(DRAFT) - Taxonomy
                           Species FERN, SHIELD, ALEUTIAN
                                Species Id ESIS857001
                                   Date 13 MAR 96



TAXONOMY

NAME - FERN, SHIELD, ALEUTIAN OTHER COMMON NAMES - FERN, SHIELD, ALEUTIAN; FERN, SHIELD; FERN, HOLLY and ALEUTIAN ELEMENT CODE - CATEGORY - Fern/F-Ally PHYLUM AND SUBPHYLUM - POLYPODIOPHYTA, CLASS AND SUBCLASS - FILICOPSIDA, ORDER AND SUBORDER - FILICALES, FAMILY AND SUBFAMILY - POLYPODIACEAE, GENUS AND SUBGENUS - POLYSTICHUM, SPECIES AND SSP - ALEUTICUM, SCIENTIFIC NAME - POLYSTICHUM ALEUTICUM AUTHORITY - TAXONOMY REFERENCES - COMMENTS ON TAXONOMY - Aleutian Shield Fern Polystichum aleuticum C. Christens. in Hulten KINGDOM: Plant GROUP: Fern or fern ally DIVISION: Polypodiophyta CLASS: Filicopsida ORDER: Filicales FAMILY: Polypodiaceae Non-technical Description (derived from 20,23,24,25,30). Plants small and erect, about six inches tall, usually solitary, fibrous-rooted with a thickened base bearing numerous chestnut-brown leaf base remnants. Leaves (fronds) present only during growing season, light, dark or olive green in color, gradually tapering at the base and tip; blades once divided into slightly overlapping segments; the segments ovate, slightly bulging at base, incised with short bristle tips. Scales sparingly distributed along main leaf axis and especially on under-surface of leaf segments, delicate, straw-colored, lance-shaped up to 3-4 mm long. Spore masses aggregated into 6-8 patches forming two rows along under-surface of leaf segments. Technical Description (derived from 30). Plants solitary, occasionally clumped, fibrous-rooted, erect to 10-18 cm tall, bearing numerous (10) - 30-50 - (or more) elaminate, persistent Taxonomy - 1 (DRAFT) - Taxonomy Species FERN, SHIELD, ALEUTIAN Species Id ESIS857001 Date 13 MAR 96 chestnut-brown stipe remnants. Leaves herbaceous, produced annually 7-15 cm long, pinnately compound; laminae linear, 3/4 or greater the leaf length, to 1.5 cm wide, tapering at base and tip. Pinna pairs (15) - 18-22 - (25), opposite to subopposite at base, alternate above, subalternate and pinnatifid at tip, sessile and disposed perpendicular to axis, 7 mm long x 4.5 mm wide at mid-lamina, gradually reduced at base and tip, not or barely touching below midleaf, succubously overlapping above, subsymmetrically ovate, incised-lobate to/and irregularly serrate with mucronate, subhyaline tips. Stipe and rachis scales crowded in vernation, tufted at pinna attachment, scattered along rachis in expanded fronds, paleaceous, brunneo-stramineous, lanceolate to 3 - (5) mm long, weakly to conspicuously lacinate at base, sparingly and occasionally retrorsely denticulate along margins; pinna scales somewhat reduced in size, otherwise similar, conspicuous on the soral surface, absent or rare on the non-soral surface except at frond tip. Sori normally 6 - 8 - (10) per pinna in two median rows. Indusia persistent, some confluent to barely overlapping, irregularly peltiform, excentrically attached; margins erose. Spores 64 per sporangium, (41) - 47-49 - (52) um greatest exospore diameter, granulose with perine weakly alveolate of low crests and papillae; maturing (? July) after mid-August. Classification and Nomenclature. The Aleutian shield fern (holly fern) Polystichum aleuticum C. Christens. has no species synonyms in scientific nomenclature (21). The complete author citation should read C. Chr. in Hulten (17) giving credit to the original description prepared by Christensen albeit published by Hulten. Common names referring to this species include: Shield Fern (unnamed) (09); Shield Fern (Holly Fern) Aleutian (10,11,12); Aleutian Shield-Fern (13,14,24,25,29,33); and Aleutian Holly-Fern (22,24,25,33). An acceptable classification above species level follows Bold et al. (06): Division Pteridophyta (Ferns); Class Filicopsida; Order Filicales; Family Polypodiaceae (04,08,09,10,11,12,13,14,33). Other authorities have placed this species within the fern families Aspidiaceae (20), Aspleniaceae (21), and Dryopteridaceae (22). Various authorities have treated this species through technical descriptions (Christensen in 17,22,23,33), short diagnoses (20,32), non-technical descriptions (23,24,25), and taxonomic comments (07,18,28). Type Specimens. Specimens of the original collection (1932) include the holotype at the Swedish Museum of Natural History, Stockholm (S) (32); isotypes at the California Academy of Sciences, San Francisco (CAS) (32); Dudley Herbarium of Stanford (DS) (32) at (CAS); and the Smithsonian Institution, Washington, D.C. (US) (32). Photographs of the type and their location have been catalogued by Lipkin (23) citing: 1) U.S. National Herbarium Vascular Plant Type Collection, microfiche #1027, column 4, row 11 (isotype); 2) University of Michigan, negative #1306; and 3) Washington State University, negatives #273437, #273438. Additional photographs appear in Hulten (17), presumably the holotype (S); and Lellinger (22) representing material collected on Adak in 1975. Additional Specimens. Specimens prepared from collections in 1975 on Adak (28) are deposited in herbaria of the University of Alaska, Fairbanks (ALA), the University of Tennessee, Knoxville Taxonomy - 2 (DRAFT) - Taxonomy Species FERN, SHIELD, ALEUTIAN Species Id ESIS857001 Date 13 MAR 96 (TENN), and the University of Oregon, Eugene (ORE). A single specimen collected in 1987 on Adak (29) is deposited at the Smithsonian Institution (US). Line Drawings. Line drawn images of habit profile are presented by Hulten (20). Hulten's figure has been reproduced and enlarged by Murray (24). An original illustration including pinna detail recently has been produced by Murray and Lipkin (25). Current Status. Fish and Wildlife Service, Interior, has recently issued (1988) a Final Rule determining endangered status for Polystichum aleuticum (14) effective March 18, 1988. Taxonomy - 3
                                  (DRAFT) - Status
                           Species FERN, SHIELD, ALEUTIAN
                                Species Id ESIS857001
                                   Date 13 MAR 96



