosch et al. 2021); 

Lissotriton (L. boscai, Fitzpatrick et al. 

2018, Bosch et al. 2021; L. lusitanica, 

Lastra Gonza ́laz et al. 2019), Triturus (T. 

dobrogicus, T. ivanbureschi, T. karelinii, 

and T. marmoratus, Fitzpatrick et al. 

2018; T. marmoratus, T. pygmaeus, 

Bosch et al. 2021), Neurergus 

(Fitzpatrick et al. 2018), Notophthalmus 

(Fitzpatrick et al. 2018; Gray et al. 2023), 

Plethodon (P. cinereus, P. cylindraceus, 

P. glutinosus, P. montanus, and P. 

shermani, DiRenzo et al. 2021), 

Salamandra (S. atra, Fitzpatrick et al. 

2018), Siren (S. lacertina, Gray et al. 

2023), and Taricha (T. granulosa, T. 

torosa, Gray et al. 2023). Studies for 

Chioglossa (Fitzpatrick et al. 2018), and 

Plethodon (Perreira and Woodley 2021) 

showed no conclusive countervailing 

evidence. 

Using the same criteria as in the 2016 

interim rule, this second interim rule 

adds the following 16 genera: 

Ambystoma, Andrias, Aneides, 

Aquiloeurycea, Calotriton, 

Chiropterotriton, Cryptobranchus, 

Desmognathus, Ensatina, Eurycea, 

Laotriton, Ommatotriton, Pachytriton, 

Proteus, Pseudobranchus, and 

Pseudotriton. This increases the total 

number of species listed by 

approximately 164. 

In conducting our analysis, the 

Service initially focused on identifying 

species for listing as injurious that 

scientific evidence demonstrates are 

capable of carrying Bsal. As we 

described above, we find that, due to 

shared characteristics by species within 

a genus, other species within these 


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genera are also highly likely to be 

carriers of Bsal, even if not every species 

in the genus has been tested to verify 

that it is a carrier of Bsal. This 

conclusion is because closely 

phylogenetically related species, such as 

those found within the same genus, 

share common traits. Our analysis found 

no conclusive evidence to the contrary 

that suggested that some species within 

such genera are not carriers. We have 

focused our findings on taxa of 

salamanders and their genera that we 

determined to be capable of carrying 

Bsal. We included genera identified as 

resistant by Martel et al. (2014) because 

carrier status was inconclusive in that 

study when histology was not done, and 

there are other studies supporting 

carrier status; these are Ambystoma, 

Hynobius, Lissotriton, and Plethodon 

(see Vulnerability and Carrier Status of 

Native Species below). Based on our 

analysis of their data and other studies, 

we have no evidence for the salamander 

genera we are not listing of being 

capable of introducing Bsal to the 

United States or otherwise transmitting 

Bsal to native populations. In addition, 

we are not listing genera where there 

was no data as of the drafting of this 

second interim rule because we do not 

have a basis for doing so, even though 

the Service recognizes that it is possible 

that untested genera may also be 

capable of carrying Bsal. 

We have determined that all species 

are injurious in the 16 genera where at 

least one species has been conclusively 

identified as a carrier of Bsal and there 

is no conclusive countervailing 

evidence suggesting that some species 

within the genus are not carriers. Where 

one species has been identified as a 

carrier, we expect that the other species 

in that genus are also carriers. This 

finding includes as injurious the 

intrageneric hybrids (crosses of species 

found within the same genus), 

intergeneric hybrids of species in two 

listed genera, and intergeneric hybrids 

from a listed and an unlisted genus. 

For this second interim rule, we 

maintain that, due to shared 

characteristics by species within a 

genus, other species within these genera 

are also likely to be carriers of Bsal if 

one species has been identified as a 

carrier, even if not every species in the 

genus has been tested to verify that it is 

a carrier of Bsal. Our updated review 

found no conclusive countervailing 

evidence that species differed within a 

genus with respect to their ability to act 

as carriers. Thus, we expect all species 

in a genus to respond similarly as 

carriers or noncarriers to Bsal. 

Therefore, based on existing scientific 

evidence, and as described in more 

detail below, we are listing all species 

in the 16 genera, including all species, 

that we now conclude constitute a 

threat to introducing and spreading Bsal 

in the United States because those 

species can carry the fungus and 

transmit it to other species that would 

be negatively impacted. 

Vulnerability and Carrier Status of 

Native Species 

Including both the final rule to the 

2016 interim rule and this second 

interim rule, we conclude that 

approximately 426 salamander species 

from around the world are carriers of 

Bsal (36 genera in 7 families). The 

United States currently has 

approximately 221 species of native 

salamanders in 23 genera 

(AmphibiaWeb 2023a), and this second 

interim rule includes 164 of those 

species (13 genera in 5 families) that we 

have determined are carriers of Bsal. Of 

the remaining 57 native species, we find 

that either they are not carriers or the 

vulnerability and carrier status is 

unknown. 

Of the 190 native U.S. salamander 

species as of the 2016 interim rule, 

carrier status had not been assessed in 

103 species from 16 genera. The 

untested genera were Amphiuma, 

Aneides, Batrachoseps, Cryptobranchus, 

Desmognathus, Dicamptodon, Ensatina, 

Eurycea, Hemidactylium, Necturus, 

Phaeognathus, Pseudobranchus, 

Pseudotriton, Rhyacotriton, 

Stereochilus, and Urspelerpes. 

Since the 2016 interim rule went into 

effect, we have evidence that eight more 

native genera, not previously tested, 

support listing with this second interim 

rule: Ambystoma, Aneides, 

Cryptobranchus, Desmognathus, 

Ensatina, Eurycea, Pseudobranchus, 

and Pseudotriton. We previously 

considered Ambystoma as resistant 

because Martel et al. (2014) had done so 

for two species, and Bsal was not 

detected during testing. However, 

Martel et al. did not perform histology 

on the Ambystoma subjects because 

they did not die, so it was undetermined 

if the individuals harbored encysted 

zoospores. 

Since the 2016 interim rule, initial or 

additional testing has been done on 

Ambystoma spp. (Fitzpatrick et al. 2018; 

Sabino-Pinto et al. 2018; Barnhart et al. 

2020; Gray et al. 2023); Aneides aeneus 

(Gray et al. 2023); Cryptobranchus (Gray 

et al. 2023); Desmognathus ocoee (Gray 

et al. 2023); Ensatina eschscholtzii (Gray 

et al. 2023); Eurycea lucifuga and E. 

wilderae (Carter et al. 2020), E. wilderae 

(DiRenzo et al. 2021), and E. bislineata, 

E. lucifuga and E. wilderae (Gray et al. 

2023); Pseudobranchus striatus (Gray et 

al. 2023); and Pseudotriton ruber (Carter 

et al. 2020) that provides support for 

carrier status of these native genera. 

There is no conclusive countervailing 

evidence. Five more native species were 

found to be lethally affected by Bsal 

(Eurycea bislineata, E. wilderae, 

Pseudotriton ruber, Ensatina 

eschscholtzii, Aneides aeneus). Gray et 

al. (2023) tested only native North 

American species specifically to access 

the conservation risk to U.S. species. 

Based on the gradient responses from 

resisting infection to lethal response 

among the genera Martel et al. (2014) 

and others tested experimentally, other 

genera could be at risk from Bsal 

infection or could serve as carriers. 

However, we are not listing species in 

those genera because the genera had not 

yet been tested or confirmed as carriers 

by the drafting of this second interim 

rule. 

Controlled Bsal experiments have 

proliferated since the discovery of the 

fungus. A study by Kumar et al. (2020) 

shows variation in experimental 

methodologies could thwart knowledge 

advancement by introducing 

confounding factors that make 

comparisons difficult among studies. 

They tested whether passage duration of 

Bsal culture (the number of times the 

fungus was transferred from its culture 

into fresh culture media), exposure 

method of the host to Bsal (water bath 

versus skin inoculation), Bsal culturing 

method (liquid versus plated), host 

husbandry conditions (aquatic versus 

terrestrial), and skin-swabbing 

frequency influenced diseased-induced 

mortality in a susceptible host species, 

the eastern newt. They found that 

disease-induced mortality was faster for 

eastern newts when exposed to a low 

passage isolate (a ‘‘young’’ Bsal isolate 

that had been passed into fresh culture 

media only 20 times), when newts were 

housed in terrestrial environments, and 

if exposure to zoospores occurred in a 

water bath. They did not detect 

differences in disease-induced mortality 

between culturing methods or swabbing 

frequencies. Their results illustrate the 

need to standardize methods among 

Bsal experiments, but they do not 

discount the results of the studies used 

to determine our results. 

Vulnerability and Carrier Status of 

Threatened and Endangered Species 

As of the drafting of this interim rule, 

20 native species of salamanders in 6 

genera are threatened or endangered 

under the ESA. As of the drafting of the 

2016 interim rule, none of the 

salamander species listed as endangered 

or threatened under the ESA in the 

United States had been specifically 


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tested for Bsal vulnerability under 

laboratory conditions. Bsal had not been 

detected in their wild populations 

(Martel et al. 2014, Bales et al. 2015). 

Since publication of the 2016 interim 

rule, several species have been 

laboratory-tested. 

One species with two federally 

endangered subspecies (eastern 

hellbender Cryptobranchus a. 

alleghaniensis and Ozark hellbender C. 

a. bishopi) has been laboratory tested 

and is considered a carrier in the second 

interim rule in this document (Gray et 

al. 2023). Notably, Cryptobranchus has 

only one species in the genus, so if the 

species is extirpated by Bsal, so is the 

genus. 

As we describe above in Vulnerability 

and Carrier Status, while the Service 

did find evidence that shows some 

species within a genus may vary in their 

specific vulnerability, the carrier status 

of tested species can be extrapolated to 

related species, including those that are 

listed as endangered or threatened, 

candidates for ESA listing, and under 

review. 

Of the other new genera that include 

native species that we have identified as 

carriers, the following 12 species are 

federally listed as endangered or 

threatened: 5 species of Ambystoma 

(California tiger salamander (A. 

californiense), frosted flatwoods 

salamander (A. cingulatum), reticulated 

flatwoods salamander (A. bishopi), 

Sonoran tiger salamander (A. 

mavortium stebbinsi), Santa Cruz longtoed salamander (A. macrodactylum)); 

and 7 species of Eurycea (Austin blind 

salamander (E. waterlooensis), Barton 

Springs salamander (E. sosorum), 

Georgetown salamander (E. naufragia), 

Jollyville Plateau salamander (E. 

tonkawae), Salado salamander (E. 

chisholmensis), San Marcos salamander 

(E. nana), and Texas blind salamander 

(E. rathbuni)). Notably, Ambystoma is 

the only genus in the family 

Ambystomatidae, so if the genus is 

extirpated by Bsal, so is the family. 

No information is available regarding 

the effect of Bsal or carrier status of the 

remaining four ESA-listed species 

native to the United States: the desert 

slender salamander (Batrachoseps 

aridus), the Alabama waterdog or black 

warrior waterdog (Necturus 

alabamensis), Neuse River waterdog (N. 

lewisi), and Red Hills salamander 

(Phaeognathus hubrichti). Three 

Plethodon species from the 2016 interim 

rule are federally listed as endangered 

or threatened: Shenandoah salamander 

(P. shenandoah), Cheat Mountain 

salamander (P. nettingi), and Jemez 

Mountains salamander (P. 

neomexicanus) (USFWS 2023). There 

were no candidate species of 

salamanders as of the drafting of this 

second interim rule. Three Plethodon 

species identified as carriers in the 2016 

interim rule remained federally listed. 

