Endangered and Threatened Wildlife and Plants; Final Rule To Remove the Plant ``Echinocereus lloydii'' (Lloyd's Hedgehog Cactus) From the Federal List of Endangered and Threatened Plants

DEPARTMENT OF THE INTERIOR

Fish and Wildlife Service

50 CFR Part 17

RIN 1018-AD91

Endangered and Threatened Wildlife and Plants; Final Rule To

Remove the Plant ``Echinocereus lloydii'' (Lloyd's Hedgehog Cactus)

From the Federal List of Endangered and Threatened Plants

AGENCY: Fish and Wildlife Service, Interior.

ACTION: Final rule.

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SUMMARY: We, the U.S. Fish and Wildlife Service, are removing the plant

Echinocereus lloydii (Lloyd's hedgehog cactus), from the Federal List

of Endangered and Threatened Species under the authority of the

Endangered Species Act of 1973, as amended (Act). Lloyd's hedgehog

cactus was listed as endangered on October 26, 1979, as a result of

threats presented by collection and highway projects. Recent evidence

indicates that Lloyd's hedgehog cactus is not a distinct species but

rather a hybrid or cross which is not evolving independently of its

parental species. Therefore, E. lloydii no longer qualifies for

protection under the Act. Removing Lloyd's hedgehog cactus from the

list constitutes our recognition of its hybrid status and removes

Federal protection under the Endangered Species Act.

DATES: This rule is effective July 26, 1999.

ADDRESSES: The complete file for this rule is available for inspection,

by appointment, during normal business hours, at the U.S. Fish and

Wildlife Service's Austin Texas Field Office, U.S. Fish and Wildlife

Service, 10711 Burnet Road, Suite 200, Austin, Texas 78758.

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FOR FURTHER INFORMATION CONTACT: Kathryn Kennedy, botanist, U.S. Fish

and Wildlife Service, 10711 Burnet Road, Suite 200, Austin, Texas

78758, (telephone 512/490-0057; facsimile 512/490-0974).

SUPPLEMENTARY INFORMATION:

Background

Echinocereus lloydii (Lloyd's hedgehog cactus), a member of the

cactus family, was first collected by F.E. Lloyd in 1909 and was named

in his honor by Britton and Rose (1922). The first plants collected by

Mr. Lloyd were from near Fort Stockton, Pecos County, Texas (Weniger

1970). Lloyd's hedgehog cactus is cylindrical with one or several

ribbed stems which grow up to about 20 centimeters (cm) (8 inches (in))

high and 10 cm (4 in) in diameter. The flowers vary a great deal in

color from lavender to magenta, are about 5 cm (2 in) in diameter, and

form mature fruits that are green tinged with pink or orange when ripe.

(Correll and Johnston 1979, Poole and Riskind 1987).

Lloyd's hedgehog cactus is known from Brewster, Culberson, Pecos,

and Presidio Counties, Texas, and Eddy County, New Mexico. It has also

been reported from the state of Chihuahua in Mexico. Currently fewer

than 15 populations are known, most occurring on private lands.

We listed Lloyd's hedgehog cactus as an endangered species on

October 26, 1979 (44 FR 61916), under the authority of the Endangered

Species Act of 1973, as amended (Act) (16 U.S.C. 1531 et. seq.) At the

time of listing, botanists considered Lloyd's hedgehog cactus a

distinct species threatened by over-collection, habitat loss or

alteration due to highway construction and maintenance, and potentially

by overgrazing.

The physical characteristics of specimens of Lloyd's hedgehog

cactus were long recognized as intermediate between those of

Echinocereus dasyacanthus (Texas rainbow cactus) and Echinocerus

coccineus (a species of claret-cup cactus). Several theories emerged as

to how this intermediacy may have arisen. One theory was that Lloyd's

hedgehog cactus represented a primitive ancestral evolutionary lineage

(ancestry), which diversified over time to give rise to two new

lineages producing E. dasyacanthus and E. coccineus. A second theory

was that Lloyd's hedgehog cactus was of more recent hybrid origin, the

result of ancient hybridization or crossing between E. dasyacanthus and

E. coccineus, but now an independent taxon or group of organisms

recognizable as a species.