STATUS

Coded Status E: Federal Endangered Non-consumptive recreational COMMENTS ON STATUS - U.S. STATUSES AND LAWS: The Aleutian shield fern (Polystichum aleuticum) 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 State of Alaska. 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. BIA -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 Bureau of Indian Affairs 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. 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. Status - 1 (DRAFT) - Status Species FERN, SHIELD, ALEUTIAN Species Id ESIS857001 Date 13 MAR 96 STATE STATUSES AND LAWS: STATE: Alaska DESIGNATED STATUS: None. INTERNATIONAL STATUSES, TREATIES, AND AGREEMENTS: CITES I. ECONOMIC STATUSES: There are no known cultural, commercial, or medicinal values associated with the species; however, it is of significant scientific value. The species is likely a relict of tertiary flora and thus is of great value in studies of island biogeography and the paleoecology of Beringia (see Lipkin (23)). The species is also of recreational value to amateur botantists. No negative values have been associated with this species. 75/01/09:40 FR 27823/ - Petition acceptance 76/06/16:41 FR 24523/ - Proposed rule/endangerd 78/04/26:43 FR 17909/ - Summary of comments 79/12/10:44 FR 70796/ - Proposal withdrawl 80/12/15:45 FR 82480/ - Notice of review 85/09/27:50 FR 39526/ - Notice of review 87/04/24:52 FR 13729/13732 - Proposed rule/endangered 88/02/17:53 FR 04626/04630 - Final rule/endangered Status - 2
     