G. Factors That Contribute to 

Injuriousness of Salamanders 

Likelihood of Release or Escape (of 

Salamanders) 

In general, there is widespread 

concern over the increasing spread of 

pathogens moved through the wildlife 

trade (for example, Chinchio et al. 2020; 

IPBES 2020). Substantial evidence 

shows that Bd has spread extensively 

throughout the world through the 

amphibian trade (Fisher and Garner 

2007; Schloegel et al. 2009; Schloegel et 

al. 2012; Galindo-Bustos 2014; Kolby 

2014; Kolby et al. 2014). Similar 

mechanisms of transmission and 

persistence in the closely related Bsal 

pathogen, along with detection of Bsal 

in captive salamanders imported by the 

pet trade into Great Britain, indicate that 

global movement of Bsal, similar to that 

of Bd, is not only possible but is already 

occurring (Cunningham 2015). Bsal was 

also found in a private pet collection in 

Germany, where it killed over half of a 

collection of approximately 200 

salamanders in the genus Salamandra 

(Sabino-Pinto et al. 2015). Although the 

origin of Bsal in the German collection 

was unknown, it is probable that Bsal is 

also present in other private or 

professional collections across Germany 

and possibly also in other European 

countries (Sabino-Pinto et al. 2015). 

Amphibian trade fairs in Spain, where 

the largest fairs in southern Europe take 

place, as well as in private collections 

in Spain, had positive test results for Bd 

and Ranavirus (Thumsova ́ et al. 2021) 

and are known to house and co-house 

sick amphibians. These collections may 

serve as a reservoir of Bsal within the 

wildlife trade or as sources of Bsal 

release into the environment. 

Considering the occurrence of Bsal in 

the global pet trade, the risk to North 

American native species, and the 

number of salamanders that are 

imported into and transported 

throughout the United States through 

trade, Bsal is likely to be introduced 

into and spread throughout native 

salamander populations in the United 

States unless immediate action is taken 

to limit the importation of salamanders 

that are likely to carry Bsal. The 2016 

interim rule has limited importation, 

and this second interim rule is intended 

to further reduce risk. 

Infected salamanders can transmit 

Bsal to other species even if the 

introduced salamander fails to establish 

a population. Evidence indicates that at 

least some of the salamanders capable of 

carrying Bsal can escape or be released 

and introduce Bsal into the 

environment. As described earlier, 

evidence exists for release of 

salamanders into the wild in the United 

States (Picco and Collins 2008; USGS 

2015a, b, c, d, e, f). As noted above in 

Invasiveness of Salamanders, the 

USGS’s Nonindigenous Aquatic Species 

database (USGS 2023) has records for 17 

salamander species that have been 

observed in the environment outside 

their native range. Of those, 14 are 

native to the United States and were 

discovered outside of their native 

ranges, and 3 are species not native to 

the United States. These findings mean 

that salamanders have been shown to 

exist, even if temporarily, outside their 

native range in the environment. Thus, 

they are capable of transmitting Bsal 

into nonindigenous ecosystems. 

Infected native species that are imported 

and escape or are released into native 

habitats would also be capable of 

carrying Bsal into native salamander 

ecosystems where Bsal has not 

previously been found. 

Infectious Bsal zoospores can also be 

released into the environment if water 

or other materials used to house 

infected salamanders enter the 

environment due to improper 

disinfection and disposal methods. The 

water and materials become passive 

carriers to introduce the fungus into the 

environment if not decontaminated or 

disposed of properly. As described 

above under Environmental Conditions 

Needed To Survive, Bsal can survive in 

filtered pond water for at least 31 days 

(Stegen et al. 2017). Bd is similarly 

known to remain viable for weeks in 

water and moist organic matter and is 

capable of being transmitted to 

uninfected specimens through such 

means. Given our assumption that Bd 

can serve as a surrogate for predicting 

Bsal’s effects in salamanders at the 

population level, and since Bd does not 

require an amphibian host to remain 

viable, we expect that Bsal can also 

persist outside salamanders (as long as 

it has sufficient water or moist soil and 

conducive temperature). Since the 

effects of desiccation or the viability of 

encysted Bsal spores in deceased hosts 

have not been thoroughly investigated, 

we also expect that Bsal can be 

transmitted on unpreserved dead 

salamanders or body parts and tissues. 

As discussed above in Introduction 

Pathways, there is evidence that Bd has 

escaped into the environment through 

untreated wastewater, increasing the 

likelihood that Bsal could also escape if 

brought in via contaminated water or 


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improperly disposed of materials. While 

standards for the treatment and 

prevention of Bd exist, in part due to 

recognition of its status as an 

internationally notifiable disease under 

the World Organisation for Animal 

Health (WOAH), the effectiveness and 

widespread application of those 

standards are uncertain given that 

international protocols for responding to 

Bd do not exist and the need to improve 

international mechanisms to respond to 

disease-related threats to biodiversity 

(Voyles et al. 2014). 

Given the number of specimens that 

have been imported into the United 

States and Canada, it is not known why 

Bsal has not yet been found in these 

countries (Muletz et al. 2014; Bales et al. 

2015; Stephen et al. 2015; Richgels et al. 

2016). A comparison of Bd, which has 

spread in the United States, to Bsal 

yields some insights. Based on genetic 

analyses and examination of historical 

specimens, Bd may have originated 

from different places, including Japan, 

South Africa, or South America (Farrer 

et al. 2011; Rodriguez et al. 2014). In 

contrast, Bsal may have originated only 

from Asia, giving it fewer pathways to 

the United States (Martel et al. 2014; 

Laking et al. 2017). Importation of 

salamanders into the United States has 

also declined in recent years, suggesting 

that the propagule pressure may also be 

a factor by limiting the number of times 

in which Bsal could possibly be 

introduced into the environment 

through trade (Lockwood et al. 2005; 

USFWS OLE 2015). Bd may have spread 

more quickly than Bsal because of its 

ability to infect frogs, whereas research 

so far has found only a few frog species 

that may carry Bsal (see Impacts on 

Wildlife Resources or Ecosystems 

below). Based on LEMIS data, frogs are 

traded in higher volumes than 

salamanders, increasing the probability 

of trade of a Bd-infected individual over 

a Bsal-infected individual. The USGS 

Nonindigenous Aquatic Species 

database also provides evidence for this 

higher level of trade, in that greater 

numbers of frogs are reported than 

salamanders. In addition, many frogs in 

trade, such as Rana catesbeiana 

(bullfrogs), are adaptable to a wide 

variety of environments and can easily 

become invasive once released in a 

watershed, as bullfrogs have become in 

the American West (Jennings and Hayes 

1994; Rosen and Schwalbe 1995; Funk 

et al. 2011; Sepulveda et al. 2015). 

Taken together with the other data we 

reviewed, this evidence suggests that 

Bsal is less likely to enter the United 

States than Bd. However, without 

action, the pathways for introduction 

and escape of Bsal are a significant and 

imminent threat. Listing salamanders 

that can carry Bsal as injurious wildlife 

to prohibit their importation targets 

those pathways, thereby minimizing 

opportunities for Bsal to be introduced, 

become established, and spread in the 

United States. 

Potential To Survive, Become 

Established, and Spread 

Even if a salamander species does not 

become established, there is evidence 

that it may be capable of carrying Bsal 

long enough in the wild to transmit 

Bsal. The USGS Nonindigenous Aquatic 

Species database has records of 17 

species and populations that have been 

observed in the United States outside of 

their native range (USGS 2023). Of 

those, 14 are native and have 

established populations in the United 

States outside of their native U.S. range: 

Eastern tiger salamander (Ambystoma 

tigrinum), northwestern salamander (A. 

gracile), blotched tiger salamander 

(Ambystoma mavortium 

melanostictum), long-toed salamander 

(Ambystoma macrodactylum), threetoed amphiuma (Amphiuma 

tridactylum), California slender 

salamander (Batrachoseps attenuatus), 

seal salamander (Desmognathus 

monticola), Santeetlah dusky 

salamander (Desmognathus santeetlah), 

black-bellied salamander 

(Desmognathus quadramaculatus), 

mudpuppy, eastern newt, red-spotted 

newt (Notophthalmus viridescens 

viridescens), large-blotched ensatina 

(Ensatina eschscholtzii klauberi), lesser 

siren (Siren intermedia), and roughskinned newt (Taricha granulosa). The 

three species from outside the United 

States are the Japanese fire-bellied newt 

(Cynops pyrrhogaster), Oriental firebellied newt (Cynops orientalis), and 

paddle-tailed newt (Paramesotriton 

(Pachytriton) labiatus), none of which 

are known to have become established. 

No foreign terrestrial salamander 

species have been detected in USGS 

surveillance for Bsal (M. Adams, USGS, 

pers. comm. 2021). As discussed earlier 

under Introduction Pathways and 

Environmental Conditions Needed To 

Survive, Bsal is expected to be able to 

survive outside of salamander hosts for 

several weeks given suitable conditions 

in water. If a salamander comes in 

contact with Bsal and then transmits it 

during a time when salamanders 

congregate, such as during breeding as 

described above under Salamander 

Biology, the potential for Bsal to survive, 

establish, and spread through animals or 

animal parts (except for purified, 

extracted genetic material, eggs, and 

gametes) is significant. As we describe 

above under How the Fungus Affects 

Salamanders, Bsal can be transmitted 

on unpreserved dead tissue where 

keratin is present, particularly skin, but 

we do not find that Bsal can be 

transmitted through reproductive tissue, 

including eggs and gametes. There is no 

evidence to suggest that Bsal can 

survive in purified, extracted genetic 

material from salamanders; in 

chemically preserved specimens, 

tissues, samples, or swabs; or in 

salamander eggs and gametes; hence, 

these parts are not covered by the 

listing. 

As Richgels et al. (2016) noted, 

‘‘Given the large number of suspected 

Bsal carriers imported into the USA 

each year (Cynops spp. and 

Paramesotriton spp., more than 100,000 

[per year]), Bsal is likely to be 

introduced if no additional risk 

mitigation steps are taken. Though 

precise estimates for the invasion 

process (proportion of imported 

individuals infected, frequency of 

release of captive individuals, and 

contact of released animals with native 

amphibians) do not exist for Bsal, the 

establishment of invasive amphibians 

common in US amphibian trade * * * 

and the patterns of global Bd spread 

* * * suggest these processes are also 

likely for Bsal.’’ The Service finds that 

the capacity of infected salamanders in 

trade to potentially infect wild 

salamanders, together with the capacity 

of Bsal to survive for an extended period 

independent of an amphibian host, 

suggests that Bsal has a high likelihood 

of surviving, becoming established, and 

spreading once it is introduced into a 

new area. 

As we noted above in Purpose of 

Listing as Injurious, even if a 

salamander found to be injurious could 

not establish a population in the wild, 

an infected or carrier salamander from 

captivity can still transmit Bsal to native 

populations if that salamander escapes 

or if material touching it is improperly 

disposed. 