While reports of interspecific hybridization (cross between two

species) between members of the genus Echinocereus were known,

hybridization between E. coccineus and E. dasyacanthus seemed highly

unlikely as the two species differ greatly in morphology (structure and

form), have different predominant pollinators (one hummingbird

pollinated, the other bee pollinated), and generally grow in different

habitats; the first being a more mesic species (average moisture) and

the latter being more typically found in more open desert. In addition,

in sites where the plants were grown or seen in proximity to each other

they were observed to bloom at different times with little if any

overlap. While many hybrids are sterile, plants of E. lloydii are

fertile and able to reproduce. In addition, because these wild

populations have persisted over time, treatment as a distinct species

was generally accepted.

Steve Brack (U.S. Fish and Wildlife Service 1985) reported locating

E. lloydii only in proximity to E. dasyacanthus and E. coccineus. This

apparent lack of isolation combined with the intermediate appearance of

the plants raised questions about the taxonomic interpretation of E.

lloydii as a distinct species. These taxonomic questions supported the

possibility that Lloyd's hedgehog cactus might be a result of recent

and sporadic hybridization events, with these wild populations simply

representing relatively unstable hybrid swarms that are not evolving

independently and are not recognizable as a species. In response to

this new information we determined that the question of the hybrid

status of Lloyd's hedgehog cactus should be further investigated.

In studies by Powell, Zimmerman, and Hilsenbeck (1991) and Powell

(1995) the progeny resulting from the artificial crossing of E.

dasyacanthus and E. coccineus and naturally occurring E. lloydii was

examined using artificial cross-pollination (cross fertilization),

morphological analyses (analysis of structure and form), pollen

stainability studies (using slide stain techniques to assess the

viability of pollen), chromosome counts, and phytochemical analysis

(plant chemical). Their research demonstrated that hybrids between E.

dasyacanthus and E. coccineus could be easily produced, closely

resembled the naturally occurring E. lloydii, and were interfertile and

able to backcross to the parental species. One theory resulting from

this work was that if fertile hybrids were produced in the wild, they

could presumably multiply and backcross to the parental species forming

the sort of persistent intermediate populations of high variability

which are found naturally in the wild. This suggests that Lloyd's

hedgehog cactus may have arisen as a result of hybridization between

these other two species of Echinocereus, both of which are common and

not protected by the Act.

The probability that Lloyd's hedgehog cactus arose through

hybridization (crossbreeding) rather than representing a persistent

ancestral condition was heightened by Powell et al.'s (1991) finding

that naturally occurring E. lloydii have tetraploid chromosome numbers

(four times the normal chromosome numbers), as do E. dasyacanthus and

E. coccineus. Tetraploid chromosome numbers are considered an advanced

or recently derived characteristic in the family Cactaceae, rather than

a primitive one. Zimmerman (1993) made additional observations on

pollinators and other ecological and phenological (the study of

periodicity in relation to climate and environment) isolating

mechanisms, examined the primitive and advanced species of the E.

dasyacanthus and E. coccineus taxonomic groups (rainbow cacti and

claret-cup cacti) and E. lloydii, and performed cladistic analyses

(analysis of the order of evolutionary decent). This work resulted in

his agreement that Lloyd's hedgehog cactus is not primitive and

probably arose as a result of hybridization.