HABITAT ASSOCIATIONS

HABITAT - TERRESTRIAL TERRESTRIAL LAND USE - Tundra Land: Shrub Brush Tundra Land: Herbaceous Tundra Land: Mixed COMMENTS ON HABITAT ASSOCIATIONS - Habitat details of the Aleutian shield fern are known only from two populations discovered on Adak Island in 1975 and 1987. Whereas little attention was paid to site characters of the 1975 population, field notes and observations were especially recorded after discovery of the 1987 population (31). Nothing can be said of the Atka collection (1932) because no data were recorded by the collector as to site location or habitat except for "very rare" (23). The index of Forest/Rangeland Associations (this document) that most closely references the general aspect of shield fern habitat are: Aleutian alpine tundra (code 53 Empetrum - Vaccinium and barren). There are no existing nor expected agricultural practices that will modify or displace the fern habitat. The species is not associated with any National Wetland Inventory types. The fern's habitat consists of exposed, weathered rock outcrops with rooting substrate confined to fissures, crevices, and thinly mantled horizontal ledges (section 5, this document). The habitat is apparently unique, restricted to rare combinations of aspect, exposure, substrate stability, and moisture (31). There is no defined composition of vegetation as per Aleutian tundra classifications proposed by Hulten (19), Skacklette (27), or Amundsen (02). Broadly circumscribed units of vegetation in proximity to shield fern habitat correspond to Anderson et al. (03) categories of Mixed Tundra consisting of low profile Shrub and Brush Tundra, upland Herbaceous Tundra, and Bare Ground Tundra, where features of surface relief (e.g. angle of slope, aspect, thickness and stability of soil mantle, and soil moisture of rhizosphere) dictate pattern and size of plant community patches (31). The 1987 population is confined within an area of approximately 100 sq. ft. on the east face of the north arm of Mt. Reed at 1600 feet elevation (29). That of the 1975 population was growing in a linear rock crease approximately 10 feet long at a lower (unmeasured) estimated elevation of 1300 feet (31). Both populations occurred similarly on near vertical, tilted rocky outcrops that are heavily fractured with crisp lines of vertical and horizontal cleavages. The sites are totally exposed to conditions of climatic weathering, mostly fog-enshrouded during the summer months. Site conditions during the winter months are undocumented, but the dormant perennating shoot crowns probably experience alternating periods of snow or ice cover, wind abrasion, and higher insolation. Soil accumulations in the rock crevices are the rooting medium, consisting of dark, extremely fine-grained particles. Moisture does not appear to be limiting anytime during the active growing season (31). Measured microsite field conditions are unknown. Weather data compiled by Lipkin (23) for Adak (22 yr. record) and Atka (1982-1983) show similar profiles for measured air temperatures and precipitation; although York (35) noted Atka is rainer than Adak. Summary data from Adak (recorded near the coast) Habitat Associations - 1 can only be used to generally predict the climate of Mt. Reed where the conditions are undoubtedly colder with much more snow, wind, and fog (23). Twenty year mean precipitation averages for Adak (02) amount to 168 cm/yr. distributed more or less evenly with slightly drier late Spring and wetter later Autumn conditions. Additional, more specific weather data for Adak is reported by Williams (34). Mean annual temperature measured from 1950-1972 was 4.9 deg. C (40.9 deg. F); and average monthly precipitation was 13.9 cm, the lowest in July and highest in November. Winds of greater than 20 knots occur 295 days per year, directed W-SW during 10 months and W during June and July (34). Ground surface solar radiation measured for 41 days between July 29 and Sept. 5, 1978 yielded data that for 19 inputs exceeded 0.5 cal./cm sq./min/ for greater than 4 hours with a maximum high of 1.21 cal./cm sq./min. on August 8, and a minimum low of 0.31 cal./cm sq./min. on Aug. 7 and 20 (34). Habitat Associations - 2
                                (DRAFT) - Food Habits
                           Species FERN, SHIELD, ALEUTIAN
                                Species Id ESIS857001
                                   Date 13 MAR 96



FOOD HABITS

TROPHIC LEVEL - AUTOTROPH Food Habits - 1
                         (DRAFT) - Environment Associations
                           Species FERN, SHIELD, ALEUTIAN
                                Species Id ESIS857001
                                   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 Terrestrial Features: Rock outcrops G Environment Associations - 1
                               (DRAFT) - Life History
                           Species FERN, SHIELD, ALEUTIAN
                                Species Id ESIS857001
                                   Date 13 MAR 96



