(DRAFT) - Taxonomy
                          Species CACTUS, HEDGEHOG, LLOYD'S
                                Species Id ESIS702015
                                   Date 13 MAR 96



TAXONOMY

NAME - CACTUS, HEDGEHOG, LLOYD'S OTHER COMMON NAMES - CACTUS, HEDGEHOG and LLOYD'S ELEMENT CODE - CATEGORY - Angiosperm PHYLUM AND SUBPHYLUM - MAGNOLIOPHYTA, CLASS AND SUBCLASS - MAGNOLIOPSIDA, ORDER AND SUBORDER - CARYOPHYLLALES, FAMILY AND SUBFAMILY - CACTACEAE, GENUS AND SUBGENUS - ECHINOCEREUS, SPECIES AND SSP - LLOYDII, SCIENTIFIC NAME - ECHINOCEREUS LLOYDII AUTHORITY - TAXONOMY REFERENCES - COMMENTS ON TAXONOMY - Lloyd's Hedgehog Cactus Echinocereus lloydii Britt. and Rose KINGDOM: Plant GROUP: Angiosperm DIVISION: Magnoliophyta CLASS: Magnoliopsida ORDER: Caryophyllales FAMILY: Cactaceae Plants with one stem or several in a clump 15 to 20 cm high, 30 cm or more in diameter; stems green, cylindroid, 15 to 30 (-60, 01) cm long, 7.5 to 11.5 cm in diameter; ribs 11 to 13 in number, tuberculate; areoles circular, 6 to 16 mm apart; spines are partially obscuring the stem, red with a gray surface coating in appearance, straight and acicular; central spines number 2 to 8 per areole, spreading, to 12 mm long; radial spines number 8 to 17 per areole, spreading irregularly, to 15 (22,02) mm long; flower is 6 to 7.5 cm long, 5 to 9 cm in diameter, with much variation in color; the outer floral bracts have purplish midribs and lavender margins, narrowly elliptic-oblong, 3 cm long, 6 mm broad, obtuse, mucronulate, undulate; inner floral bracts are lavender or magenta to orangish red, cuneate- lanceolate, 4 cm long, 1.2 cm broad, rounded, mucronulate, entire; filaments 6 to 9 mm long; anthers variable from yellow to pink, 1 mm long; the style is 25 mm long; stigmas are green, 9 to 20 lobes, 9 mm Taxonomy - 1 (DRAFT) - Taxonomy Species CACTUS, HEDGEHOG, LLOYD'S Species Id ESIS702015 Date 13 MAR 96 long; the fruit is green tinged with pink or orange, ovoid, 25 to 30 (-50, 02) mm long, 12 to 20 mm in diameter, with white spines about 1.5 mm long; the seeds are black, strongly papillate, 1 mm long, about 1.5 mm broad. Chromosome number n=22 (02,03,04,05,06,07; Koole 45, UTEP). According to Weniger, the Eddy County, New Mexico populations are erroneously identified (02). However, Benson, Zimmerman and Knight feel these specimens to be correctly identified as Echinocereus lloydii (03,08,09). Backeberg placed E. lloydii as a variety of E. roetteri (05). Zimmerman thinks that roetteri might be the valid name for this taxon (08). Zimmerman also feels that Benson's interpretation of E. lloydii is too restricted, and that some of the localities asscribed to E. pectinatus var. minor actually represent E. lloydii (08). In the European horticultural literature, the Chihuahuan population of E. lloydii is being called E. pectinatus "var. rectispinus", according to Zimmerman (08). Due to the morphological variability within the species, both Knight and Zimmerman have suggested that the taxon may be of hybrid origin (between E. dasyacanthus and E. triglochidiatus) but stabilized reproductively (08,09). The Pecos County populations intergrade with E. dasyacanthus, resulting in a spectacular array of different flower colors (08). According to Benson, an invalid name of uncertain application, but perhaps referable to E. lloydii, is E. viridiflorus var. intermedius (03). However, Leuck submerges this name in E. viridiflorus var. cylindricus (10). The lectotype (F.E. Lloyd s.n.) is housed at the U.S. National Herbarium (US 691964) (03). Other specimens may be found at the following herbaria: NY, POM, UNM, and UTEP. Descriptions, photographs, and/or illustrations are found in several works: (01,02,03,04,05,06,07). Taxonomy - 2
                                  (DRAFT) - Status
                          Species CACTUS, HEDGEHOG, LLOYD'S
                                Species Id ESIS702015
                                   Date 13 MAR 96