The conclusion that plants recognized as E. lloydii arose through

hybridization raised questions about the integrity or cohesiveness of

populations and whether they were a sufficiently distinct, isolated and

independently evolving genome (genetic entity) that they should be

recognized as distinct species. Powell et al. (1991) and Powell (1995),

in their phytochemical, morphological, and crossing studies detected no

unique characters or reproductive isolation that would demonstrate any

independent evolution had occurred. Though their study lacked

comprehensive examination and interpretation of populations in the

field and throughout the known range, they suggested that populations

recognized as E. lloydii might represent mere hybrids, and should

probably at best be recognized only as an illegitimate species

recognized nomenclaturally (by scientific name) for purposes of

identification. They designated their artificially produced hybrids as

Echinocereus X lloydii.

Zimmerman (1993) examined geographical distribution, correlations

with geographic variation across the range of E. lloydii and its

parental

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species, and population characteristics at several sites in the wild.

He found that E. lloydii was only found in areas where both E.

dasyacanthus and E. coccineus occur. Further, sites with plants known

as E. lloydii were not uniform in appearance, and exhibited great

variation among individuals consistent with a pattern of backcrossing

or introgression with the parental species. Zimmerman could find no

evidence of reproductive isolation in the field. Zimmerman found that

blooming time overlapped both parental species, and hybrid individuals

did not exhibit any significant habitat preference that would provide

any significant separation from the parental species, concluding that

E. lloydii is not a legitimate species. Zimmerman's review of the

nomenclature resulted in the recommendation that plants formerly

recognized as E. lloydii should properly be referred to as Echinocereus

X roetteri var. neomexicanus.

Previous Federal Action

Federal action concerning Lloyd's hedgehog cactus began with

Section 12 of the original Endangered Species Act of 1973, which

directed the Secretary of the Smithsonian Institution to prepare a

report on those plants considered to be endangered, threatened, or

extinct. This report, designated as House Document No. 94-51 was

presented to Congress on January 9, 1975. A notice was published on

July 1, 1975 (40 FR 27823), of our acceptance of the report of the

Smithsonian Institution as a petition to list these species, including

Echinocereus lloydii, under Section 4(c)(2), now section 4(b)(3)(A) of

the Act.

The report was published in the Federal Register on July 1, 1975

(40 FR 27823-27924), and provided notice of our intention to review the

status of the plant taxa named within. On June 16, 1976, we published a

proposed rulemaking in the Federal Register (41 FR 24523-24572)

proposing the listing of approximately 1,700 vascular plant species as

endangered under Section 4 of the Act. Echinocereus lloydii was

included in this list. In response to our proposal of June 16, 1976,

four hearings were held in July and August of 1976, in the following

locations: Washington, D.C.; Honolulu, Hawaii; El Segundo, California;

and Kansas City, Missouri. We held a fifth public hearing on July 9,

1979, in Austin, Texas for seven Texas cacti, including E. lloydii, and

one fish.

We published a final rule in the Federal Register on June 24, 1977

(42 FR 32373-32381, codified at 50 CFR 17) detailing the regulations to

protect Endangered and Threatened plant species. These regulations

codified the prohibitions of the Act and established procedure for the

permitting of certain activities under the Act. We published a final

rule to list the Lloyd's hedgehog cactus as an endangered species on

October 26, 1979 (44 FR 61916).

We initiated our review of new information and the status of

Lloyd's hedgehog cactus in 1994 and a draft proposed delisting rule was

forwarded to the Washington Office on April 4, 1995. However, a listing

moratorium (Public Law 104-6, April 10, 1995) and rescission of listing

program funding in Fiscal Year 1996 disrupted our listing program. This

moratorium was lifted and our listing program funding was restored on

April 26, 1996. We issued guidance on May 16, 1996 (61 FR 24722),

setting priorities for restarting the listing program that included

processing of proposed delistings already in the Washington Office. The

proposed rule for delisting Lloyd's hedgehog cactus was published on

June 14, 1996 (61 FR 30209). The public comment period on the proposed

rule closed August 13, 1996.

