[Federal Register: June 24, 1999 (Volume 64, Number 121)]

[Rules and Regulations]               

[Page 33796-33800]

From the Federal Register Online via GPO Access [wais.access.gpo.gov]





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.


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.

[[Page 33797]]

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).



    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 


[[Page 33798]]

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 


    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

[[Page 33799]]

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 


    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-


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, 


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.


    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:


    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]