Actaea rubifolia - (Kearney) Kartesz
Appalachian Bugbane
Other Common Names: Appalachian bugbane
Synonym(s): Cimicifuga rubifolia Kearney
Taxonomic Status: Accepted
Related ITIS Name(s): Cimicifuga rubifolia Kearney (TSN 18758)
Unique Identifier: ELEMENT_GLOBAL.2.152653
Element Code: PDRAN07060
Informal Taxonomy: Plants, Vascular - Flowering Plants - Buttercup Family
 
Kingdom Phylum Class Order Family Genus
Plantae Anthophyta Dicotyledoneae Ranunculales Ranunculaceae Actaea
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Concept Reference
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Concept Reference: Kartesz, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. 2nd edition. 2 vols. Timber Press, Portland, OR.
Concept Reference Code: B94KAR01HQUS
Name Used in Concept Reference: Cimicifuga rubifolia
Taxonomic Comments: Treated by Kartesz (1999) as Actaea rubifolia; also known as Cimicifuga rubifolia (e.g., in Kartesz (1994)).
Conservation Status
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NatureServe Status

Global Status: G3
Global Status Last Reviewed: 26Jan2001
Global Status Last Changed: 29Feb1984
Rounded Global Status: G3 - Vulnerable
Reasons: Cimicifuga rubifolia has a limited range in the U.S. in the Cumberland Plateau area of Alabama, Tennessee, Kentucky, and one county in Virginia and near the lower Ohio River valley in southeastern Illinois, southwestern Indiana, and western Kentucky. This species is generally to significantly rare throughout its range and occurs in mature forests. Although it is closely related to Cimicifuga species that are heavily traded in the medicinal plant industry, currently it does not appear that C. rubifolia is threatened by wild-collection. However, habitat loss due to logging and land-use change continues to deplete its habitat. Approximately fifty occurrences are known for the species.
Nation: United States
National Status: N3

U.S. & Canada State/Province Status
Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
United States Alabama (SH), Illinois (S2), Indiana (S1), Kentucky (S2), Pennsylvania (SNA), Tennessee (S3), Virginia (S1)

Other Statuses

Comments on USESA: This species was considered "C2" by the US Fish and Wildlife Service and has been under review for federal endangered species status listing.

NatureServe Global Conservation Status Factors

Range Extent Comments: Primary concentration of known populations in Ridge and Valley province of eastern Tennessee and southwestern Virginia; with a disjunct area of occurrence in southwestern Indiana (one county), southern Illinois (four counties), western Kentucky (four counties), and northwestern Tennessee. Also in Alabama (Kral 1983).

Number of Occurrences: 21 - 80
Number of Occurrences Comments: Approximately fifty extant occurrences: Alabama (one post-1970 record); Kentucky (four--1A, 3E; plus 1H); Tennessee (25-35--1A, 1AB, 6B, 2BC, 9C, 8E, 8U; plus 9H); Virginia (7, in 2 counties); Indiana (4, in 1 county--1A, 1B, 1C, 1E); Illinois (4 in 4 counties--1A, 3E).

Population Size Comments: This species is generally to significantly rare throughout its range. This species was considered "C2" by the US Fish and Wildlife Service and has been under review for federal endangered species status listing. C. rubifolia is included on the Illinois threatened and endangered species list for 4 counties. There are only a few occurrences known in Kentucky (pers. comm. D. White, October 2000). In January 2000, The Oak Ridge Reservation (Tennessee) listed it as threatened (Oak Ridge Experimental Laboratory Online 2000). At least three known populations have several thousand plants (ramets).

Overall Threat Impact Comments: The major threat to this shade-loving plant is the destruction of the hardwood overstory. The forest canopy can be lost to logging and other forest projects, leaving the ground unprotected (Pyne 1994, Collins 1992, Kral 1983). This in turn increases insolation and the prospect of erosion (Pyne 1994, Kral 1983). Black cohosh is not usually grazed by animals such as cattle, although rootstocks may suffer damage by trampling when such animals are present in woodlands. Despite the characteristically steep terrain in which the species occurs, grazing is a threat at some sites (Ramsey 1993b, Collins 1992). Several populations in Virginia are nearing extirpation along the tributaries of the Cumberland River. Human encroachment coupled with subsequent urbanization, development, and habitat manipulation have caused the degradation and loss of the habitat of this species (Pyne 1994, Collins 1992, Ramsey and Chester 1981). Construction of dams can lead to flooding and subsequent destruction of populations occurring on lower slopes. Some populations in Tennessee have been lost to this activity (Pyne 1994). Insect predators on C. rubifolia flowers and seeds may be a threat, although the degree to which it is a threat is not known. Seed production for a given population may be significantly impacted by insect predators.