LIFE HISTORY

HABIT: Herb (07,13,14,17,20,22,23,24,25,30,33). LIFE CYCLE: Perennial (07,13,14,17,20,22,23,24,25,30,33). The Aleutian shield fern is a non-evergreen, herbaceous perennial as is typical of many arctic-alpine species of ferns. Seasonally produced fronds do not overwinter, being renewed each year from the perennating shoot crown. Fronds of the next season's growth are initiated during the Summer and Fall of the current growing season; but remain tightly coiled and protected within the sheaflike cluster of persistent stipe remnants until the following Spring (31). TYPE OF REPRODUCTION: Sexual (07,17,20,22,23,30,32,33). All stages of the Aleutian shield fern's life cycle have not been observed. As is typcial of all "true" ferns, one full generation consists of a dibiontic (2 living phases) alternation between haploid (1N) and diploid (2N) phases. Spores that may germinate will form a delicate, inconspicuous haploid plant that bears male and/or female sex organs when mature. Subsequent to fertilization, an embryo develops and grows into the perennial diploid plant (that which has been collected to date). Sexual reproductive success hinges on the ability of the species to complete both haploid and diploid phases (31). This species regularly produces spores (17,30,31) that show no evidence of abnormal development. Many ferns are known to reproduce asexually or "pseudosexually" by replicating either/or both phases without union of sperm and egg. Such diasporic or apogamous mechanisms do not appear operational in this species. The apparent lack of specialized asexual mechanisms is supported by the depauperate number of individuals and spotty REPRODUCTIVE PHENOLOGY: Germination dates are unknown (30,31). Leafing dates occur in in Aleutian Spring - June(?) (30,31). Spore maturation and dispersal dates are from July to September (30,31). This species appears only to propagate (though infrequently) through spores. Although spores are produced annually, successful sowing and germination are infrequent. If spores germinate in the Fall after dispersal or in the following Spring is unknown (31). Annual production of leaves (is assumed to) occurs after the habitat becomes snow-free during the Aleutian Spring. Eyerdam's 1932 Atka specimen collected on July 5 bears fully developed leaves and mature spores. Adak specimens collected on Sept. 19, 1985, also contained mature fronds and ripened spires; while plants observed on on Aug. 17, 1987, only contained unripened, immature spores. Ripened spores are readily released during the season of production (31). SEX OR SPORE STATUS: Homosporous (30,31). Most species of the genus Polystichum produce a bisexual haploid phase, often with in-crossing barriers Life History - 1 (DRAFT) - Life History Species FERN, SHIELD, ALEUTIAN Species Id ESIS857001 Date 13 MAR 96 that inhibit or precent self-fertilization (15). The low, highly restricted gene pool of this species, coupled with an out-crossing requirement, realistically may account for the low number of populations and individuals (31). POLLINATION, SPORE AND SEED DISSEMINATION: Wind and/or seepage moisture are the spore disseminating agents for this species (23,31). Spore dispersal affected by wind may result in most spores being blown off the islands or, at least, away from suitable habitat (23,31). SEED BIOLOGY: Virtually nothing is known about the spore biology of this species. Spores are produced annually, in sporangia, clustered into fertile patches (sori) on the undersurface of mature leaves. Six to eight patches containing numerous sporangia (estimated >25) are produced on most of the segments of each leaf. Sixty-four spores are produced per sporangium measuring between 41 and 52 um in exospore diameter (31). POPULATION BIOLOGY: The following discussion is purely conjectural except for data relating to population density. One population of unknown number of individuals is known to (have) occur(ed) on Atka (17). Two populations are known from Adak, one containing an estimated number of 12-15 individuals (partially depleted by collecting) in 1975 (23,28), and another discovered in 1987 containing 7 individuals (5 sexually mature plants, 2 immature plants) now reduced to 6 individuals because of substrate release of one plant (29). Undoubtedly population isolation, low gene pool, virtual lack of interpopulational crossing, and specific requirements of habitat