Our listing priority guidance for Fiscal Year 1997, finalized

December 5, 1996 (61 FR 64475), precluded the final delisting decision

and processing of this final rule. Our 1997 guidance determined that,

given limited resources, enacting conservation protection for the

backlog of listing actions for high priority imperiled species merited

priority. Delistings and reclassifications actions were given our

lowest priority.

With the publication of listing priority guidance for Fiscal Years

1998 and 1999 on May 8, 1998 (63 FR 25502), we returned to a more

balanced listing program. Delisting and reclassification actions are

now in the lowest priority position within Tier 2 actions. With

resources allocated to all types of Tier 2 listing actions, work on the

final determination for Lloyd's hedgehog cactus resumed.

In our June 14, 1996 (61 FR 30209), proposed rule, all interested

parties were requested to submit factual reports or information that

might contribute to the development of a final rule. One hundred and

fifteen letters of notification were sent to appropriate Federal and

State agencies, county governments, scientific organizations, and other

interested parties requesting comment. Newspaper notices were published

in the Carlsbad Current-Argus on June 22, 1996, The El Paso Times on

June 25, 1996, the Fort Stockton Pioneer on June 27, 1996, and in the

Van Horn Advocate on June 27 and July 4. We received five responses,

all supporting delisting. One response was from the U.S. Forest

Service, three were from botanists familiar with Lloyd's hedgehog

cactus and one was from the president of a landowner's group. One

response included a scientific paper published in 1995 after the

proposed rule had been drafted and transmitted to Washington, which was

not previously reviewed. This paper is cited in this final rule, and is

a slight extension of earlier work supporting the hybrid nature of

Lloyd's hedgehog cactus.

During the public comment period we invited peer review of the

conclusions and supporting information from four qualified systematic

botanists. In response we received two responses, both concurring that

Lloyd's hedgehog cactus is not a distinct species.

Summary of Factors Affecting the Species

After a thorough review and consideration of all information

available, we have determined that Lloyd's hedgehog cactus should be

removed from the List of Threatened and Endangered Plants. Procedures

found at section 4(a)(1) of the Act (16 U.S.C. 1531 et seq.) and

regulations implementing the delisting provisions of the Act (50 CFR

Part 424) were followed. The regulations at 50 CFR 424.11(d) state that

a species may be delisted if (1) it becomes extinct, (2) it recovers,

or (3) the original classification data were in error.

Since the time of listing, additional study has shown that Lloyd's

hedgehog cactus is not a distinct species but a hybrid. After a review

of the species' taxonomy, we conclude, based on the best scientific and

commercial information available, that the original listing decision

was based on a taxonomic interpretation subsequently demonstrated to be

incorrect. Lloyd's hedgehog cactus no longer qualifies for protection

under the Act because it does not conform with the definition of

species.

A species may be determined to be an endangered or threatened

species due to one or more of the five factors described in Section

4(a)(1). At the time of listing it was believed that Lloyd's hedgehog

cactus was a distinct species and that several of these factors were

relevant to its status. These factors and their application to

Echinocereus lloydii Britt. & Rose (Lloyd's hedgehog cactus) were

discussed in detail in the final rule (44 FR 61916) and included:

A. The present or threatened destruction, modification, or

curtailment of its habitat or range. The

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primary concern in our prior rulemaking was that Lloyd's hedgehog

cactus was vulnerable from past and potential habitat destruction due

to highway construction and maintenance, and the potential destructive

impacts of overgrazing in the rural rangeland habitat.

B. Overutilization for commercial, recreational, scientific, or

educational purposes. At the time of the final rule and continuing

today, Echinocereus lloydii is in world-wide demand by collectors of

rare cacti. Removal of plants from the wild has resulted in the

depletion of natural populations.

C. Disease or predation. At the time of listing it was felt that

Echinocereus lloydii, particularly young plants, could suffer possible

adverse affects from trampling by grazing cattle. The final rule

reported that light grazing did not seem to affect the species,

however, intensified grazing could threaten the continued existence of

E. lloydii.

D. The inadequacy of existing regulatory mechanisms. At the time

Echinocereus lloydii was listed, the states of Texas and New Mexico had

no laws protecting endangered and threatened plants. Since the listing,

both states have enacted protective laws and regulations for plants.