STATUS

Coded Status E: Federal Endangered Commercial Non-consumptive recreational Ornamental COMMENTS ON STATUS - U.S. STATUSES AND LAWS: The Lloyd's hedgehog cactus (Echinocereus lloydii) has been designated an Endangered species pursuant to the Endangered Species Act of 1973 (50 CFR 17.12; P.L. 93-205, 87 Stat. 884; 16 U.S.C. 1531-1540), as amended. The species has this status wherever found including the States of New Mexico and Texas. Populations of this species which occur in Mexico are not listed pursuant to the Endangered Species Act. Removal and reduction to possession of any Federally listed plant from an area under Federal jurisdiction is unlawful (50 CFR 17.61 and 17.71). This species is protected by the Lacey Act (P.L. 97-79, as amended; 16 U.S.C. 3371 et seq.) which makes it unlawful to possess any wild plant (including roots, seeds, and other parts) within U.S. territorial or special maritime jurisdiction (as defined in 18 U.S.C. 7); or to import, export, transport, sell, receive, acquire, or purchase in interstate or foreign commerce any wild plant (including roots, seeds, and other parts) taken, possessed, transported, or sold in violation of any State law or regulation. It is also unlawful to import, export, transport, sell, receive, acquire, or purchase any wild plant (including roots, seeds, and other parts) taken or possessed in violation of any U.S. law, treaty, or regulation or in violation of Indian tribal law. RESPONSIBLE FEDERAL AGENCIES: USFWS -Responsible for the management/recovery, listing, and law enforcement/protection of this species. BLM -Responsible for the law enforcement/protection of this species with applicable State and Federal laws on public land under their control (43 CFR 4140). Also responsible for management/recovery on Bureau of Land Management lands. DOD -Responsible for the law enforcement/protection of this species with applicable State and Federal laws on public land under their control. Also responsible for management/recovery on Department of Defense lands. Status - 1 (DRAFT) - Status Species CACTUS, HEDGEHOG, LLOYD'S Species Id ESIS702015 Date 13 MAR 96 NPS -Responsible for the law enforcement/protection of this species with applicable State and Federal laws on public lands under their control. Also responsible for conservation (Nat. Park System Organic Act - 16 U.S.C. 1, 2-3)/management/recovery on National Park Service lands. Taking, possessing, or disturbing of Federally listed species is prohibited on NPS lands (36 CFR 2.1, 2.2, and 2.3). USFS -Responsible for the law enforcement/protection of this species with applicable State and Federal laws on public lands under their control. Also responsible for management/recovery on Forest Service lands. The Forest Service is responsible for integrating management, protection, and conservation of Federally listed species into the Forest Planning process (36 CFR 219.19 and 219.20). All Federal agencies have responsibility to ensure that any action authorized, funded, or carried out by that agency is not likely to jeopardize the continued existence of the species or result in the destruction or adverse modification of Critical Habitat (50 CFR 402), and to utilize their authorities to carry out programs for the conservation of the species. STATE STATUSES AND LAWS: STATE: New Mexico DESIGNATED STATUS: Protected ADMINISTRATIVE AGENCY: New Mexico Department of Natural Resources STATE STATUTE: New Mexico Statutes Annotated 1978, Endangered Plant Species Program STATE: Texas DESIGNATED STATUS: Endangered ADMINISTRATIVE AGENCY: Texas Parks and Wildlife Department STATE STATUTE: Chapter 88 of the Texas Parks and Wildlife Code, Texas Parks and Wildlife Rules for Protected, Threatened and Endangered Native Plant Species, Sec 57.401-57.413 INTERNATIONAL STATUSES, TREATIES, AND AGREEMENTS: This species is listed in Appendix I of CITES. ECONOMIC STATUSES: This species has horticultural value and is commercially available from cactus and succulent dealers. It's rarity makes it particularly desirable to collectors. It has value as a component of New Mexico and Texas's natural heritage of biotic diversity. 75/07/01:40 FR 27823/27924 - Notice of review 76/06/16:41 FR 24523/24572 - Proposed Endangered Status - 2 (DRAFT) - Status Species CACTUS, HEDGEHOG, LLOYD'S Species Id ESIS702015 Date 13 MAR 96 79/10/26:44 FR 61916/61917 - Listing as Endangered 85/07/22:50 FR 29901/29909 - Five year review Status - 3
     