Impacts on Wildlife Resources or 

Ecosystems 

If Bsal is introduced into the United 

States, we expect the species with lethal 

vulnerability would be at greatest risk. 

However, disease outbreaks can result 

from a combination of biotic and abiotic 

factors, including species vulnerability, 

exposure, host behavior, host immunity, 

co-infections, and environmental 

conditions (Wobeser 2007). Therefore, 

the vulnerability of individuals under 

laboratory conditions is an incomplete 

predictor of disease effects (Wobeser 

2007). Native salamander species 

known to be tolerant of Bsal infection 

under experimental conditions may 


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demonstrate more severe clinical 

disease when infection is combined 

with additional stressors in the wild, as 

has been found for other diseases, 

including those in amphibians (Wobeser 

2007; Kerby et al. 2011; Kiesecker 2011). 

For example, Bodinof et al. (2011) noted 

that Bd may be found more frequently 

in hellbenders that are 

immunocompromised or that Bd 

infection increases the adverse effects of 

other co-infections. Considering these 

cumulative factors, as well as the lack 

of testing for the majority of native 

salamander species, our assessment of 

risk in native species is likely 

conservative. 

Bsal can severely affect wildlife 

resources. At least nine native species 

are lethally vulnerable to Bsal, and at 

least one is tolerant to Bsal infection. At 

least 164 native species may act as 

carriers or sources of infection for other 

species. While not all species have been 

tested for their response to Bsal, based 

on the high rates of infection that have 

been observed, the fungus may have 

significant negative effects on additional 

species. 

As described above in EcosystemLevel Effects, salamanders are important 

parts of the ecosystems in which they 

occur. They are often the most abundant 

vertebrates in their ecosystems, and, as 

a vital part of the food web, they are 

both important prey for and predators of 

many species (Holomuzki et al. 1994; 

Regester et al. 2006). In some places, 

they are considered keystone species 

that help control some invertebrate 

populations and affect cycling of 

nutrients in an ecosystem, contributing 

significantly to overall ecosystem 

health. For example, by consuming 

arthropods that would otherwise release 

carbon dioxide into the atmosphere by 

decomposing leaf litter in forests, 

salamanders slow carbon emissions 

from leaf litter decomposition, which 

has implications for the global carbon 

cycle (Best and Welsh 2014). As 

described earlier, invertebrate species 

that depend on salamanders for aspects 

of their life cycle or ecology are likely 

to be adversely affected if their host 

species declines in response to a Bsal 

introduction. Loss of these keystone 

species would result in significant 

ecosystem-level change. 

Salamanders constitute much of the 

vertebrate biomass of forests, and they 

play an important role in ecosystems as 

insect consumers, shapers of the 

landscape, and climate mediators 

(Burton and Likens 1975; Davic and 

Welsh 2004; Wyman 1998; Best and 

Welsh 2014). If native U.S. salamander 

species do experience declines from 

Bsal infection as the fire salamander 

experienced in the Netherlands 

(Spitzen-van der Sluijs et al. 2013), we 

expect detrimental ecological effects. 

Nine native salamanders are 

documented as lethally vulnerable. 

The eastern newt, one of the lethally 

vulnerable species (Martel et al. 2014; 

Gray et al. 2023), is one of the most 

widespread salamander species in North 

America (Roe and Grayson 2008, Martel 

et al. 2014). As top predators in pond 

ecosystems, eastern newts regulate frog 

tadpole abundance and, therefore, affect 

the amount and type of nutrients 

available in the ponds, keeping them in 

ecological balance (Morin et al. 1983; 

Morin 1995). If eastern newt 

populations decline because of Bsal 

infection in the wild, imbalances could 

result in ponds and ecosystems 

throughout the Eastern United States. 

Eastern newts also travel long distances 

between aquatic and terrestrial habitats 

(Roe and Grayson 2008), so if the 

species was to be eliminated from an 

area, the amount of nutrients available 

in upland areas would also be affected. 

The rough-skinned newt is another 

native U.S. species known to be lethally 

vulnerable to Bsal (Martel et al. 2014; 

Gray et al. 2023) and is geographically 

widespread along the Pacific coast of 

North America from Santa Cruz, 

California, to southeastern Alaska 

(AmphibiaWeb 2023a). The roughskinned newt plays an important role in 

ecosystems through its consumption of 

invertebrates that break down leaf litter 

and release carbon into the atmosphere 

(Davic and Welsh 2004). If roughskinned newt populations do 

experience severe declines from Bsal 

infection, atmospheric inputs of carbon 

may be altered, as has been observed 

with other species (Wyman 1998; Best 

and Welsh 2014). 

The green salamander (Aneides 

aeneus) was found by challenge tests to 

be lethally vulnerable when all 10 

salamanders in the study became 

infected and 5 died as a result (Gray et 

al. 2023). This species is State-listed as 

endangered in Indiana, Ohio, Maryland, 

and Mississippi and threatened in 

Pennsylvania (AmphibiaWeb 2023a); it 

is found in a narrow range from 

southwestern Pennsylvania southwest 

to Alabama. Of 15 Blue Ridge two-lined 

salamanders (Eurycea wilderae) that 

were challenge-tested, all became 

infected and 10 of those died (Gray et 

al. 2023; see also mortality in Carter et 

al. 2020; DiRenzo et al. 2021), and 6 of 

9 infected northern two-lined 

salamanders (of 9 Eurycea bislineata 

challenged; Gray et al. 2023). Red 

salamanders (Pseudotriton ruber), 

widely found from New York State to 

Alabama and Florida, had 100 percent 

mortality (Carter et al. 2019; Gray et al. 

2023). The Ensatina salamander 

(Ensatina eschscholtzii) is the only 

species in its genus, although it has 

seven subspecies, all found on the West 

Coast; two subspecies were documented 

as having lethal results (Gray et al. 

2023). 

Other taxa besides salamanders may 

also be negatively affected by Bsal. 

Several species of anurans have been 

found to carry Bsal (midwife toad Alytes 

obstetricans, Stegen et al. 2017; firebellied toad Bombina microdeladigitora, 

Nguyen et al. 2017; Cuban treefrog 

Osteopilus septentrionalis, Towe et al. 

2021; Gray et al. 2023). However, little 

is known about the negative effects of 

the disease on the anurans. None of 

these species is native to the United 

States, but all are imported in trade. 

Therefore, there is a risk of spreading 

Bsal to native frogs and toads, the 

susceptibility of which we do not know, 

and also a risk of spreading to 

salamanders. As explained above, we 

are not adding any frogs or toads to the 

list of injurious wildlife because they 

are in a different order (Anura), and we 

did not include the possibility of adding 

the order Anura in the 2016 interim 

rule, which would give the public a 

chance to comment. 

As Richgels et al. (2016) noted, some 

parts of the United States may reach 

temperatures above the thermal range of 

Bsal on a seasonal basis. However, 

wildlife and habitats would suffer losses 

if local populations of salamanders 

affected by Bsal prior to temperatures 

rising as part of the regular seasonal 

cycle suffered declines (and possible 

extirpation) and were unable to return 

to pre-infection levels in those 

ecosystems. 

Gray et al. (2023) estimated mean 

infectious and lethal doses for the North 

American species they tested with 

sufficient infection and mortality data 

and derived an amplification potential. 

Species with high amplification 

potential may contribute 

disproportionately to transmission 

events because they are easy to infect, 

less likely to die quickly from infection, 

and likely to be more infectious due to 

greater pathogen loads on their skin. 

Species that were susceptible to 

infection but did not die from Bsal are 

likely to be carriers. Considering all the 

variables, there is an immense potential 

for amphibian communities in North 

America to harbor carrier species that 

serve as reservoirs, amplification 

species that disproportionally transmit 

Bsal, and Bsal-susceptible species that 

are at high risk of population decline 

and extirpation. 


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For the above reasons, we conclude 

that the negative impact to wildlife 

resources or ecosystems is expected to 

be high if Bsal is introduced into U.S. 

ecosystems. 

Impacts to Threatened and Endangered 

Species and Their Habitats 

As of publication of the 2016 interim 

rule, none of the salamander species 

listed as endangered or threatened 

under the ESA in the United States had 

been specifically tested for Bsal 

vulnerability under laboratory 

conditions; Bsal had not been detected 

in their wild populations (Martel et al. 

2014, Bales et al. 2015). As of the final 

rule to the 2016 interim rule in this 

document, only the eastern hellbender 

(of which two subspecies are federally 

endangered) has been tested (Gray et al. 

2023) and is considered a carrier. Of the 

genera that include native species that 

we have identified as carriers, 20 

salamander species are federally listed 

as threatened or endangered and 2 

salamanders are candidates or proposed 

for listing. Because not all species have 

been tested, it is possible that the fungus 

will negatively affect other ESAprotected species. 

Impacts to Human Beings, Forestry, 

Horticulture, and Agriculture 

We do not expect direct effects to 

forestry, horticulture, or agriculture. 

Trees and other plants are also not 

affected. Bsal does not appear to infect 

humans or other animals except for 

salamanders and a few anurans. 

Indirectly, the introduction or 

establishment of Bsal would have 

negative effects on humans primarily 

from the loss of native wildlife 

biodiversity. These losses would affect 

the aesthetic, recreational, and 

economic values currently provided by 

native wildlife and healthy ecosystems. 

However, other indirect links to human 

health may occur. Many salamander 

species prey on mosquito larvae, and if 

the salamander numbers decline (such 

as from Bsal), the population of 

mosquitoes is likely to increase. Insect 

repellants used in surface waters for 

mosquito control have been linked to 

salamander larvae mortality and 

deformities, thus reducing predation on 

mosquito larvae, also leading to 

increased numbers of mosquitoes 

(Almeida et al. 2018). Similarly, a 

correlation has been made for Bd 

causing declines of mosquito-eating 

frogs, which then led to increased 

numbers of malaria-carrying mosquitoes 

(Springborn et al. 2022). Educational 

values would also be diminished 

through the loss of biodiversity and 

ecosystem health. However, we are not 

listing the species because of the 

indirect impacts to humans, forestry, 

horticulture, or agriculture, but rather 

due to their impacts to wildlife and 

wildlife resources. 

Wildlife or Habitat Damages That May 

Occur From Control Measures 

Richgels et al. (2016) stated, ‘‘[T]here 

are few known viable treatment or 

management options for responding to 

the introduction of Bsal * * * Strategies 

focused on prevention or reduction of 

introduction events remain the best 

control option for emerging diseases.’’ 

As discussed below in Ability To 

Prevent or Control the Spread of 

Pathogens or Parasites, current control 

strategies appear to focus on treating 

salamanders in a controlled laboratory 

setting. We are not aware of control 

measures that are effective in treating 

infected free-ranging salamanders over a 

large-scale area that could eliminate 

Bsal without killing the salamanders 

themselves, have low side effects, or do 

not require significant resources to 

implement. In addition, the life history 

of salamanders makes it highly unlikely 

that all individuals, including those that 

are infected, could be captured and 

treated. Many species are long-lived and 

inhabit areas that may be hard to reach. 

Furthermore, the effects on other 

wildlife of chemically treating an area, 

if such a treatment becomes available to 

eradicate infected salamanders and if 

capturing and treating individually is 

not practical, is unknown but is likely 

to be severe. 