Lloyd's hedgehog cactus is on the New Mexico State List of Plant

Species (9-10-10 NMSA 1978; NMFRCD Rule No. 91-1) and on the Texas List

of Endangered, Threatened, or Protected Plants (Chapter 88, Texas Parks

and Wildlife Code).

On July 1, 1975, Appendix II of the Convention on International

Trade in Endangered Species of Wild Fauna and Flora (CITES) was amended

to include all members of the family Cactaceae. CITES is an

international treaty established to prevent international trade that

may be detrimental to the survival of plants and animals. A CITES

export permit must be issued by the exporting country before an

Appendix II species may be shipped. CITES permits may not be issued if

the export will be detrimental to the survival of the species or if the

specimens were not legally acquired. However, CITES does not regulate

take or domestic trade.

E. Other natural or manmade factors affecting its continued

existence. The final rule contained some discussion of the low numbers

of populations and the resulting restricted gene pool as a factor that

could intensify the adverse effects of other threats.

The determination that Lloyd's hedgehog cactus should be delisted

is based upon evidence that it is a hybrid that does not qualify for

protection under the Act, rather than on the control of threats. Since

Lloyd's hedgehog cactus is a hybrid which continues to be produced by

the two parent species, the number of E. lloydii populations is no

longer significant.

We have carefully assessed the best scientific and commercial

information available regarding the conclusion that Echinocereus

lloydii is a hybrid that does not qualify for protection under the Act

in determining to make this rule final. Based on this evaluation, the

preferred action is to remove Lloyd's hedgehog cactus from the list of

Endangered and Threatened Plants.

In accordance with 5 U.S.C. 553(d), we have determined that this

rule relieves an existing restriction and good cause exists to make

this rule effective immediately. Delay in implementation of this

delisting would cost government agencies staff time and monies on

conducting Section 7 consultation on actions which may affect the

Lloyd's hedgehog cactus, when this hybrid should no longer come under

the protection of the Act. Lifting the existing restrictions associated

with the listing of this species will enable Federal agencies to

minimize any delays in project planning and implementation for actions

that may affect Lloyd's hedgehog cactus.

Effects of the Final Rule

This action removes Lloyd's hedgehog cactus from the List of

Endangered and Threatened Plants. The Act and its implementing

regulations set forth a series of general prohibitions that apply to

all endangered plants. All prohibitions of section 9(a)(2) of the Act,

implemented by 50 CFR 17.61, currently apply to Lloyd's hedgehog

cactus. These prohibitions, in part, make it illegal for any person

subject to the jurisdiction of the United States to import or export,

transport in interstate or foreign commerce in the course of a

commercial activity, sell or offer for sale this species in interstate

or foreign commerce, or to remove and reduce to possession the species

from areas under Federal jurisdiction. In addition, for plants listed

as endangered, the Act prohibits the malicious damage or destruction on

areas under Federal jurisdiction and the removal, cutting, digging up,

or damaging or destroying endangered plants in knowing violation of any

State law or regulation, including State criminal trespass law. These

prohibitions will no longer apply to Lloyd's hedgehog cactus.

The requirements of Section 7 of the Act will also no longer apply

to Lloyd's hedgehog cactus and Federal agencies will no longer be

required to consult on their actions that may affect Lloyd's hedgehog

cactus.

The 1988 amendments to the Act require that all species which have

been delisted due to recovery be monitored for at least 5 years

following delisting. Lloyd's hedgehog cactus is being delisted because

the taxonomic interpretation that it is a valid species has been found

to be incorrect, and Lloyd's hedgehog cactus is an unstable hybrid

rather than a distinct taxon. Therefore no monitoring period following

delisting is required.