are the species' principal limiting factors. Reproduction, survival, and mortality rates are unknown. All populations may be very old, possibly declining, as the natural course of mortality is not offset by growth of new individuals (23). Recovery potential through natural processes is doubtful. Some enhancement might be possible through cultivation if a protocol could be developed that would introduce biotypes from different populations through crossing experiments. Such manipulations would require destructive sampling of existing populations to obtain spores. Until more source material is discovered, propagation is not advisable, if not, impracticable (29,31). ECOLOGICAL/EDAPHIC FACTORS: Habitat characteristics of the 1987 Aleutian shield fern population consist of exposed, weathered rock outcrops with rooting substrate confined to fissures, crevices, and thinly mantled horizontal ledges. The site is fully exposed to conditions of climatic weathering. Dense fog blankets the habitat during the summer months, except for brief periods when clear weather prevails. Hence, during the growing season, incident solar radiation is intercepted by fog that significantly lowers insolation values at the substrate Life History - 2 (DRAFT) - Life History Species FERN, SHIELD, ALEUTIAN Species Id ESIS857001 Date 13 MAR 96 surface. Ground surface solar radiation values measured for 41 days during the summer, 1978 on Adak are profiled by Williams (34). Temperature and precipitation data (02,23,34) indicate mean annual values of 4.9 deg. C and 13.9-16.8 cm/yr. These values may only predict general climatic conditions on Mt. Reed where temperatures are undoubtedly lower and much more snow, wind, and fog prevail (23). Winds of greater than 20 knots occur 295 days per year (34). Winds of greater force are experienced at the upper elevations on Mr. Reed (31). Fern habitat is situated between 1300 and 1600 feet elevation on the east exposure of the north arm of Mt. Reed. It is located in the South Adak region of folded, faulted and intensely altered basaltic flows and pyroclastic rocks (Finger Bay Volcanics) (23,26). The steep (50-75 deg.) upper talus/rubble slopes of Mt. Reed are interrupted by near vertical, tilted rocky outcrops bearing crisp, horizontal and vertical, fractures. Soils of such upland sites consist of cryandepts and cryofolists and lithic cryofolists (23) that are stabilized by low profile mixed Shrub-Forb-Herb vegetation. Soil that accumulates to the surface of outcrop creases consists of dark, extremely fine-grained particles. Texture and stickiness indicate high (constant) moisture and mixed mineral/organic composition (31). Natural disturbance to the habitat results from wasting of the outcrops. Superficial weather-abraiding effects may delay or prematurely halt the annual growth cycle in years with abnormal weather cycles. More severs impacts may result from freeze-thaw cycles that loosen and pare away blocks of the outcrop. Downslope creep of the vegetated talus slopes could contribute to destabilization of upslope outcrops. Such threats may be magnified by hiking or climbing near the site creating breaks in the vegetation mat or dislodging rock (31). TROPHIC STATUS: Phototrophic (31). CHARACTERISTIC DOMINANCE: Component (29,31). The Aleutian shield fern is affiliated with a loosely defined enclave of species without obvious characteristics of dominance (29,31). COMMUNITY ECOLOGY: Species associated with the 1987 shield fern population are: vascular plants; Cardamine bellidifolia, Carex spp., Cassiope lycopodioides, Empetrum nigrum, Geum rossii, Lycopodium selago, Oxyria digyna, Polygonum viviparum, Saxifraga punctata subsp. insularis, and Saxifraga unalaschensis; bryophytes, Mnium magnifolium and Rhytidiadelphus squarrosus (29). It is unlikely that forces of competitive displacement, positively or negatively, regulate population size or dynamics among all species present in the habitat (29). SPECIES INTERRELATIONSHIPS: Not known; relationships remote or absent (31). Life History - 3 (DRAFT) - Life History Species FERN, SHIELD, ALEUTIAN Species Id ESIS857001 Date 13 MAR 96 OTHER LIFE HISTORY DESCRIPTORS: None. Life History - 4
                           (DRAFT) - Management Practices
                           Species FERN, SHIELD, ALEUTIAN
                                Species Id ESIS857001
                                   Date 13 MAR 96