HABITAT ASSOCIATIONS

HABITAT - TERRESTRIAL TERRESTRIAL LAND USE - Transportation, communications, and Util Shrub and Brush Rangeland Orchards, Groves, Vineyards, Nurseries, Bare Exposed Rock COMMENTS ON HABITAT ASSOCIATIONS - Echinocereus lloydii occurs in the shrub and brush rangeland of the Chihuahuan Desert. A major highway (IH 10, with a associated fenceline) runs through part of the habitat in Pecos County, TX. Echinocereus lloydii grows on sandy, gravelly, or rock soils on slopes and hillsides, as well as on bare rock ledges (02,03), and on fine-textured alluvial floors (08). The rock or gravel is usually limestone, occasionally with weathered metamorphic rock. The climate is hot (64 to 68 degrees F mean annual temperature) and arid (8 to 14 inches mean annual precipitation) (11). The elevation varies between 900 and 1650 meters (03). Ussually E. lloydii grows on open, fully exposed sites with very scattered forbs, grasses and brush (02). Sometimes it occurs in dense mesquite scrub among tall grasses (08). Some associates are Agave lecheguilla (02), Prosopis glandulosa (Weniger 183, UNM), Opuntia engelmannii, O. phaeacantha, O. imbricata, O. leptocaulis, Echinocereus dasyacanthus, E. stramineus, Coryphantha vivipara var. radiosa, Ferocactus hamatacanthus var. hamatacanthus, and Echinocactus horizonthalonius (12). A large area of scrub is visible from IH 10, not far east of Tunas Springs, has been cleared and might be a vineyard; more such developments might be forthcoming) (08). See the field "N-BIOLOGY" (Ecological/Edaphic Factors) for specific geological and soil associations. In Texas, one population of Lloyd's hedgehog cactus occurs in northeastern Culberson County in the Guadalupe Mountains National Park at the trailhead to the Williams Ranch at Highway 62-180. Habitat Associations - 1
                                (DRAFT) - Food Habits
                          Species CACTUS, HEDGEHOG, LLOYD'S
                                Species Id ESIS702015
                                   Date 13 MAR 96



FOOD HABITS

TROPHIC LEVEL - AUTOTROPH Food Habits - 1
                         (DRAFT) - Environment Associations
                          Species CACTUS, HEDGEHOG, LLOYD'S
                                Species Id ESIS702015
                                   Date 13 MAR 96



ENVIRONMENTAL ASSOCIATIONS

G = General A = Adult LIM = Limiting RA = Resting Adult J = Juvenile FA = Feeding Adult RJ = Resting Juvenile BA = Breeding Adult FJ = Feeding Juvenile P = Pupae L = Larvae E = Egg RL = Resting Larvae FL = Feeding Larvae
LIFESTAGE ENVIRONMENTAL ASSOCIATIONS G Terrestrial Features: Cliffs/ledges G Availability of fence rows: roadside ditches and grassy G G G G G G Environment Associations - 1
                               (DRAFT) - Life History
                          Species CACTUS, HEDGEHOG, LLOYD'S
                                Species Id ESIS702015
                                   Date 13 MAR 96