Many species in the genus Cimicifuga are used in the herbal industry. However, C. racemosa and C. rubra are in relatively higher demand than C. rubifolia, perhaps because fewer trials have been conducted (pers. comm. R. Klein, October 2000). Therefore, threats to native populations due to collection for the medicinal use are probably minimal at this time, especially since it is relatively unknown to herbalists and wildcrafters (pers. comm. R. Klein, October 2000; pers. comm. M. McGuffin, December 2000). However, trends in the herbal products market and population dynamics should be monitored, since many congenerics are used commercially and this species is currently relatively rare. Invasive weedy species threaten this species in degraded habitats (Southern Appalachian Species Viability Project 2002).

Short-term Trend: Decline of <30% to relatively stable
Short-term Trend Comments: Populations have probably been lost in recent years to logging, erosion, impoundment following dam construction and other activities, but the extent to which this has happened is not well-documented.

Other NatureServe Conservation Status Information

Distribution
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Global Range: Primary concentration of known populations in Ridge and Valley province of eastern Tennessee and southwestern Virginia; with a disjunct area of occurrence in southwestern Indiana (one county), southern Illinois (four counties), western Kentucky (four counties), and northwestern Tennessee. Also in Alabama (Kral 1983).

U.S. States and Canadian Provinces

Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
Color legend for Distribution Map

U.S. & Canada State/Province Distribution
United States AL, IL, IN, KY, PAexotic, TN, VA