MANAGEMENT PRACTICES

RESULT MANAGEMENT PRACTICE Beneficial Stocking captive-reared wild-strain animals Beneficial Maintaining undisturbed/undeveloped areas Beneficial Transplanting wild animals Beneficial Restricting/regulating human disturbance of populations Beneficial Controlling/Removing Exotic Vertebrates Adverse Collecting Existing Collecting Adverse Low Gene Pool Existing Low Gene Pool COMMENTS ON MANAGEMENT PRACTICES - Past and Present Reasons for Status. The Aleutian shield fern is is among the rarest vascular plants in North America with a total known distribution restricted only to Atka and Adak Islands (13,14,23,24,25,28,29). Causes for such rarity are probably most profoundly historical factors related to Pleistocene and Holocene paleoclimates (31). That the Aleutian shield fern is derived from Asiatic ancestral stock is a reasonable assumption, since it is most closely related to species now found mainly in mountainous areas of the Sino-Himalayan region. Plant colonization of the Aleutians undoubtedly resulted, in part, from sources situated to the west (Kamchatka and Siberia). Spores of ancestral stock most likely were conveyed from the west by winds during the Late Tertiary (Pliocene) or more recently during the Pleistocene or Holocene. Once founded on the Aleutians, this species may have radiated along the island chain, but probably was never plentiful. Episodic glaciation through the Pleistocene likely limited and/or decimated populations to the extent that only biotypes remain today (31). The recent Holocene climate of the Aleutians is estimated at 10,000 yr. BP (01,05), following deglaciation of Wisconsin ice that capped the mountainous island masses. The Holocene hypsithermal was replaced by a climate of "...lower temperatures and greater storminess..." (16) beginning about 3000 yr. BP. Post-hypsithermal decay of the Aleutian climate may account for population decline and isolation evidenced today by extreme distributional restriction. It is conceivable also, that the Aleutian shield fern became founded in the Aleutians as recently as the period of early Holocene climatic amelioration; and that population decline and isolation has resulted only in the past 3000 years (31). Rarity of this species equates temporal factors of: source and availability of founding propagules (presumably spores); time(s) of arrival; success in establishment; repreoduction and radiation, or lack thereof; and persistence in place (29). The current status of this species would indicate that few and isolated populations comprise a restricted gene pool with little or no inter- or intra-populational gene flow. The potential for outcrossing between populations is remote if at all, and intra-populational gene flow Management Practices - 1 (DRAFT) - Management Practices Species FERN, SHIELD, ALEUTIAN Species Id ESIS857001 Date 13 MAR 96 appears equally unlikely given the low number of reproductively mature individuals and the idiosyncracies of the fern sexual life cycle (29,31). The apparent inability of this species to significantly/frequently reproduce is consistent with field observations made from both populations occurring on Adak. Plants of both populations lack any evidence of producing specialized asexual diasporic structures or haploid alternate phase plantlets (31). Two juvenile plants (sexually immature) were discovered in the 1987 Adak population suggest that some reproduction still does occur. However, the origin and age of these individuals can be presumed only as recent and derived from within the population. The expectation that they will develop to sexual maturity is uncertain (31). Reproductive rate and frequency may be so low that premature failure or insufficient population replenishment may not offset later mortality (23). Future Perceived Threats. Population decline due to natural consequences of habitat alteration, scarcity, low gene pool, and biological fitness of the species has been compounded by unintended anthropogenic insults on both Atka and Adak Islands. Collecting for scientific purposes is a known cause for loss of individuals from all known populations. There is now an adequate specimen record for this rare species, and, until additional populations are discovered, future collecting should be strictly prohibited (29). Unknown negative impacts may have resulted from introduced ungulates on both islands. European reindeer were introduced onto Atka in 1914 and have severely overgrazed the west end of the island (13,14,23). Caribou, introduced onto Adak in 1958 (13), have not yet shown the same degree of impact. At least for the Adak populations, they are situated on steep rocky outcrops where caribou foraging would be improbably (29). The most predictable future threats are mediated by human activity. Site visitation to known populations or surface-resource land development will gradually alter or certainly destroy suitable habitat. Such threats can easily minimized or eliminated by careful management. Areas where the fern is known to occur should be declared critical habitat, but markers or exclosures should be avoided so as not to draw attention to or destabilize these areas (29). UNAPPROVED PLAN: Currently, no recovery plan. Development of a recovery plan - if determined to be beneficial to the species - will be initiated in 1989. No approved recovery plan due to the recent date of listing and current lack of information on reproductive ecology and limiting factors. Current and planned activities to benefit the species include education and field surveys to locate additional specimens/populations. The Service is also coordinating with the U.S. Navy to restrict human access to the Adak Island population site. Recovery activities in the future may include controlling exotic Management Practices - 2 (DRAFT) - Management Practices Species FERN, SHIELD, ALEUTIAN Species Id ESIS857001 Date 13 MAR 96 vertebrates with fencing, transplanting and/or cultivating individuals for reintroduction and restricting access and development. Management Practices - 3
                                   (DRAFT) - References
                              Species FERN, SHIELD, ALEUTIAN
                                  Species Id ESIS857001
                                      Date 13 MAR 96



     