LIFE HISTORY

HABIT: Succulent (01,02,03,04,05,06,07). LIFE CYCLE: Perennial (02,03,04,05,06,07,08). TYPE OF REPRODUCTION: Sexual (02,08). REPRODUCTIVE PHENOLOGY: Germination and leafing dates are unknown. Budding and Anthesis occur in April to May (02,08). Fruiting occurs in mid-summer and seed/fruit dispersal dates are unknown (02,08). SEX OR SPORE STATUS: Monoclinous (03,04,05,06,07). Some individuals are partially male-sterile (few pollen grains, as compared to normal Echinocereus flowers) (08). This is consistent with the hypothesis of a hybrid origin of this species (08). POLLINATION, SPORE AND SEED DISSEMINATION: Pollen dissemination agents are insects (02). The seed dissemination agents are probably vertebrates (08). Seedlings are not concentrated close to the parent plants; presumably the ripe fruits are removed and consumed by vertebrates, so that the seeds do not fall close to the female plant (08). SEED BIOLOGY: Seeds germinate easily and rapidly in cultivation (02). The seed biology in nature is unknown. POPULATION BIOLOGY: According to Weniger (02), the population in Pecos County has declined drastically over the last 15 years. Here the population is composed of various sized individuals, but small seedlings were not observed (02). The cacti were numerous, with an individual every hundred feet, but most were destroyed by the highway or taken by collectors (02). A few miles east of the type locality, Zimmerman saw several dozen plants in six hours of searching; small seedlings were few and inconspicuous (08). The status of the other populations is unknown, but most of the individuals collected have been old adults (08). The plants 23 to 26 miles south of Fort Stockton are sparse; the 6 plants seen there thus far were all adults (08). In Pecos County, the complete range of intermediates between E. lloydii and the much-more-common E. dasyacanthus suggests introgression, which might eventually result in dilution (swamping) of the E. lloydii genepool (08). ECOLOGICAL/EDAPHIC FACTORS: E. lloydii grows on sandy, gravelly, or rock soils on slopes and Life History - 1 (DRAFT) - Life History Species CACTUS, HEDGEHOG, LLOYD'S Species Id ESIS702015 Date 13 MAR 96 hillsides, as well as on bare rock ledges (02,03), and on fine textured alluvial valley floors (08). The rock or gravel is usually limestone, occasionally with weathered metamorphic rock. The climate is hot (64 to 68 degrees F mean annual temperature) and arid (8 to 14 inches mean annual precipitation) (11). The elevation varies between 900 and 1650 meters (03). Usually E. lloydii grows on open, fully exposed sites with scattered forbs, grasses, and brush (02), although sometimes it occurs in dense mesquite scrub among tall grasses (08). Apparent nurse-plants (shrubs and perennial grasses) commonly are associated with E. lloydii plants; a definite requirement for nurse- plants is documented for seedlings of E. triglochidiatus var. gonacanthus, which has a similar growth-habit to E. lloydii, elsewhere in the Chihuahuan Desert (13). In Texas, geologic formations in the area of E. lloydii in Culberson County are the Brushy Canyon formation (mostly sandstone; Permian), and old and young Quaternary deposits (15). The soils in the area of occurrence of this species in Culberson County are not mapped (19). In Pecos County, the area of occurrence includes Holocene alluvium and fan deposits, Quaterary deposits, the Perdiz Conglomerate and similar gravel deposits (Quaternary and Tertiary), the Tessey and Capitan limestones (both Permian), and the Washita and Fredericksburg groups (limestone, dolomite, chert, and marl; lower Cretaceous) (15). E. lloydii occurs on aridisols and mollisols in Pecos County (16). The soil associations and series in this area are Ector, Ector-Rock Outcrop, Hodgins, Iraan, Lozier, Reagan, Sanderson, and Upton (16). In the vicinity of the Presidio County locality are young and old Quaternary deposits and Bolson deposits (clay, silt, sandstone, and conglomerates; Pleistocene to Miocene) (15). The soil orders are aridisols, entisols, and mollisols; and the soil associations are Glendale-Anthony-Toyah and