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
AL Madison (01089)*
IL Gallatin (17059), Hardin (17069), Jackson (17077)*, Johnson (17087), Massac (17127), Mchenry (17111), Pope (17151), Saline (17165)*
IN Harrison (18061), Posey (18129)
KY Caldwell (21033), Crittenden (21055), Livingston (21139), Lyon (21143)
TN Anderson (47001), Blount (47009), Claiborne (47025), Davidson (47037), Giles (47055), Grainger (47057)*, Hamblen (47063), Hancock (47067), Hawkins (47073), Jefferson (47089), Knox (47093), Loudon (47105), Meigs (47121), Monroe (47123), Montgomery (47125), Roane (47145), Stewart (47161), Sullivan (47163)
VA Scott (51169), Smyth (51173)*, Wise (51195)*
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
05 Lower Wabash (05120113)+, Lower Cumberland-Sycamore (05130202)+, Lower Cumberland (05130205)+, Red (05130206)+, Blue-Sinking (05140104)+, Lower Ohio-Bay (05140203)+, Saline (05140204)+, Tradewater (05140205)+, Lower Ohio (05140206)+
06 North Fork Holston (06010101)+*, South Fork Holston (06010102)+, Holston (06010104)+, Watts Bar Lake (06010201)+, Lower Little Tennessee (06010204)+, Upper Clinch (06010205)+, Powell (06010206)+, Lower Clinch (06010207)+, Emory (06010208)+, Middle Tennessee-Chickamauga (06020001)+, Wheeler Lake (06030002)+*, Lower Elk (06030004)+
07 Upper Fox (07120006)+, Big Muddy (07140106)+*
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Basic Description: Perennial herb to 1.5m tall; leaves sheathing, ternately compound, leaflets 3-5(7)lobed with heart-shaped bases and irregularly serrate margins; inflorescence an elongated, terminal, raceme of apetalous flowers.
General Description: Cimicifuga rubifolia is a tall, slender perennial herb. The single shoot rises from a horizontal rhizome. Stems are roundish, often maroon-tinted, unbranched and wand-like in flower. Leaves are few and concentrated toward the stem-base; leaflets are ovate to round, prominently 3-5(7) lobed and resemble red maple in size and outline; its bases are usually heart-shaped, upper surfaces usually deep green and smooth. The inflorescence is an elongated, terminal, openly cylindrical raceme of whitish flowers. The flowers are symmetrical, lack petals and have 2-5 sepals which fall off upon or shortly after the bud opens. The fruit is asymmetrically oblong, with numerous small, chaffy seeds in 2 irregular rows. (Kral 1983).
Technical Description: "Rhizomes horizontal, ca. 10 X 1.5-2 cm, knotted, bearing numerous strong fibrous roots; Stems 3-22 dm, erect, dark purplish-brown, + sulcate on the faces, smooth and glabrous or with a few lax delicate haris up to the inflorescence; Leaves ternate-biternate, the central and lateral divisions usually trifoliate but sometimes unifoliate, (thus, leaves may possess 3, 5, 7, or 9 leaflets), very rarely more than 9; Petioles 2-5 dm, angled, deeply sulcate on upper face near base, + densely pubescent with lax hairs in the groove especially toward the summit; Terminal leaflet of the central segment 9-30 X 9-25 cm, very broadly ovate, equilateral, sharply acuminate at the apex, usually deeply cordate, coarsely and irregularly dentate, deeply and acutely palmately 3-5 lobed, thin dark green above, paler beneath, glabrous above, sparsely short ciliate, sparsely to + densely pubescent with long appressed, delicate, lustrous haris along the lower veins; very veiny, 5-7 veins prominent; Other leaflets 8-24 X 6-22 cm; Inflorescence a simple panicle of 2-6 slender racemes, the terminal raceme much the longest ca. 15-30 cm; Rachis and pedicels sparsely to rather densely puberulent or short pubescent. Pedicels ca. 2 mm, slender, + thickened toward the summit, subtended by 3 bracts, 1 lance-subulate ca. 2 mm, and 2 laterally disposed, ovate-triangular, acute ca. 1 mm; Flowers white; sepals 5, fugacious, 4.5-5 mm, flattened especially toward the summit; anthers ca. .5 mm; pistils 1(--2), sessile, ca. 2 mm; style and stigma only slightly differentiated from the body of the ovary, stigma minute (_ mm), style slightly recurved, crested; Follicles 8-21 mm, sessile, strongly compressed laterally, oblong, beaked by the short blunt, hardened, apically + enlarged, ascending style which departs subapically from the ventral side, at an angle of ca. 45 degrees, pale green, walls thin, + chartaceous, veins prominent, reticulate; Seeds usually 6, the 4 middle ones in two rows, the other two solitary, ca. 3 X 1.5 mm, lenticular, cylindrical in appearance, sides oblong in outline, reddish-brown, densely covered with reddish-brown, thin, chaffy scales which are generally wider than those of C. arizonica. 2n = 16." (Ramsey 1965).
Diagnostic Characteristics: Cimicifuga rubifolia stands apart from the two other North American species, C. americanum and C. racemosa, by having 3-9 (very rarely more) leaflets, deeply cordate terminal leaflets which possess 5-9 prominent, basally arising veins, 3 bracts subtending the pedicel, oblong fruits, scaly seeds which are cylindric in outline and no staminodes (Ramsey 1987). Analyses of terminal leaflets in the three species revealed that the mean length (14.4 cm), width (15.6 cm) and sinus depth (99% falling between 1.5 and 3.0 cm) of C. rubilfolia was significantly different than C. americanum and C. rubifolia (Ramsey 1986).

According to Kearney (1897), C. rubifolia can be further separated from C. racemosa by its solitary sessile ovary, short style and partially two-ranked seeds. However, it is mostly distinguished from that species by its bibracteolate and much shorter pedicels, much fewer stamens (with filaments one-half as long and anthers one-half as large as C. racemosa), thin-walled pods (about twice as large) and chaff-covered, lenticular seeds (Kearney 1897).

Cimicifuga rubifolia can be distinguished from C. americanum by its sessile, solitary ovary, much shorter and straighter style and much shorter beak to the pod (Kearney 1897). Kearney (1897) also made the following distinctions:

The bractlets of C. rubifolia's pedicel are always basal but C. americanum's bractlets are usually borne near the middle of the pedicel. The pubescence of C. rubifolia's inflorescence is rather sparse and straight, not glandular; the pubescence of C. americanum is dense, glandular, almost granular. C. rubifolia does not have any petals, whereas C. americanum has two obovate, deeply lobed, cucullate petals. The filaments and anthers of C. rubifolia are both less than 7mm long; C. americanum's filaments and anthers are about 7 mm long.