References

***** REFERENCES FOR ALL NARRATIVES EXCEPT N-OCCURRENCE ***** 01 Ager, T.A. 1982. Vegetational History of Western Alaska During the Wisconsin Glacial Interval and Holocene. pp. 75-92. In D.M. Hopkins et al. Paleoecology of Beringia. Academic Press, New York. 02 Amundsen, C.C. 1985. Central Aleutian Tundra: Ecological Manifestations of Maritime Tundra Landscapes in the Central Aleutian Islands (Amchitka, Adak) Alaska. Final Report prepared for Department of Energy, ORO-4180-10. 77 pp. + appendices I,II,III. 03 Anderson, J.R., E.E. Hardy, J.T. Roach, and R.E. Witmer. 1976. A Land Use and Land Cover Classification System for Use with Remote Sensor Data. Geol. Sur. Prof. Paper 964. 28 pp. 04 Ayensu, E.S., and R.A. DeFilipps. 1978. Endangered and Threatened Plants of the United States. Smithsonian Institution and World Wildlife Fund. Washington, D.C. 403 pp. 05 Black, R.F. 1976. Influence of holocene climatic changes on Aleut expansion into the Aleutian Islands, Alaska. Anthropological Papers, Univ. Alaska. 18: 31-42. 06 Bold, H.C., C.J. Alexopoulos, and T. Delevoryas. 1987. Morphology of Plants and Fungi. 5th Ed. Harper and Row, New York. 912 pp. 07 Christensen, C. 1938. On Polystichum aleuticum C. Chr., a new North American species. Amer. Fern J. 28: 111-112. 08 Department of Interior. Fish and Wildlife Service. 1975. Threatened or Endangered Fauna or Flora: Review of Status of Vascular Plants and Determination of "Critical Habitat." Federal Register 41: 27826. 09 Department of Interior. Fish and Wildlife Service. 1976. Endangered and Threatened Species: Plants. Federal Register 41: 24560. 10 Department of Interior. United States Fish & Wildlife Service. 1980. Endangered and Threatened Wildlife and Plants: Review of Plant Taxa for Listing as Endangered or Threatened Species. Federal Register 45: 82532. 11 Department of Interior. Fish and Wildlife Service. 1983. Endangered and Threatened Wildlife and Plants: Supplement to Review of Plant Taxa for Listing: Proposed Rule. Federal Register 43: 53657. 12 Department of Interior. Fish and Wildlife Service. 1985. Endangered and Threatened Wildlife and Plants; Review of Plant Taxa for Listing as Endangered or Threatened Species; Notice of Review. Federal Register 50: 39573. 13 Department of Interior. Fish and Wildlife Service. 1987. Endangered and Threatened Wildlife and Plants; Proposed Endangered Status for Polystichum aleuticum. Federal Register 52: 13729-13732. 14 Department of Interior. Fish and Wildlife Service. 1988. Endangered and Threatened Wildlife and Plants; Determination of Endangered Status for Polystichum aleuticum. Federal Register 53: 4626-4630. 15 Evans, A.M. 1988. Personal communication. Department of Botany, References - 1 (DRAFT) - References Species FERN, SHIELD, ALEUTIAN Species Id ESIS857001 Date 13 MAR 96 University of Tennessee, Knoxville, Tennessee. 16 Heusser, C.J. 1978. Postglacial vegetation on Adak Island, Aleutian Islands, Alaska. Bull. Torrey Bot. Club 105: 18-23. 17 Hulten, E. 1936. New or notable species from Alaska. Contributions to the flora of Alaska I. Svensk Bot. Tidskr. 30: 515-528. 18 Hulten, E. 1941. Flora of Alaska and Yukon, 1-10. Part 1, pp. 30. Lunds. Universitets Arsskrift N.F., Avd. 2, 37: 1-46:1 (1941-1950). 19 Hulten, E. 1960. Flora of the Aleutian Islands. 2nd Ed. J. Cramer, Weinheim/Bergstr. 376 pp. + 533 maps + 33 figs. 20 Hulten, E. 1968. Flora of Alaska and Neighboring Territories. Stanford University Press, Stanford. 1008 pp. 21 Kartesz, J.T. and Rosemarie Kartesz. 1980. A Synonymized Checklist of the Vascular Flora of the United States, Canada, and Greenland. The University of North Carolina Press, Chapel Hill. 500 pp. 22 Lellinger, D.B. 1985. A Field Manual of the Ferns & Fern-Allies of the United States and Canada. Smithsonian Institution Press, Washington, D.C. 389 pp. 23 Lipkin, R. 1985. Status Report of Polystichum aleuticum C. Chr. U.S. Fish and Wildlife Service, Office of Endangered Species, Anchorage. 21 pp. + appendix A (4 figs.), appendix B (9 figs.), appendix C (5 figs.), appendix D (3 figs.). 24 Murray, D.F. 1980. Threatened and Endangered Plants of Alaska. U.S. Department of Agriculture, Forest Service, and the U.S. Department of the Interior, Bureau of Land Management. (pp. 12-13) 59 pp. 25 Murray, D.F. and R. Lipkin. 1987. Candidate Threatened and Endangered Plants of Alaska, with Comments on other Rare Plants. University of Alaska Museum, Fairbanks, Alaska. (pp. 14-15) 76 pp. 26 Scholl, D.W., H.G. Greene, and M.S. Marlow. 