H. Measures That Reduce or Remove 

Injuriousness of Salamanders 

Ability To Prevent Escape and 

Establishment 

As discussed below in Ability To 

Prevent or Control the Spread of 

Pathogens or Parasites, the ability and 

effectiveness of measures to prevent or 

control Bsal is currently low. While less 

certain, we also expect the ability to 

prevent escape and establishment is also 

low. Nonregulatory actions, such as 

implementing voluntary Best 

Management Practices or individual 

State action, are possible. The Service, 

for example, is working with partners 

on such efforts as HabitattitudeTM, a 

national campaign that encourages 

responsible consumer actions with 

respect to pet ownership. Such actions 

include finding alternatives to releasing 

pets into the environment. In November 

2015, PIJAC (currently known as the Pet 

Advocacy Network) asked its member 

entities to voluntarily ban their 

importation of paddle-tailed newts and 

Oriental fire-bellied newts to prevent 

the unintentional introduction of Bsal 

(PIJAC 2015). Voluntary actions, such as 

applying heat and antifungal medication 

therapy as described in Blooi et al. 

(2015a) and Blooi et al. (2015b), may 

help reduce the threat posed by Bsal for 

specimens held in captivity. However, 

at this time it is not possible to 

determine the likelihood of success of 

such measures in preventing the 

introduction, establishment, and spread 

of Bsal in the United States. 

As described above under 

Invasiveness of Salamanders and 

General Information About Bsal, 

nonnative salamanders have escaped 

into the United States, and Bd, a related 

fungus, has also escaped and 

established in the United States. While 

treatment options exist that may help 

reduce the threat posed by Bsal for 

imported and captive-held specimens, 

those options have not been 

standardized and their effectiveness 

remains uncertain for large-scale 

regulatory purposes. Treatment options 

for free-ranging specimens are not 

practical at this time. Therefore, we 

expect the likelihood of the ability to 

prevent escape and establishment of 

Bsal through infected salamanders to be 

low. Although voluntary actions are 

vital to help minimize the threat of 

invasive species, the Service is highly 

concerned about the extensive damage 

that introduction of Bsal would do to 

our Nation’s natural resources. Thus, we 

concluded that we cannot rely on 

voluntary actions alone to address the 

severity of the threat that Bsal poses and 

that other measures to prevent escape 

and establishment are not sufficient to 

ensure Bsal is not successfully 

introduced. 

Therefore, we find that we cannot rely 

on these approaches to prevent escape 

and establishment of Bsal and that our 

current capacity to prevent escape and 

establishment is low. 

Potential To Eradicate or Manage 

Established Populations 

While some introduced salamanders 

in the United States have been 

successfully controlled, such as the 

lesser siren (which was eliminated from 

a backyard pond outside its native U.S. 

range), others, such as the three-toed 

amphiuma, have not been as 

successfully controlled. However, 

evidence for control is sparse. Given the 

high rates of infection among 

salamanders tested by Martel et al. 

(2014), and the lack of control measures 

for Bsal that could be employed outside 

of a controlled facility, it is likely that 

Bsal would persist once introduced into 

the environment given appropriate 

environmental conditions, especially if 


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a tolerant or susceptible salamander 

established a population and continued 

to spread Bsal. 

Ability To Rehabilitate Disturbed 

Ecosystems 

Bsal infection can lead to the loss of 

keystone species in the ecosystem. The 

ability to rehabilitate disturbed 

ecosystems is expected to be low. We 

considered whether the Service’s 

National Fish Hatchery System (NFHS) 

could be used to maintain salamanders 

in refugia while areas are treated, 

assuming the salamanders could be 

treated for the fungus. However, it is 

impractical to equip NFHS facilities to 

be able to rapidly protect numerous 

salamander populations and maintain 

them for an extended time, such as 

might be required due to the 

introduction of Bsal. Although, as 

described in the next section, a few 

options exist to treat individual 

salamanders, none have been identified 

that can be used to clear Bsal from a 

widespread area. Consequently, we 

expect that, once Bsal has been 

introduced, it will persist and spread 

with little opportunity for widespread 

disinfection of ecosystems. 

Studies have also questioned the 

effectiveness of captive-breeding 

programs to address such threats as 

infectious disease to amphibians, 

including salamanders (Harding et al. 

2015). However, a recent study showed 

enhanced resistance following a second 

exposure of Bsal in eastern newts, both 

for increased survival and decreased 

Bsal loads by QPCR (L. Rollins-Smith, 

Vanderbilt University, pers. comm. 

2021). Also, another study found a 

second higher Bsal dose led to decreases 

in Bsal infection intensity over time as 

compared to salamanders exposed only 

once to a lower dose that sustained 

infections over time (DiRenzo et al. 

2021). Since that study was not 

designed specifically to study the effects 

of immune priming, these results are 

indicative but not proven. Therefore, it 

may be possible to stimulate an immune 

response in captive salamander 

populations that would allow them to 

be reintroduced into ecosystems where 

Bsal may still exist; however, this 

response has not been demonstrated for 

Bd, and research is needed in a broader 

array of conditions and species and to 

determine how resistance or immunity 

works. 

Therefore, the ability to rehabilitate 

disturbed ecosystems is expected to be 

low because the Service would be 

unable to ensure that all salamander 

populations expected to be affected by 

Bsal could be treated and protected in 

the wild. 

Ability To Prevent or Control the Spread 

of Pathogens or Parasites 

The ability and effectiveness of 

nonregulatory measures to prevent or 

control Bsal on a widespread scale from 

live specimens is currently low. The 

risk is compounded beyond the effect of 

a more common type of injurious listing 

(where the species cause harm by being 

invasive) by having two separate 

variables that can each spread—the 

fungus and the host species 

(salamanders). Few options can ensure 

potentially infected salamanders do not 

carry Bsal, and none exist on a broad 

scale. 

Blooi et al. (2015a) has shown that 

treating salamanders infected with Bsal 

by exposing them ‘‘to 25 °C [77 °F] for 10 

days resulted in complete clearance of 

infection and clinically cured all 

experimentally infected animals. This 

treatment protocol was validated in 

naturally infected wild fire 

salamanders.’’ The authors found that 

temperature treatment could be an 

effective option given the host 

salamander’s thermal tolerance. 

However, the treatment does have some 

shortcomings. Not all salamander 

species can tolerate the thermal regime 

required, and the researchers noted that 

there is a ‘‘narrow margin between the 

temperature able to limit [Bsal] and the 

upper thermal limit most urodelans 

tolerate.’’ Blooi et al. (2015a) also noted 

that there is some uncertainty as to 

whether the method is completely 

effective. Evidence of Bsal was found 

after thermal treatment, although it is 

possible that the evidence consisted of 

dead cells only. While thermal 

treatment is promising, the paper’s 

introduction noted that it was intended 

to ‘‘help to develop treatment protocols’’ 

and, therefore, is not intended to serve 

as the standalone standard treatment. As 

the treatment has not been standardized 

as a protocol for use at the landscape 

scale for salamanders in the wild or 

throughout trade, its ability to prevent 

introduction or control the spread of 

Bsal is low or uncertain. 

In the 2016 interim rule and 

corresponding economic analysis, one 

of the five alternatives that we 

considered was requiring a health 

certificate upon import stating that the 

animal being moved is free of Bsal, in 

lieu of or in addition to listing. During 

our evaluation for the 2016 interim rule, 

we considered whether it was practical 

for an exporting foreign nation to 

provide a health certificate stating that 

a possible carrier of Bsal has been tested 

and found to be free of the fungus or 

treated with antifungal drugs and 

thermal procedures to ensure that any 

Bsal that the salamanders might be 

carrying has been killed. We 

acknowledged that these testing and 

treatment methods existed and may be 

effective under certain circumstances. 

Requiring a health certificate would 

help ensure that Bsal does not escape 

from an exporting nation by being 

carried on an infected salamander. 

However, considering information from 

the public comments and other more 

recent information, we have significant 

concerns about this requirement’s 

feasibility for large-scale regulatory use 

for exporting countries given the 

effectiveness and sensitivity of current 

testing methods (including the return of 

false negatives), lack of validation and 

sufficient testing capacity, lack of 

standardized treatment methods, and 

agency resources required to conduct 

inspections, interpret results, and issue 

health certificates. The cost of testing 

could also be prohibitive for some 

exporters, since the cost of testing may 

be per animal. In the United States, 

qPCR testing can run around $25 to $65 

per salamander, and the salamander 

may wholesale for only $5. 

In May 2017, the WOAH listed 

infection with Bsal as an emerging 

disease in its Aquatic Animal Health 

Code; now WOAH considers Bsal as a 

disease of amphibians, with a chapter in 

the Aquatic Animal Health Code 

(WOAH 2021a) and a chapter in the 

manual (WOAH 2021b). The WOAH is 

the intergovernmental organization 

responsible for improving animal health 

worldwide. The WOAH chapter on Bsal 

provides recommendations that ‘‘may 

include’’ 13 named species of Asian, 

European, and North American newts 

and salamanders (WOAH 2021b); those 

species were a portion of the species 

covered by our 2016 interim rule. The 

recommendations include that the 

consignment be accompanied by an 

international aquatic animal health 

certificate issued by the Competent 

Authority of the exporting country. 

That recommendation came after the 

2016 interim rule took effect. The 2016 

draft economic analysis did not explain 

the costs of health certification because 

it was unclear how much testing, 

treatment, and the health-certification 

processes would cost. Because the 

details for these recommendations were 

not available for regulatory 

consideration for the 2016 interim rule’s 

public and peer comments and because 

much of those details are still not clear, 

we are not adding a health certification 

to this second interim rule. However, 

we believe there are valuable 

recommendations in the WOAH chapter 

8.2 (WOAH 2021a) that we support for 

the public to voluntarily help prevent or 


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control the spread of Bsal. They include, 

but are not limited to (summarized from 

WOAH 2021a; details found there): 

• quarantine; 

• treating or disposing of shipment 

water, equipment, containers, and 

packaging in a biosecure manner; and 

• treating effluent and waste 

materials (fomites) to inactivate Bsal. 

The European Union, Switzerland, 

and United Kingdom have each 

implemented Bsal-specific healthcertification requirements, and their 

regulations are similar to each other’s. 

Imports to these countries are 

prohibited unless they are from WOAH 

member countries and accompanied by 

a health certificate. The European 

Union’s implementing decision lays out 

testing and quarantine protocols along 

with providing a model health 

certificate. The United Kingdom costs 

for Bsal quarantining starts at £250 

($340) per consignment monitored and 

£40 ($54) for testing where required, 

and these could be untenable for most 

importers. Although some countries 

may have the necessary expertise to 

certify that salamanders are free of Bsal, 

not all exporting nations may have the 

necessary skills or resources, nor do we 

know which ones do. At some point in 

the future we may be able to propose 

health-certification criteria that are 

reliable and tenable, with the costs 

borne by the trade. 

Scientists and diagnostic laboratories 

are also working to standardize 

laboratory protocols, but there are 

currently no standardized sampling, 

testing, and screening protocols in the 

United States, and we do not know the 

standards, if any, in the various 

countries from which salamanders are 

currently being or may be exported. 