Ecology Comments: Black cohosh flowers in the early fall from August to October, with the peak flowering period occurring about September 5-10, depending upon location throughout the range. The flowering period often lasts about 70 days (Pellmyr 1986). Fruiting occurs from late August to October (Ramsey 1993a).

Under garden conditions, individuals may obtain sufficient size to flower in their first year (Cook 1993). This, however, was not observed in the field. In most instances, plants remained small and changed little in size during their first two years (Cook 1993).

Reproduction is accomplished primarily through sexual means. Although asexual reproduction through rhizome fragmentation does occur, it is a rare phenomenon (Cook 1993). Within a given population, the number of seeds and seedlings produced peryear varies greatly due to varible environmental factors, including precipitation (Cook 1993). The production of very large numbers of seedlings (even on an occassional basis) should be sufficient to sustain the populations over time (Cook 1993).

Flowers are apetalous, consisting mainly of stamens and pistils. While no nectar is produced, there is a heavy investment in the production of pollen. Lack of nectar in these flowers makes them inferior in competition with nectar-producing plants for pollinating bumblebees (Heinrich 1975). This condition has lead some to believe that the species is primarily wind pollinated (Leppik 1964, Eames 1961), but that does not seem to be the case. Cimicifuga rubifolia is dependent upon a high density of pollinators in the vicinity and the presence of nectar producing plants intermixed within the population to attract pollinators (Pellmyr 1986).

Pellmyr (1986) examined animal pollination systems that effect this species. Four orders of insects were discovered to be visitors to C. rubifolia. Bumblebees (Hymenoptera) were the most frequent visitors and effective pollinators, trapping pollen between their legs and abdomen. Syrphid flies (Diptera) occurred with low frequency, but were also effective pollinators. Several species of small beetles (Coleoptera) and flies were observed crawling around on the flowers from stamen to stamen but rarely touched the stigmas (Pellmyr 1986). There is a large difference in height between the stigma and anthers, making bumblebees and larger Syrphid flies the best pollinators for this species (Pellmyr 1986).

Studies by Cook (1993) have found that populations or patches within populations of C. rubifolia often show a relatively high level of genetic divergence. It has been speculated that this may be an artifact of two factors, inbreeding within populations and the pooling of small breeding populations with different allelic frequencies into a single population. In the latter, pollinator flights between populations are rare as bees are often attracted more preferentially toward other plants like Impatiens pallida and Polymnia canadensis. Consequently, distribution of pollen over long distances from one individual to another is a rare event (Cook 1993). Additionally, seed dispersal is primarily through gravity, leading to the germination and growth of closely-related individuals in close proximity to each other.

Seed and flower predators reduce propagule production. The larval stage of Eupithecia cimicifugata (Lepidoptera; Geotmetridae) can strip the flowers directly off the stalk. In many experiments they were able to remove all the flowers and seed from 15-20% of a population, another 60% of all ramets in some locations suffered some loss from the larva (Pellmyr 1986).

Seeds of C. rubifolia may show the same epicotyl dormancy as described previously for C. racemosa (Cook 1993). Cook (1993) found that C. rubifolia follicles may not open until after the first frosts. Because of this exposure to cold temperatures before emergence of the radicle, dormancy is initiated. Two periods of stratification are typically required before germination can occur.

Observations of seedlings in the field led Cook (1993) to speculate that germination was enhanced in areas with little or no litter present. Future studies are required to further investigate this possibility.

The life history and demography of C. rubifolia is similar to those of other long-lived perennial species (Cook 1993). During extended periods of drought, plants may undergo extended periods of dormancy of up to at least 3 years. Also during such periods, relatively large portions of populations may be dormant.

Habitat Comments: Cimicifuga rubifolia is typically found at or near the base of north-facing slopes on talus and rocky soils derived from dolomite (Ramsey 1993d). Occupied sites are typically cool, moist and occur within mixed mesophytic forests between 270 and 480 m in elevation (although plants have been found up to 900 m) (Ramsey 1993d, Gleason and Cronquist 1991, Keener 1977). These sites frequently possess soils high in calcium and magnesium content (i.e., derived from calcareous shales, dolomite or limestone) (Ramsey 1993c, Kral 1983).