1970. Eocene age of the Adak 'Paleozoic(?)' rocks, Aleutian Islands, Alaska. Geol. Soc. Amer. Bull. 81: 3583-3592. 27 Shacklette, H.T., et al. 1969. Vegetation of Amchitka Island, Aleutian Islands, Alaska. U.S. Geol. Surv. Prof. Paper 648. 66 pp. 28 Smith, D.K. 1985. 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A Contribution to the Ecology of the Plants of Atka Island, with Notes on the Flora of Adak Island, Alaska. References - 2 (DRAFT) - References Species FERN, SHIELD, ALEUTIAN Species Id ESIS857001 Date 13 MAR 96 Ph.D. Dissertation, University of Texas, Austin. 165 pp. ***** REFERENCES FOR N-OCCURRENCE NARRATIVE ONLY ***** 01 Amundsen, C.C. 1985. Central Aleutian Tundra: Ecological Manifestations of Maritime Tundra Landscapes in the Central Aleutian Islands (Amchitka, Adak) Alaska. Final Report prepared for Department of Energy, ORO-4180-10. 77 pp. + appendices I,II,III. 02 Black, R.F. 1976. Influence of holocene climatic changes on Aleut expansion into the Aleutian Islands, Alaska. Anthropological Papers, Univ. Alaska 18:31-42. 03 Christensen, C. 1938. On Polystichum aleutium C. Chr., a new North American species. Amer. Fern J. 28:111-112. 04 Friedman, B.F. 1984. (pp. 3-4 - 3-28) in Feasibility Study Data Collection Program for the Proposed Hydroelectric Project at Atka, Alaska. Northern Technical Services, Inc. and Van Gulik and Associates, Inc. Final Report prepared for Alaska Power Authority. 05 Fries. 1984. in litt., fide Lipkin 1985. 06 Hulten, E. 1936. New or notable species from Alaska. Contributions to the flora of Alaska I. Svensk Bot. Tidskr. 30: 515-528. 07 Hulten, E. 1940. History of botanical exploration in Alaska and Yukon territories from the time of discovery to 1940. Bot. Not. 289-346. 08 Hulten, E. 1941. Flora of Alaska and Yukon, 1-10. Part 1, pp. 30. Lunds. Universitets Arsskrift N.F., Avd. 2, 37L 1-46:1 (1941-1950). 09 Hulten, E. 1960. Flora of the Aleutian Islands. 2nd Ed. J. Cramer, Weinheim/Bergstr. 376 pp. + 533 maps + 33 figs. 10 Hulten, E. 1968. Flora of Alaska and Neighboring Territories. Stanford University Press, Stanford. 1008 pp. 11 Lipkin, R. 1985. Status Report of Polystichum aleuticum C. Chr. U.S. Fish and Wildlife Service, Office of Endangered Species, Anchorage. 21 pp. + appendix A (4 figs.), appendix B (9 figs.), appendix C (5 figs.), appendix D (3 figs.). 12 Murray, D.F. 1980. Threatened and Endangered Plants of Alaska. U.S. Department of Agriculture, Forest Service, and the U.S. Department of the Interior, Bureau of Land Management. (pp. 12-13) 59 pp. 13 Murray, D.F., and R. Lipkin. 1987. Candidate Threatened and Endangered Plants of Alaska, with Comments on other Rare Plants. University of Alaska Museum, Fairbanks, Alaska. (pp. 14-15) 76 pp. 14 Smith, D.K. 1985. Polystichum aleuticum from Adak Island, Alaska, a second locality for the species. Amer. Fern J. 75: 72. 15 Smith, D.K. Personal information and communication. Department of Botany, University of Tennessee, Knoxville, TN. (1987). 16 Smith, D.K. 1987. Polystichum aleuticum Chr. on Adak Island, Alaska: Status Report for 1987. U.S. Fish and Wildlife Service, Office of Endangered Species, Anchorage, Alaska. 15 pp. + appendices (7 figs.). 17 Talbot, S.S. 1986. A Search for Polystichum aleuticum (POLYPODIACEAE) on Adak Island, Alaska: Status Report Supplement. References - 3 (DRAFT) - References Species FERN, SHIELD, ALEUTIAN Species Id ESIS857001 Date 13 MAR 96 U.S. Fish and Wildlife Service, Office of Endangered Species, Anchorage, Alaska. 1-21 pp. 18 United States Geological Survey, National Mapping Division. 1982. Ecoregions and Land-Surface Forms of Alaska and Hawaii. in Ecoregion, Land-Surface Form and Hydrologic Unit Maps of the United States. Biological Services Program. Fish and Wildlife Service/OBS-82/09 (1982). 19 United States Geological Survey. 1980. 'adapted' Hydrologic Unit Map of Alaska and Hawaii. in Ecoregions, Land-Surface Form and Hydrologic Unit Maps of the United States. Biological Services Program. Fish and Wildlife Service/OBS-82/09 (1982). 20 Wagner, D. 1979. Systematics of Polystichum in western North America north of Mexico. Pteridologia 1: 1-64. 21 Welsh, S.L. 1974. Anderson's Flora of Alaska and adjacent parts of Canada. Brigham Young University Press, Provo, Utah. 724 pp. 22 Williams, M.P. 1978. in litt. U.S. Fish and Wildlife Service Files, Anchorage, Alaska. References - 4