Nickel-Canutio (18). In New Mexico, geologic formations in the area of E. lloydii in Dona Ana County are the Camp Rice and Sierra Ladrones formations (Quaternary), and the following Cretaceous formations: Sarten, Beartooth Sandstone, Anapra, Mesilla Valley, Muleros, and Smeltertown (17). Also in Dona Ana County, E. lloydii occurs on the Nickel-Upton association, Rock outcrop-Argids association, and Tencee-Upton association (20). In Eddy County, near Carlsbad, the formations are Quaternary alluvium, dunes, terraces, and pediments (17). The Sitting Bull Falls locality is on Permian limestone. In New Mexico in general, E. lloydii occurs on aridisols, entisols, and mollisols (20). The rather general locality of "near Carlsbad" includes numerous soil series: Atoka, Anthony, Arno, Dev, Ector, Cottonwood, Gypsum-land, Harkey, Karro, Kermit, Berino, Duneland, Pajarito, Pima, Reagan, Bippus, Simona, Tonuco, and Upton (20). In the Sitting Bull Falls area, the following soil complexes occur: Deama-Rock outcrop, Rock outcrop-Deama complex, and Rock outcrop-Tortugas-Ustiluvents complex (20). In Otero County, geologic formations in the area of E. lloydii are Pennsylvanian (the Magdalena Group) and Permian (Yeso and Abo- Hueco Formations) limestones with Tertiary intermediate to silicic intrusions (17). Also, in Otero County, the cactus is found on the Nickel-Tencee association and Rock outcrop (20). The very general locality of "30 miles west of Artesia" falls within Chaves County for which geologic associations include formations that are Quaternary Life History - 2 (DRAFT) - Life History Species CACTUS, HEDGEHOG, LLOYD'S Species Id ESIS702015 Date 13 MAR 96 alluvium, dunes, terraces, and pediments (17). In Chaves County, the soils in the area of occurrence are Ector-Rock complex, Lozier-Tencee complex, Pecos-Dev association, Reakor-Pecos association, and the Tencee cobbly loam (20). TROPHIC STATUS: Phototrophic (03,04,05,06,07). CHARACTERISTIC DOMINANCE: Component (08). COMMUNITY ECOLOGY: Echinocereus lloydii is a component of the Chihuahuan Desert, xeromorphic shrub, a stable community (14). In Pecos County, the complete range of intermediates between E. lloydii and the much-more-common E. dasyacanthus suggests introgression, which might eventually result in dilution (swamping) of the E. lloydii gene-pool (08). SPECIES INTERRELATIONSHIPS: Apparent nurse-plants (shrubs and perennial grasses) commonly are associated with E. lloydii plants; a definite requirement for nurse- plants is documented for seedlings of E. triglochidiatus var. gonacanthus, which has a similar growth-habit to E. lloydii, elsewhere in the Chihuahuan Desert (13). OTHER LIFE HISTORY DESCRIPTORS: At least in Pecos County, Texas, the purity of the lloydii gene- pool may be threatened by introgression from E. dasyacanthus, a much commoner and more widespread species (08). The apparent ease of hybridization between these species is consistent with the hypothesis that E. lloydii originated through former hybridization between the ancestors of E. dasyacanthus and E. troglochidiatus, which are sympatric with each other in many parts of west Texas and southern New Mexico (08). In the Jarilla Mountains, Otero County, New Mexico, the population identified as lloydii consists of (or is involved in) an apparent hybrid swarm involving at least three species of Echinocereus (E. dasyacanthus, E. troglochidiatus var. neomexicanus, and E. viridiflorus) that normally are sympatric without hybridization (08). These species are very different in appearance, and display pollination syndromes adapted for large bees, hummingbirds, and small bees, respectively (08). The hybrids involving E. viridiflorus are sterile or nearly so, consistent with the different ploidy level of that species; E. viridiflorus is a diploid, while E. lloydii, E. troglochidiatus var. neomexicanus, and E. dasyacanthus all are tetraploid (08). Life History - 3
                           (DRAFT) - Management Practices
                          Species CACTUS, HEDGEHOG, LLOYD'S
                                Species Id ESIS702015
                                   Date 13 MAR 96