Assay sensitivity can vary between 

laboratories. A wide variety of 

laboratory equipment, reagents, 

techniques, protocols, and personnel 

experience is available, thus 

contributing to non-standardized 

techniques that can lead to variable or 

inconsistent results. Each laboratory has 

different equipment and uses different 

reagents (L. Sprague, USFWS, pers. 

comm., 2021). The North American Bsal 

Task Force recommends corroborative 

assays or further testing to validate a 

positive PCR or qPCR result, although 

WOAH does not state this; they also 

recommend quarantining under various 

scenarios (North American Bsal Task 

Force 2022). Considering the lack of 

amphibian quarantine facilities at ports 

of import and the unknown standards of 

testing for certification, we conclude it 

is currently not sufficient to rely on 

methods similar to those of the 

European Union and United Kingdom 

for preventing the introduction and 

establishment of Bsal. We need more 

information on this, which we requested 

below in K. Information Requested in 

question (10). 

Some treatment options also exist, 

such as treatment with antifungal 

medications that can be applied on 

animals that do not tolerate 25 °C (77 °F) 

(A. Martel, University of Ghent, pers. 

comm. 2015; Blooi et al. 2015b). It may 

be possible to treat amphibians in the 

wild for Bd with antifungals by 

capturing individuals and soaking them 

in a bath of the chemical, then releasing 

them back into the environment. As 

Hardy et al. (2015) showed for Cascades 

frogs (Rana cascadae), this process does 

not seem to be as effective as desired 

given possible side effects, but it may 

delay the eventual outcome of an 

outbreak enough to help individuals 

persist in the population. However, this 

process left unanswered questions about 

its applicability for salamanders and 

whether reinfection from fomites could 

still occur. Blooi et al. (2015b) identified 

a method for treating infected fire 

salamanders for Bsal with a 

combination of antifungals and 

temperature control that successfully 

cleared the fungus. However, such 

treatment worked only for controlled 

settings, such as those found in a 

laboratory or conservation facility, and 

side effects are unknown. 

It is impractical to treat widespread 

areas in the natural environment given 

the likely cost, personnel, and time 

needed to locate and treat all 

salamanders in the wild. Additionally, 

without a standardized process it is 

impractical to set required protocols 

that would apply to all specimens in 

trade. The possibility also exists for 

unknown and unintended consequences 

from such large-scale treatments, such 

as possible side effects from the 

widespread use of antifungal drugs. 

While promising, the treatment has not 

been standardized for use in a 

widespread manner for any salamander 

species. As we have noted above under 

Environmental Conditions Needed To 

Survive, Bsal is likely capable of 

persisting in the environment without a 

host by transmission to infected 

materials. Even if all individuals of a 

population could be successfully 

treated, the threat of reintroduction from 

environmental contamination would 

still exist. 

Even without the capacity to treat 

animals, research has shown that it is 

possible to identify whether a 

salamander is infected with Bsal, which 

would allow animals to be screened 

prior to importation. Blooi et al. (2013) 

presented a method of sampling 

salamanders and testing them to 

determine whether they carry Bsal. 

However, if certification occurs prior to 

importation, the process would require 

trained technicians in the country of 

origin, specialized equipment, and 

certainty that the salamanders would 

not become infected following 

certification. The results of those tests 

must then be interpreted by qualified 

health professionals and documented 

through a health-certification process. 

We cannot rule out the possibility of 

false negative tests or falsified 

documentation that could allow Bsal to 

be introduced into the United States. 

Post-import health certification by the 

Service at ports of entry would require 

holding salamanders at least 1 day for 

processing, testing, and diagnosis. 

Quarantine facilities would be needed 

while samples are processed and the 

health of the salamanders is certified. In 

addition, wildlife inspectors at ports of 

entry would need to be trained in 

testing and diagnosis procedures. 

In comparison, the certification 

process for listed salmonid species is 

standardized and salmonids are easier 

to test. For example, the tests for 

regulated salmonids are non-molecular, 

validated, require pathogen culture, and 

are, therefore, more straightforward. 

There are also only 12 countries 

(Australia, Canada, Chile, Denmark, 

Finland, Iceland, Isle of Man, New 

Zealand, Northern Ireland, Norway, 

Scotland, and Wales) that have officials 

certified to sign health certificates to 

export salmonids (generally as eggs) to 

the United States. Generally, only a few 

countries export to the United States in 

a given year. For example, in 2019, the 

Service received requests to import 

salmonid eggs from Canada, Denmark, 

Finland, and Iceland. The Service 

concluded that there is currently 

insufficient certainty that a certification 

program utilizing the method described 

by Blooi et al. (2013) would be effective 

in preventing the introduction or spread 

of Bsal in the United States. However, 

we may consider a certification system 

in the future and have posed a question 

below for public comment. 

Given the expected severity of 

consequences of Bsal introduction, 

imported salamanders that could be 

carriers may need to be treated, even 

with a health-certification process, 

which is not practical at this time for 

implementation by the Service as a 

broad-scale regulatory tool. The studies 

that have been conducted have not been 

standardized or agreed to as a suitable 

diagnostic or treatment effort on a large 

scale for treating all of the specimens 

that would potentially be imported. Not 

all species will tolerate treatment, and 


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reliable diagnostic capacity is needed to 

verify that animals do not carry Bsal 

following treatment. If an outbreak 

occurs, it would not be practical to 

locate and treat all free-ranging 

individuals in the wild in U.S. 

ecosystems. While antifungal agents 

could be applied to all animals, either 

in the laboratory or perhaps applied 

over a large geographic area, we are 

concerned about side effects on the 

animals being treated or nontarget 

species. We are also concerned about 

possible negative environmental effects 

if a chemical was widely applied 

(Gyllenhammar et al. 2009; Hasselberg 

et al. 2008). 

Researchers are also looking into the 

composition of the skin microbiome of 

various salamander species to gauge if it 

is possible to determine natural 

resistance to Bsal. For example, Bletz et 

al. (2018) found that European fire 

salamanders maintain complex skin 

microbiotas that have some Bsalinhibiting properties, but the bacterial 

numbers are too low to protect 

sufficiently against Bsal. Currently, we 

do not know if skin microbiota can be 

enhanced to inhibit Bsal sufficiently 

because of the complexities involved. 

In contrast to live specimens, 

salamanders that are chemically 

preserved with common scientific and 

museum collection protocols present no 

risk of introducing or transmitting Bsal. 

Tissue samples fixed in 10 percent 

formalin or embedded in paraffin 

(usually both in sequence) after routine 

histological processing (or both), 

including those from amphibians 

known to, or suspected of, carrying Bsal, 

contain only nonviable material and, 

thus, are not considered injurious (M. 

Forza ́n, Bsal Task Force, pers. comm. 

2021). In addition, experimental trials 

have demonstrated that Bsal is killed 

when exposed to 70 percent ethanol for 

at least 60 seconds (Van Rooij et al. 

2017). Therefore, skin swabs preserved 

in 70 percent ethanol are not considered 

injurious and may be used for PCR 

testing since this chemical does not 

damage DNA quality. However, freezing 

a salamander as a way to kill Bsal spores 

is not a proven method to kill Bsal to 

our knowledge. For Bd, analysis of some 

sample results suggest that the freezeshock treatment reduced the probability 

that Bd will grow, but all the Bd strains 

had at least a portion of samples that 

grew and produced zoospores following 

a freeze shock of ¥12 °C (10.4 °F) for 24 

hours (Voyles et al. 2017). Since we 

know of no evidence that unpreserved 

dead salamanders, including species 

that may be frozen, are not capable of 

carrying Bsal, they are considered 

injurious. 

In conclusion, various methods for 

mitigating the spread of Bsal by 

salamanders have been studied. We 

reviewed studies on thermal treatments, 

health certifications, anti-fungal 

medications, and resistance by skin 

microbiota, and none individually or in 

combination are sufficiently effective, 

safe, and broadly applicable to negate 

the need to list salamanders as 

injurious. 

Any Potential Ecological Benefits to 

Introduction 

No known benefits would result from 

Bsal or salamanders carrying Bsal 

occurring in the United States. The risks 

to native wildlife and wildlife resources 

greatly outweigh any unlikely benefits. 

Moreover, we are aware of no other 

potential ecological benefits for the 

introduction of Bsal or of Bsal-infected 

or Bsal-carrier salamanders into the 

United States. 

I. Summary and Conclusion for Interim 

Rule 

Overall, there is a high risk to the 

wildlife and wildlife resources of the 

United States from salamanders that are 

capable of carrying Bsal. The United 

States harbors 221 species of 

salamanders, more than any other 

country. Of the 23 native genera, 13 

genera were found to be vulnerable to or 

carriers of Bsal as of the drafting of this 

second interim rule. We find that the 

fungus is lethal to at least 9 native 

salamander species in 6 genera and that 

164 native species are considered 

carriers of Bsal. As of the drafting of this 

document, the vulnerability to disease 

and carrier status of 10 genera have not 

been tested or do not have conclusive 

results as carriers, many of which may 

have species vulnerable to this 

potentially deadly fungus. Under wild 

conditions, the disease may stress 

species to a point below the lethal 

threshold, and if these species are 

stressed by other factors, Bsal could 

cause cumulative harm to additional 

species. The benefits that these native 

salamander species provide to 

ecosystems, and in turn the ecosystem 

services that benefit people, are 

significant. The Service concludes that 

preventing Bsal from infecting native 

salamanders will prevent harmful 

effects to the wildlife and wildlife 

resources of the United States and 

merits listing of salamanders capable of 

carrying Bsal as injurious. 

Salamanders capable of carrying Bsal 

have the potential to escape and spread 

Bsal into the environment. Species 

capable of carrying Bsal can survive 

long enough in the wild to transmit the 

fungus or can transmit it to other 

carriers while in transit. Bsal can also be 

introduced and infect native 

salamanders by people improperly 

disposing of material that comes in 

contact with infected salamanders and 

can persist long enough in the 

environment without a host to represent 

a threat. 

Substantive evidence exists that all 

species within a genus where at least 

one species has been identified as a 

carrier of Bsal can also be a threat. Our 

review found no conclusive 

countervailing evidence. We find that, 

due to shared characteristics by species 

within a genus, other species within 

these genera are also highly likely to be 

carriers of Bsal, even if not every species 

in the genus has been tested to verify 

that it is a carrier of Bsal. For these same 

reasons, hybrids of species found in a 

genus that has at least one carrier 

species are also expected to be carriers. 

The main pathway for the global 

spread of Bsal is the international trade 

in salamanders. The most likely 

pathway of Bsal into the United States 

would be from salamanders being 

imported into the United States for 

commercial trade, including native 

species that are propagated outside the 

United States and subsequently 

imported into the United States. Listing 

salamanders that can carry Bsal as 

injurious wildlife will significantly 

confine this pathway and limit the 

capacity of Bsal to be introduced, 

become established, and spread in the 

United States. 

The current capacity to prevent 

escape and establishment is low. 

Rehabilitation of disturbed ecosystems 

is expected to be low, if not impossible. 

The ability and effectiveness of 

measures to prevent or control 

established Bsal is currently low. There 

are no known benefits of Bsal. 