Some protection for Lloyd's hedgehog cactus will remain in place.

All native cacti, including hybrids, are on Appendix II of CITES. CITES

regulates international trade of cacti, but does not regulate trade

within the United States or prevent habitat destruction.

National Environmental Policy Act

We have determined that an Environmental Assessment, as defined

under the authority of the National Environmental Policy Act of 1969,

need not be prepared in connection with regulations adopted pursuant to

Section 4(a) of the Endangered Species Act of 1973, as amended. A

notice outlining the basis for this determination was published in the

Federal Register on October 25, 1983 (48 FR 49244).

References Cited

Benson, L. 1982. The cacti of the United States and Canada. Stanford

University Press, Stanford, California. 1044 pp.

Britton, N.L. and J.N. Rose. 1922. The Cactaceae. Vol. III 258:37-

38.

Correll, D.S., and M.C. Johnston. 1979. Manual of the vascular

plants of Texas. The University of Texas at Dallas, Richardson,

Texas. 1881 pp.

Poole, J.M., and D.H. Riskind. 1987. Endangered, threatened, or

protected native plants of Texas. Texas Parks and Wildlife

Department, Austin, Texas.

Poole, J.M., and A.D. Zimmerman. 1985. Endangered species

information system species record, Echinocereus lloydii. U.S. Fish

and Wildlife Service, Division of Endangered Species, Washington,

D.C.

Powell, A.M., A.D. Zimmerman, and R.A. Hilsenbeck. 1991.

Experimental documentation of natural hybridization in Cactaceae:

origin of Lloyd's hedgehog cactus, Echinocereus X lloydii. Plant

Systematics and Evolution 178:107-122.

Powell, A.M. 1995. Second generation experimental hybridizations in

the Echinocereus X Lloydii complex (Cactaceae), and further

documentation of dioecy in E. coccineus. Pl. Syst. Evol. 196:63-74.

U.S. Fish and Wildlife Service. 1985. Minutes of the Joint Meeting,

Region 2 Plant Recovery Teams, January 10-11, 1985. Region 2,

Albuquerque, New Mexico.

[[Page 33800]]

Weniger, D. 1979. Status report on Echinocereus lloydii. U.S. Fish

and Wildlife Service, Albuquerque, New Mexico. 11 pp.

Weniger, D. 1970. Cacti of the southwest. University of Texas Press,

Austin, Texas. 249 pp.

Zimmerman, A.D. 1993. Systematics of Echinocereus X roetteri

(Cactaceae), including Lloyd's hedgehog-cactus. Southwestern Rare

and Endangered Plants; Proceedings of the Southwestern Rare and

Endangered Plant Conference. Forestry and Resources Conservation

Division of the New Mexico Energy, Minerals, and Natural Resources

Department. Miscellaneous Publication 2:270-288.

Authors

The primary author of this document is Kathryn Kennedy, Austin

Ecological Services Field Office (refer to ADDRESSES section).

List of Subjects in 50 CFR Part 17

Endangered and threatened species, Exports, Imports, Reporting and

recordkeeping requirements, Transportation.

Regulation Promulgation

Accordingly, part 17, subchapter B of chapter I, title 50 of the

Code of Federal Regulations is amended, as set forth below:

PART 17--[AMENDED]

1. The authority citation for Part 17 continues to read as follows:

Authority: 16 U.S.C. 1361-1407; 16 U.S.C. 1531-1544; 16 U.S.C.

4201-4245; Pub. L. 99-625, 100 Stat. 3500; unless otherwise noted.

Sec. 17.12 [Amended]

2. Section 17.12(h) is amended by removing the entry for

``Echinocereus lloydii'' under ``FLOWERING PLANTS'' from the List of

Endangered and Threatened Plants.

Dated: May 13, 1999.

Jamie Rappaport Clark,

Director, Fish and Wildlife Service.

[FR Doc. 99-16029 Filed 6-23-99; 8:45 am]

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