MANAGEMENT PRACTICES

RESULT MANAGEMENT PRACTICE Beneficial Restricting Poaching Beneficial Controlling/Restricting Noncommercial Harvest Beneficial Regulating commercial harvest levels Adverse Collecting Existing Collecting Adverse Commercial Exploitation Existing Commercial Exploitation Adverse Gas/Oil Development Existing Gas/Oil Development Adverse Highway/Railroads Existing Highway/Railroads Adverse Soil compaction by heavy equipment in mine areas Existing Soil compaction by heavy equipment in mine areas COMMENTS ON MANAGEMENT PRACTICES - Part of the Echinocereus lloydii population in Pecos County, TX, was destroyed by the construction of Interstate Highway 10 (02). Highway and roadside maintenance threatens any individuals in the immediate vicinity. In this same general area, some plants are in a producing oil field which, so far, has had minimal effects on the species (02). By far the most serious threat comes from private and commercial collecting of the greatly prized and beautiful, rare collector's item (02). Future threats might be brush control management techniques, such as clearing or herbicides, and farming (a large area of scrub is visible from IH 10, not far east of Tunas Springs, has been cleared and might be a vineyard; more such developments might be forthcoming) (08). UNAPPROVED PLAN: No target date for the initiation of a Recovery Plan has been established as of 1986. There is an ongoing study to determine whether Echinocereus lloydii is actually a distinct taxon. Until this is determined, a Recovery Plan will not be undertaken. In the interim it is recommended that the species be managed through enforcement of existing collecting and trade regulations, and protection of habitat from Federal actions contiune through Section 7 of the Endangered Species Act. Management Practices - 1
                                   (DRAFT) - References
                            Species CACTUS, HEDGEHOG, LLOYD'S
                                  Species Id ESIS702015
                                      Date 13 MAR 96



     

References

References - 1 (DRAFT) - References Species CACTUS, HEDGEHOG, LLOYD'S Species Id ESIS702015 Date 13 MAR 96 ***** REFERENCES FOR ALL NARRATIVES EXCEPT N-OCCURRENCE ***** 01 Warnock, B.H. 1974. Wildflowers of the Guadalupe Mountains and Sand Dune Country, Texas. Sul Ross State Univ., Alpine. 02 Weniger, D. 1979. Status report on Echinocereus lloydii. On file at: U.S. Fish and Wildlife Service. 03 Benson, L. 1982. The cacti of the United States and Canada. Stanford Univ. Press, Stanford. 04 Weniger, D. 1970. Cacti of the southwest. Univ. of TX Press, Austin. 05 Benson, L. 1969. Cactaceae in flora of Texas, Vol. II. Texas Research Found., Renner. 06 Correll, D.S. and M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner. 07 Britton, N.L. and J.N. Rose. 1963. The Cactaceae, Vol. III. Dover Public., Inc., NY. 08 Zimmerman, A.D. 1984. Pers. comm. Department of Biology, University of Texas, Austin, TX 78712. 09 Knight, P. 1984. Pers. comm. Department of Natural Resources, Villagra Building, Santa Fe, NM 87503. 10 Leuk, E.E., II. 1980. Biosystematic studies in the Echinocereus viridiflorus complex. Ph.D. dissertation. Univ. of OK, Norman. 11 Godfrey, C.L., G.S. McKee and H. Oakes. 1973. General soil map of Texas. TX Agri. Exp. Sta., Texas A and M Univ., College Station. 12 Department of Interoir. 1979. Environmental Assessment. 13 Lozano, R. and W. Reid. 1982. Claret cup cactus at White Sands National Monument. Cact. and Succ. J. (US) 54:196-201. 14 Diamond, D.D. 1984. Pers. comm. 15 Bureau of Economic Geology. 1965-1983. Geologic atlas of Texas. Univ. of Texas, Austin. 16 Rives, J.L. 1980. Soil survey of Pecos County, Texas. USDA. 17 Mew Mexico Geological Society. 1982. New Mexico Highway Geologic Map. 18 Soil Conservation Service. 1973. General soil map, Presidio County, Texas. 19 Rives, J.L. 1985. Pers. comm. 20 Ruiz, J.E. 1985. Pers. comm. ***** REFERENCES FOR N-OCCURRENCE NARRATIVE ONLY ***** 01 Zimmerman, A.D. 1984. Pers. comm. Department of Biology, University of Texas, Austin, TX 78712. 02 Knight, P. 1984. Pers. comm. Department of Natural Resources, Villagra Building, Santa Fe, NM 87503. 03 Benson, L. 1982. The cacti of the United States and Canada. Stanford Univ. Press, Stanford. 04 Weniger, D. 1970. Cacti of the southwest. Univ. of TX Press, Austin. 05 Warnock, B.H. 1974. Wildflowers of the Guadalupe Mountains and Sand Dune Country, Texas. Sul Ross State Univ., Alpine. References - 2