The Service is listing live or dead 

specimens, hybrids, including parts, as 

injurious, but not eggs, gametes, 

preserved specimens or parts (including 

tissue), or purified extracted genetic 

material, where ‘‘preserved’’ means the 

preservation techniques kill the 

pathogen and thereby prevent 

transmission of Bsal. We find the risk of 

transmission of Bsal to other 

salamanders is high from both live and 

unpreserved dead specimens. Any 

salamanders that are infected and 

lethally vulnerable may die in transport 

and continue to carry Bsal into the 

United States. The risk is also high from 

improper disposal of materials that 

might be contaminated by those live or 

unpreserved dead specimens. Dead 

specimens, including those that are or 

were frozen, are considered injurious. 

Although under the authority of 18 


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U.S.C. 42 we cannot list contaminated 

materials (fomites) as injurious, by 

listing the carriers of Bsal, we seek to 

prevent the introduction of those 

materials. Conversely, material that is 

not injurious includes purified extracted 

genetic material because it carries a low 

risk of infection by Bsal; however, other 

tissue samples, such as skin swabs, are 

listed unless they have been chemically 

preserved to deactivate any live Bsal. 

Swabs that are preserved by exposing to 

70 percent ethanol for at least 60 

seconds are not considered injurious. 

The Service is not listing specimens that 

are chemically preserved. We conclude 

the risk of infection from such 

specimens is low. 

The Service is not adding eggs or 

gametes to the listing because there is 

no evidence that Bsal affects salamander 

reproductive tissue, such as eggs or 

gametes. The Service is not listing 

genera that we concluded are not 

carriers of Bsal because we do not have 

direct evidence that they are capable of 

introducing Bsal to the United States or 

otherwise transmitting it to native 

populations. We are also not listing 

genera where there is no data, even 

though it is possible that untested 

genera may also be capable of carrying 

Bsal. 

For the reasons stated, the Service 

finds the 16 genera comprising 

approximately 164 species of 

salamanders to be injurious to the 

wildlife and wildlife resources of the 

United States. The potential for Bsal 

introduction into the United States is 

high, the United States has suitable 

conditions for Bsal survival, and the 

consequences of introduction into the 

United States are expected to be 

significant and occur across a wide 

range of the United States. By listing 

species that can carry Bsal, we are 

taking preemptive action to help ensure 

the fungus does not enter the United 

States and infect native salamander 

populations and cause severe individual 

mortality, population declines, and 

ecosystem and economic harm. 

J. Required Determinations 

Regulatory Planning and Review 

Executive Order 12866 (E.O. 12866), 

as reaffirmed by E.O. 13563 and E.O. 

14094, provides that the Office of 

Information and Regulatory Affairs 

(OIRA) in the Office of Management and 

Budget (OMB) will review all significant 

rules. OIRA has determined that this 

rulemaking action is not significant. 

Executive Order 14094 reaffirms the 

principles of E.O. 12866 and E.O 13563 

and states that regulatory analysis 

should facilitate agency efforts to 

develop regulations that serve the 

public interest, advance statutory 

objectives, and are consistent with E.O. 

12866 and E.O. 13563. Regulatory 

analysis, as practicable and appropriate, 

shall recognize distributive impacts and 

equity, to the extent permitted by law. 

E.O. 13563 emphasizes further that 

regulations must be based on the best 

available science and that the 

rulemaking process must allow for 

public participation and an open 

exchange of ideas. We have developed 

this rule in a manner consistent with 

these requirements. 

The economic analysis for the second 

interim rule is provided in this section 

of the preamble. We are presenting the 

alternatives considered to identify 

whether there is a more effective option 

that can achieve the desired goals of the 

rule. The Service considered three 

alternatives for the economic analysis: 

Alternative 1: No action; Alternative 2: 

listing all species in 16 genera in which 

there is at least one confirmed carrier 

and all species in the genus are likely 

to be a carrier; and Alternative 3: listing 

all salamanders. We eliminated two 

alternatives that were considered for the 

2016 interim rule. One was ‘‘listing 

species that were identified by Martel et 

al. (2014) and other scientific sources to 

be carriers of Bsal.’’ This alternative was 

eliminated because new research 

provided evidence that this narrow 

approach could allow potential carriers 

to be imported. The other alternative 

was ‘‘requiring a health certificate 

stating that the animal being moved is 

free of Bsal, in lieu of or in addition to 

listing.’’ This alternative was eliminated 

because we do not have enough 

information to develop a reliable health 

certificate system at this time. 

To establish the baseline, analysis of 

current market conditions for imports 

and domestically bred salamanders is 

necessary. However, available U.S. 

salamander market data are minimal. 

The analysis uses two data sources to 

estimate the imported salamander 

industry: the Service’s LEMIS (USFWS 

OLE 2021) for data on the number of 

imported salamanders (live, dead 

specimens, hybrids, or parts) and the 

data submitted during the comment 

period for the 2016 interim rule by the 

Pet Industry Joint Advisory Council 

(PIJAC 2016) on live salamander pricing 

(updated to 2021 dollars). Due to 

limited data availability, we cannot 

estimate domestically bred salamander 

sales that are transported between the 

enumerated jurisdictions. We expect 

impacts to domestically bred 

salamanders to be minimal because, as 

determined by the 2017 court decision 

discussed above in this document, none 

of the alternatives prohibit interstate 

transport between States within the 

continental United States. We are 

requesting public comment on the 

number and sales of salamanders (by 

species) that are domestically bred and 

the percentage that are transported 

between the enumerated jurisdictions 

(see Information Requested below). 

We establish the baseline as the years 

2017 to 2019. This baseline accounts for 

the changes in imports and sales that 

resulted in January 2016 from the first 

interim rule (after the rule published, 

only 20 salamanders were imported in 

2016); the April 2017 court decision that 

overturned the prohibition on interstate 

transport between States within the 

continental United States; and the 

outlier 2020 data due to the pandemic. 

The second interim rule prohibits the 

importation of live or dead specimens, 

hybrids, including parts, as injurious, 

but not eggs, gametes, preserved 

specimens or parts (including tissue), 

unless an exemption is issued for 

scientific purposes. From 2017 to 2019, 

11 genera of live salamanders (of which 

5 genera are herein being listed; table 1), 

and 16 genera of salamander specimens 

(that is, dead salamanders or parts; of 

which 6 genera are herein being listed; 

table 2) were imported with no 

discernible trend. Live imports of the 

genera being listed herein are minimal, 

totaling 373 salamanders and 

approximately $18,000 (table 1). From 

2017 to 2019, approximately 1,000 

salamander specimens were imported 

(table 2), of which 25 percent would 

qualify as injurious under the second 

interim rule. The values for live 

salamanders with scientific purposes 

and for salamander specimens are 

unknown. No other salamanders (dead 

or parts) were imported during the 

baseline period, except for eggs, which 

would not be affected by this rule. 


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TABLE 1—TOTAL NUMBER OF ALL LIVE IMPORTED SALAMANDERS, 2017–2019 (2021$) 

[Data from USFWS OLE 2021] 

Genus * Purpose Live 

salamanders 

Estimated 

sales 

Ambystoma ..................................................................... Commercial .................................................................... 90 $8,000 

Scientific ......................................................................... 13 (**) 

Amphiuma ....................................................................... Commercial .................................................................... 1 <$1,000 

Andrias ............................................................................ Zoos ................................................................................ 4 $3,000 

Bolitoglossa ..................................................................... Commercial .................................................................... 5 <$1,000 

Cynops ............................................................................ Personal ......................................................................... 1 (**) 

Desmognathus ................................................................ Commercial .................................................................... 119 $3,000 

Eurycea ........................................................................... Commercial .................................................................... 137 $3,000 

Necturus .......................................................................... Commercial .................................................................... 163 $4,000 

Pleurodeles ..................................................................... Scientific ......................................................................... 108 (**) 

Pseudotriton .................................................................... Commercial .................................................................... 10 <$1,000 

Salamandra ..................................................................... Scientific ......................................................................... 100 (**) 

Total all genera ........................................................ ......................................................................................... 751 ** $24,000 

Total new genera ..................................................... ......................................................................................... 373 ** $18,000 

* Genera in bold are listed under the second interim rule. Amphiuma, Cynops, Pleurodeles, and Salamandra are listed under the final rule to 

the 2016 interim rule. 

** The value of live salamanders for scientific purposes is unavailable, and estimating this value is beyond the scope of this analysis. 

TABLE 2—TOTAL NUMBER OF ALL IMPORTED SALAMANDER SPECIMENS,1 2017–2019 

[Data from USFWS OLE 2021] 

Genus 2 Purpose Salamander 

specimens 1

Ambystoma ................................................................................................................... Scientific .................................................... 200 

Amphiuma * .................................................................................................................. Commercial ............................................... 200 

Bolitoglossa * ................................................................................................................ Scientific .................................................... 99 

Commercial ............................................... 80 

Cryptobranchus ............................................................................................................ Traveling Exhibit ....................................... 1 

Desmognathus ............................................................................................................. Scientific .................................................... 24 

Eurycea ........................................................................................................................ Scientific .................................................... 9 

Laotriton ........................................................................................................................ Scientific .................................................... 2 

Necturus * ..................................................................................................................... Commercial ............................................... 6 

Notophthalmus ............................................................................................................. Scientific .................................................... 9 

Nototriton * .................................................................................................................... Scientific .................................................... 6 

Oedipina * ..................................................................................................................... Scientific .................................................... 2 

Paramesotriton ............................................................................................................. Scientific .................................................... 4 

Plethodon ..................................................................................................................... Scientific .................................................... 6 

Salamandra .................................................................................................................. Scientific .................................................... 187 

Triturus ......................................................................................................................... Scientific .................................................... 11 

Tylototriton .................................................................................................................... Scientific .................................................... 111 

Total ...................................................................................................................... ................................................................... 957 

1Specimens are animals that are preserved for scientific or museum use; however, it is unknown whether these specimens would meet the 

preservation standards and be exempt from listing. 2Genera in bold are listed under the second interim rule. Genera with * are not listed under either rule. 

Alternative 1 is the no action 

alternative and is the status quo. We 

would not list additional species of 

salamanders as injurious. Retail sales of 

imported salamanders would continue; 

there would be no prohibition on 

transportation between the enumerated 

jurisdictions; and imports would 

continue. Salamander and ancillary 

industries would not incur any 

additional costs unless Bsal is 

introduced in the United States. 

Alternative 1 would not reduce the 

risk of introducing Bsal to the United 

States, and any benefits that accrue 

under Alternative 2 (this second interim 

rule) would not accrue under 

Alternative 1. Under Alternative 1, Bsal 

would continue to pose risk to native 

species and other wildlife resources in 

the United States. Furthermore, 

Alternative 1 does not meet the purpose 

of the listing, which is to prevent the 

introduction, establishment, and spread 

of Bsal in the wild in the United States. 

Therefore, we expect that greater 

financial and natural resources losses 

would be incurred due to managing and 

responding to Bsal if the fungus 

establishes and spreads in the United 

States compared to taking action now to 

prevent and minimize its introduction. 

Alternative 2 (second interim rule) 

lists all species in 16 genera for which 

there is at least one confirmed carrier 

and all species in that genus are likely 

to be a carrier. From 2017 through 2019, 

live individuals imported from genera 

that would be listed under Alternative 

2 were in Ambystoma, Desmognathus, 

Eurycea, and Pseudotriton, and 

specimens (dead individuals or parts) 

were in Ambystoma, Cryptobranchus, 

Desmognathus, Eurycea, and Laotriton. 

Under this alternative, live commercial 

imports totaled about 370 salamanders 

and $14,000, which represented 

approximately 47 percent of all live 


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salamander imports and 77 percent of 

sales. Live imports for scientific and 

zoological purposes totaled 17 

salamanders and represented 8 percent 

of scientific imports. All specimens 

under this alternative (235 specimens) 

were imported for scientific purposes, 

and importers would be eligible to 

apply for a permit. Under Alternative 2, 

imports of these genera would 

discontinue unless the importer is 

approved for a scientific permit. 

In the long term, the second interim 

rule is expected to benefit the economy. 

Efforts to control or eradicate invasive 

species (in this case, an invasive 

pathogen on a host wildlife species) and 

manage the costs they incur to society, 

once they have become established, are 

generally recognized as being less 

effective and more expensive than 

efforts to prevent potentially invasive 

species from establishing in the first 

place (Cuthbert et al. 2022). Emerging 

pathogens are currently 

underrepresented in databases of the 

cost of invasives, so adding them would 

greatly increase the estimated costs in 

the framework of biological invasions 

(Diagne et al. 2021). As a result, sectors 

of the economy that will not need to 

expend resources to control or manage 

injurious wildlife will be expected to 

gain from a timely listing process. 

Alternative 3 proposes listing all 804 

species of salamanders in the world. 

Although some species may or may not 

serve as carriers of Bsal, this alternative 

takes immediate action against those 

genera for which current scientific 

research and analysis has provided 

evidence are carriers of Bsal, along with 

other genera that may eventually be 

found to be carriers of Bsal. Under 

Alternative 3, all salamander imports 

would be prohibited (tables 1 and 2). 

This alternative would have the largest 

impact on salamander imports and the 

highest probability of preventing the 

introduction of Bsal in the wild. We did 

not select this option because we do not 

have enough evidence at this time that 

all genera could be carriers. However, 

evidence could be established in the 

future, or another reason could surface, 

such as the appearance of a 

hypervirulent variant of the fungus. 

We considered other alternatives that 

we rejected because we do not have 

sufficient information at this time that 

they could be effectively implemented 

to prevent introduction, establishment, 

and spread of Bsal from salamanders. 

For example, we do not have the 

capacity to establish and enforce a 

quarantine system or confidence in its 

effectiveness at preventing Bsal. We 

noted in the 2016 interim rule that, 

absent concerns regarding the 

effectiveness and sensitivity of current 

testing methods (including the return of 

false negatives), the lack of validation 

and sufficient testing capacity, and 

agency resources required to conduct 

inspections, interpret results, and issue 

health certificates, it may be possible to 

establish a health certification for 

salamanders that are free of Bsal. These 

concerns remain, and no such health 

certification has been established. 

However, this situation does preclude 

us from establishing health certification 

in the future if circumstances change. 

Appropriate conditions may also be 

included in injurious wildlife permits 

under the authority of and consistent 

with the purposes of 18 U.S.C. 42. 

We also considered encouraging 

partners to take nonregulatory action, 

such as voluntary best management 

practices or individual State action. The 

Service will pursue such actions as we 

move forward, and we are working with 

partners on such efforts as 

HabitattitudeTM, which encourages 

responsible consumer behaviors with 

respect to pet ownership. Voluntary 

actions, such as applying heat therapy 

as described in Blooi et al. (2015a) and 

Blooi et al. (2015b), may help reduce the 

threat posed by Bsal, but 

standardization and widespread 

application of the methods remain as 

challenges. Although voluntary actions 

are vital to help minimize the threat of 

invasive species, the Service is highly 

concerned about the extensive damage 

that introduction of Bsal would do to 

our Nation’s natural resources and 

concluded that we cannot rely on 

voluntary actions alone in this instance 

to address the severity of the threat that 

Bsal poses. 

Regulatory Flexibility Act 

Under the Regulatory Flexibility Act 

(as amended by the Small Business 

Regulatory Enforcement Fairness Act 

(SBREFA) of 1996), whenever a Federal 

agency is required to publish a notice of 

rulemaking for any proposed or final 

rule, it must prepare and make available 

for public comment a regulatory 

flexibility analysis that describes the 

effect of the rule on small entities (i.e., 

small businesses, small organizations, 

and small government jurisdictions) (5 

U.S.C. 601 et seq.). However, no 

regulatory flexibility analysis is required 

if the head of an agency certifies that the 

rule would not have a significant 

economic impact on a substantial 

number of small entities. Thus, for a 

regulatory flexibility analysis to be 

required, impacts must exceed a 

threshold for ‘‘significant impact’’ and a 

threshold for a ‘‘substantial number of 

small entities.’’ See 5 U.S.C. 605(b). 

SBREFA amended the Regulatory 

Flexibility Act to require Federal 

agencies to provide a statement of the 

factual basis for certifying that a rule 

would not have a significant economic 

impact on a substantial number of small 

entities. 

The U.S. Small Business 

Administration defines a small business 

as one with annual revenue or 

employment that meets or is below an 

established size standard for industries 

described in the North American 

Industry Classification System (NAICS). 

To assess the effects of the rule on small 

entities, we focus on (1) entities that 

import animals or animal parts and 

hybrids of listed genera and (2) entities 

with sales of animals, animal parts, and 

hybrids that are transported between the 

enumerated jurisdictions listed in 18 

U.S.C. 42(a)(1) and 50 CFR 16.3. Small 

entities affected by the rule are 

represented by categories and standards 

from the NAICS. The NAICS categories 

pertaining to this rule are those entities 

with: 

(1) receipts less than $32.0 million for 

‘‘Pet and Pet Supplies Stores’’ (NAICS 

459910); 

(2) receipts less than $2.75 million for 

‘‘All Other Animal Production’’ (NAICS 

112990); 

(3) receipts less than $34.0 million for 

‘‘Zoos and Botanical Gardens’’ (NAICS 

712130); 

(4) receipts less than $34.5 million for 

‘‘Colleges, Universities and Professional 

Schools’’ (NAICS 611310); and 

(5) fewer than 1,000 employees for 

‘‘Research and Development in the 

Physical, Engineering, and Life 

Sciences’’ (NAICS 541715). 

Under the second interim rule, we 

expect the effect on entities that import 

the 16 genera to be small. From 2017 to 

2019, seven businesses imported live 

salamanders from some of those genera, 

which represented 0.1 percent of all pet 

and pet-supplies establishments and 

less than 0.1 percent of all other animalproduction establishments. Three 

businesses imported the listed 

specimens for scientific purposes, 

which represented less than 0.1 percent 

of all universities and research facilities 

(USFWS OLE 2021). We expect the 

effect on entities that sell the 16 genera 

between the enumerated jurisdictions to 

be small as well, because the interim 

rule does not prohibit interstate 

transport between the 49 States in the 

continental United States. Furthermore, 

pet stores outside the 49 States in the 

continental United States represent less 

than 1 percent of all stores and less than 

1 percent of total pet store sales (USCB 

2017). 


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Federal Register / Vol. 90, No. 6 / Friday, January 10, 2025 / Rules and Regulations 2219 

Therefore, we certify that this interim 

rule will not have a significant 

economic effect on a substantial number 

of small entities as defined under the 

Regulatory Flexibility Act (5 U.S.C. 601 

et seq.). An initial regulatory flexibility 

analysis is not required. Accordingly, a 

small entity compliance guide is not 

required. 

The second interim rule makes no 

changes in the compliance requirements 

of any business. The Service is unaware 

of any duplicative, overlapping, or 

conflicting Federal rules. Several States 

implement similar acts that are more 

restrictive than the Federal law. 

Congressional Review Act 

The interim rule is not a major rule 

under 5 U.S.C. 804(2), the Congressional 

Review Act. This rule: 

a. Would not have an annual effect on 

the economy of $100 million or more. 

The rule listing 16 genera of 

salamanders, including approximately 

164 species, would prohibit an 

estimated 125 live salamanders 

imported per year and prohibit the 

transport of domestically bred 

individuals between the enumerated 

jurisdictions. In addition, businesses 

would also face the risk of fines if 

caught transporting these salamanders 

or their parts between the enumerated 

jurisdictions in the shipment clause of 

18 U.S.C. 42(a)(1), which is codified in 

Federal regulations at 50 CFR 16.3. The 

penalty for violation of this law is not 

more than 6 months in prison and not 

more than a $5,000 fine for an 

individual and not more than a $10,000 

fine for an organization. 

b. Would not cause a major increase 

in costs or prices for consumers, 

individual industries, Federal, State, or 

local government agencies, or 

geographic regions. 

c. Would not have significant adverse 

effects on competition, employment, 

investment, productivity, innovation, or 

the ability of United States-based 

enterprises to compete with foreignbased enterprises. 

Unfunded Mandates Reform Act (2 

U.S.C. 1501 et seq.) 

In accordance with the Unfunded 

Mandates Reform Act (2 U.S.C. 1501), 

the Service makes the following 

findings: 

a. This rule would not produce a 

Federal mandate. In general, a Federal 

mandate is a provision in legislation, 

statute, or regulation that would impose 

an enforceable duty upon State, local, or 

Tribal governments, or the private 

sector. 

b. The rule would not have a 

significant or unique effect on State, 

local, or Tribal governments or the 

private sector. A statement containing 

the information required by the 

Unfunded Mandates Reform Act (2 

U.S.C. 1531 et seq.) is not required. 

Takings 

In accordance with Executive Order 

12630 (Government Actions and 

Interference with Constitutionally 

Protected Private Property Rights), the 

rule does not have significant takings 

implications. A takings implication 

assessment is not required. This rule 

would not impose significant 

requirements or limitations on private 

property use. While import and 

transport between the enumerated 

jurisdictions of any of the listed species 

is prohibited, 18 U.S.C. 42(a) does not 

prohibit any person who owns one of 

the listed species at the time of listing 

from continuing to possess the 

salamander or engaging in intrastate 

transport and other activities within 

their State or territory, as allowed under 

State, Tribal, or territorial law. 

Federalism 

In accordance with Executive Order 

13132 (Federalism), this interim rule 

does not have significant federalism 

effects. A federalism assessment is not 

required. This rule would not have any 

direct effects on States, on the 

relationship between the Federal 

Government and the States, or on the 

distribution of power and 

responsibilities among the various 

levels of government. Therefore, in 

accordance with Executive Order 13132, 

we determine that this rule does not 

have sufficient federalism implications 

to warrant the preparation of a 

federalism assessment. 

Civil Justice Reform 

In accordance with Executive Order 

12988, the Office of the Solicitor has 

determined that the interim rule does 

not unduly burden the judicial system 

and meets the requirements of sections 

3(a) and 3(b)(2) of the Executive order. 

The interim rule has been reviewed to 

eliminate drafting errors and ambiguity, 

was written to minimize litigation, 

provides a clear legal standard for 

affected conduct rather than a general 

standard, and promotes simplification 

and burden reduction. 

Paperwork Reduction Act of 1995 (44 

U.S.C. 3501 et seq.) 

This rule does not contain any new 

collections of information that require 

approval by the Office of Management 

and Budget (OMB) under the Paperwork 

Reduction Act of 1995 (44 U.S.C. 3501 

et seq.). OMB has previously approved 

the information collection requirements 

associated with filing declarations and 

the importation of injurious wildlife and 

assigned the following OMB Control 

Numbers: 

• 1018–0012, ‘‘Declaration for 

Importation or Exportation of Fish or 

Wildlife, 50 CFR 14’’ (expires 03/31/ 

2024, and in accordance with 5 CFR 

1320.10, an agency may continue to 

conduct or sponsor this collection of 

information while the submission is 

pending at OMB), and 

• 1018–0078, ‘‘Injurious Wildlife; 

Importation Certification for Live Fish 

and Fish Eggs (50 CFR 16)’’ (expires 01/ 

31/2024, and in accordance with 5 CFR 

1320.10, an agency may continue to 

conduct or sponsor this collection of 

information while the submission is 

pending at OMB). 

An agency may not conduct or 

sponsor, and a person is not required to 

respond to, a collection of information 

unless it displays a currently valid OMB 

control number. 

National Environmental Policy Act 

We have reviewed this rule in 

accordance with the criteria of the 

National Environmental Policy Act 

(NEPA) and our Departmental Manual 

in 516 DM. This rule does not constitute 

a major Federal action significantly 

affecting the quality of the human 

environment. Under Department of the 

Interior agency policy and procedures, 

this rule is covered by a categorical 

exclusion (516 DM 8.5 C(9)), and 

preparation of a detailed statement 

under NEPA is not required because it 

adds species to the list of injurious 

wildlife under 50 CFR subchapter B, 

part 16, which prohibits the importation 

into the United States and shipment 

between some jurisdictions of wildlife 

found to be injurious (for further 

information on the categorical 

exclusion, see 80 FR 66554, October 29, 

2015). The categorical exclusion states, 

‘‘The adding of species to the list of 

injurious wildlife regulated under the 

Lacey Act (18 U.S.C. 42, as amended) as 

implemented under 50 CFR subchapter 

B, part 16, which prohibits the 

importation into the United States 

* * * of wildlife found to be injurious.’’ 

We have also determined that the rule 

does not involve any of the 

extraordinary circumstances listed in 43 

CFR 46.215 that would require further 

analysis under NEPA. 

Government-to-Government 

Relationship With Tribes 

In accordance with the President’s 

memorandum of April 29, 1994, 

‘‘Government-to-Government Relations 

with Native American Tribal 


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2220 Federal Register / Vol. 90, No. 6 / Friday, January 10, 2025 / Rules and Regulations 

Governments’’ (59 FR 22951), Executive 

Order 13175, and the Department of the 

Interior’s manual at 512 DM 2, we 

readily acknowledge our responsibility 

to communicate meaningfully with 

recognized Federal Tribes on a 

government-to-government basis. In 

accordance with Secretarial Order 3206 

of June 5, 1997 (American Indian Tribal 

Rights, Federal-Tribal Trust 

Responsibilities, and the Endangered 

Species Act), we readily acknowledge 

our responsibilities to work directly 

with Tribes in developing programs for 

healthy ecosystems, to acknowledge that 

Tribal lands are not subject to the same 

controls as Federal public lands, to 

remain sensitive to Indian culture, and 

to make information available to Tribes. 

For this interim rule, we sent a letter to 

the leaders of the almost 580 federally 

recognized Tribes providing some 

background and asking for their 

comments. We received none. We have 

evaluated potential effects on federally 

recognized Indian Tribes and have 

determined that there are no potential 

effects. This rule involves the 

importation of salamanders and 

shipment of salamanders between the 

enumerated jurisdictions of the 

shipment clause of 18 U.S.C. 42, also set 

forth in 50 CFR 16.3. We are unaware 

of such movement in these species by 

Tribes. 

Effects on Energy 

Executive Order 13211 requires 

agencies to prepare statements of energy 

effects when undertaking certain 

actions. This rule is not expected to 

affect energy supplies, distribution, and 

use. Therefore, this action is a not a 

significant energy action and no 

statement of energy effects is required. 

Clarity of Rule 

We are required by Executive Orders 

12866 and 12988 and by the 

Presidential Memorandum of June 1, 

1998, to write all rules in plain 

language. This means that each rule we 

publish must: 

a. Be logically organized; 

b. Use the active voice to address 

readers directly; 

c. Use clear language rather than 

jargon; 

d. Be divided into short sections and 

sentences; and 

e. Use lists and tables wherever 

possible. 

If you feel that we have not met these 

requirements, send us comments by one 

of the methods listed in ADDRESSES. To 

help us revise the rule, your comments 

should be as specific as possible. For 

example, you should tell us the 

numbers of the sections or paragraphs 

that are unclearly written, which 

sections or sentences are too long, and 

the sections where you feel lists or 

tables would be useful. 

K. Information Requested 

We are soliciting public comments 

and supporting data for this second 

interim rule to add 16 new genera to the 

current list of 20 genera of salamanders 

that are listed as injurious amphibians 

under 18 U.S.C. 42, including comments 

and supporting data on the economic 

information as described above in the 

Required Determinations. As stated 

above in this document, we are not 

soliciting comments regarding the 

listing of the genera that were listed in 

the 2016 interim rule. We will review 

the public comments for the preparation 

of a second final rule. 

You may submit your comments and 

materials concerning this second 

interim rule by one of the methods 

listed in ADDRESSES. We will not accept 

comments sent by email or fax or to an 

address not listed in ADDRESSES. We 

will post your entire comment— 

including your personal identifying 

information—on https://

www.regulations.gov. If your written 

comments provide personal identifying 

information, you may request at the top 

of your document that we withhold this 

information from public review. 

However, we cannot guarantee that we 

will be able to do so. 

Comments and materials we receive, 

as well as supporting documentation we 

used in preparing this second interim 

rule, will be available for public 

inspection on https://

www.regulations.gov under Docket No. 

FWS–HQ–FAC–2015–0005, or by 

appointment, during normal business 

hours at the Service’s office in Falls 

Church, VA (see FOR FURTHER

INFORMATION CONTACT). 

We are soliciting public comments 

and supporting data to gain additional 

information, and we specifically seek 

comment on the following questions: 

(1) How many of the new genera 

listed by this interim rule are currently 

in domestic production for wholesale or 

retail sale, and in how many and which 

States? 

(2) How many businesses sell 

salamanders from the genera listed by 

this interim rule, and how many 

businesses transport these listed genera 

between enumerated jurisdictions? 

(3) How many businesses breed 

salamanders of one or more of the 

genera listed by this interim rule? 

(4) What species listed as threatened 

or endangered by one or more States 

would be affected by the introduction of 

Bsal? 

(5) What provisions in this interim 

rule should the Service have considered 

with regard to: (a) the impact of the 

provision(s) (including any benefits and 

costs), if any, and (b) the alternatives, if 

any, that the Service should consider, as 

well as the costs and benefits of those 

alternatives, paying specific attention to 

the effect of the rule on small entities? 

(6) How could this interim rule be 

modified to reduce costs or burdens for 

some or all entities, including small 

entities, consistent with the Service’s 

requirements? For example, we seek 

comment on the distinct benefits and 

costs, both quantitative and qualitative, 

of (a) the prohibitions on importation 

and (b) the prohibitions on transport 

between enumerated jurisdictions of the 

genera listed by this rule. What are the 

costs and benefits of the modifications? 

(7) Is there any evidence suggesting 

that Bsal has been introduced into the 

United States or may have already 

established? 

(8) Is there evidence suggesting that 

any of the genera listed by this interim 

rule are not carriers of Bsal? If so, which 

ones? 

(9) Is there evidence suggesting that 

additional salamander genera are 

carriers of Bsal and should be listed as 

injurious? If so, which ones? 

(10) Could a reliable health certificate 

within the Service’s authority be 

developed that would allow Bsal-free 

salamander imports? Are there 

treatments that would ensure 

salamanders imported into the United 

States are reliably free of Bsal, and how 

could compliance be monitored? 

(11) Are there other means of 

preserving or treating salamander 

specimens, parts, or products that are 

not identified in this rule and that are 

proven adequate to render Bsal nonviable? 

(12) Should the Service add eggs or 

other reproductive material of listed 

salamanders to the list of injurious 

wildlife because they may also carry 

Bsal? 

(13) What are relevant Federal, State, 

or local rules that may duplicate, 

overlap, or conflict with this interim 

rule? 

We will also submit the rule for peer 

review concurrent with public 

comments. In conducting peer review, 

we will follow guidance from the Office 

of Management and Budget ‘‘Final 

Information Quality Bulletin for Peer 

Review’’ (OMB 2004) and the Service’s 

own guidance. 

References Cited 

A complete list of all references used 

in this rulemaking is available at https:// 


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Federal Register / Vol. 90, No. 6 / Friday, January 10, 2025 / Rules and Regulations 2221 

www.regulations.gov under Docket No. 

FWS–HQ–FAC–2015–0005. 

Authors 

The primary authors of this rule are 

the staff members of the U.S. Fish and 

Wildlife Service’s Branch of Aquatic 

Invasive Species. 

List of Subjects in 50 CFR Part 16 

Animal diseases, Imports, Reporting 

and recordkeeping requirements, 

Transportation, Wildlife. 

Regulation Promulgation 

For the reasons discussed in the 

preamble, the U.S. Fish and Wildlife 

Service amends part 16, subchapter B of 

chapter I, title 50 of the Code of Federal 

Regulations, as set forth below: 

PART 16—INJURIOUS WILDLIFE 

■ 1. The authority citation for part 16 

continues to read as follows: 

Authority: 18 U.S.C. 42. 

■ 2. Revise § 16.14(a) to read as follows: 

§ 16.14 Importation of live or dead 

amphibians or their eggs. 

(a) The importation, transportation, or 

acquisition of any live or dead specimen 

or hybrid, including parts (except for 

eggs or gametes; parts or tissues that 

have been chemically preserved, 

chemically treated, or heat treated so 

that the pathogen Batrachochytrium 

salamandrivorans, if present, is 

rendered non-viable; and molecular 

specimens consisting of only the nucleic 

acids from organisms), of all species in 

the genera Ambystoma, Andrias, 

Aneides, Aquiloeurycea, Calotriton, 

Chioglossa, Chiropterotriton, 

Cryptobranchus, Cynops, 

Desmognathus, Ensatina, Euproctus, 

Eurycea, Hydromantes, Hynobius, 

Ichthyosaura, Laotriton, Lissotriton, 

Neurergus, Notophthalmus, 

Ommatotriton, Onychodactylus, 

Pachytriton, Paramesotriton, Plethodon, 

Pleurodeles, Proteus, Pseudobranchus, 

Pseudotriton, Salamandra, 

Salamandrella, Salamandrina, Siren, 

Taricha, Triturus, and Tylototriton is 

prohibited except as provided under the 

terms and conditions set forth at § 16.22 

of this part. 

* * * * * 

Shannon Estenoz, 

Assistant Secretary for Fish and Wildlife and 

Parks. 

[FR Doc. 2024–31203 Filed 1–8–25; 8:45 am] 

BILLING CODE 4333–15–P 


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