Common plant associates include Tilia heterophylla, Liriodendron tulipifera, Dirca palustris, Acer saccharum, A. rubrum, Celtis occidentalis, Liquidambar styraciflua, Impatiens capensis, Actaea pachypoda, Astilbe biternata, Lactuca canadensis, Ptilimnium capillaceum, Cimicifuga racemosa, Polygontum biflorum, Heuchera americana, Sedum ternatum, Tiarella cordifolia, Stellaria pubera, Anemone canadensis, Lindera benzoin, Sanguinaria canadensis, Phacelia bipinnatifida, Trautvetteria carolinensis, Hepatica acutiloba, Erythronium americanum and Camptosorus rhizophyllus (Ramsey 1993c). For detailed lists of associates from sites throughout its range, see Cook (1993).

The single occupied location in Alabama is a rich, damp, mixed mesophytic forest at the base of a north-facing slope (Tennessee Valley Authority Regional Heritage Program 1992).

In Illinois, C. rubifolia grows on sandstone and limestone in rather deep ravines (Ramsey 1993c). Mohlenbrock (1975) listed the habitat in Illinois for this species as "rich woods." For detailed lists of associates at two Illinois sites, see Cook (1993).

The primary habitat in Indiana is mesic upland forests (Homoya 1992).

In Kentucky, C. rubifolia habitat includes ravines, rocky places near creeks at the base of hills and piney bluffs along rivers (Kentucky Natural Heritage Program 1992). For detailed lists of associates, see Cook (1993).

Occupied habitat in Tennessee includes rich soil on river bluffs, north-facing hillsides and talus slopes, moist dolomite ledges in ravines, as well as rocky and shady woods below limestone bluffs (Pyne 1994, Tennessee Ecological Services Division 1992, Tennessee Valley Authority Regional Heritage Program 1992, Chester 1975, Small 1933). Cimicifuga rubifolia was collected near the base of caprock on a north-facing slope in Savage Gulf in 1977 (Wofford et al. 1979). Common associated canopy species include: sugar maple, beech, cucumbertree, bigleaf magnolia, white basswood, buckeye and chinquapin oak (Pyne 1994). Herb layer associates include: Aristolochia macrophylla, Chelone lyoni, Claytonia caroliniana, Disporum maculatum, Dryopteris intermedia, Mitella diphylla, Polygonatum pubescens, Trillium erectum, Phlox divaricata, Actaea spp., Delphinium tricurve, Sedum ternatum, Dentaria spp., Sanicula spp., Hepatica spp., Carex platyphylla, Cimicifuga racemosa, Caulophyllum spp., Erythronium spp., Heuchera spp. and Viola blanda (Pyne 1994, Pelmyr 1986). For additional lists of associates, see Cook (1993).

The habitat in Virginia is rocky, north-facing limestone talus slopes, rich mesic slopes, limestone bluffs, rocky woods along rivers, shaded deciduous woods and edges and open mixed hardwood forests (Tennessee Valley Authority Regional Heritage Program 1992, Virginia Division of Natural Heritage 1992b, Ramsey 1964). For a detailed list of associates at one Virginia site, see Cook (1993).

Economic Attributes
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Economically Important Genus: Y
Economic Comments: A RELATED SPECIES, CIMICIFUGA RUBRA, ANOTHER SPECIES IN THE GENUS, IS SOMETIMES USED AS A SUBSTITUTE FOR C. RACEMOSA, THE MOST POPULAR "BLACK COHOSH" ON THE MARKET.
Management Summary
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Stewardship Overview: Threats to C. rubifolia include the destruction of hardwood canopies resulting in increased isolation and erosion, long-term, high intensity grazing and habitat development and manipulation through dam and housing construction, road building, etc. Management requirements include the elimination of livestock grazing in occupied habitat and the implementation of appropriate low-impact logging activities (e.g., selective logging). Monitoring requirements include the tracking of extant populations with respect to on-going management regimes (number of individuals, flowering plants, fruiting plants, successful recruitment, habitat and threats). Research should be centered on the effects of forest management techniques on the species, its reproductive biology and ecology, structure of populations, rate of recruitment into populations and conditions required for germination. The restoration potential of the species needs to be assessed.
Restoration Potential: The restoration potential of this species is not well known. No efforts to restore the species to favorable habitats have been undertaken.
Preserve Selection & Design Considerations: Preserve designs should encompass sufficient acreage and habitat to maintain a viable population of C. rubifolia over the long-term. Preserve designs should include buffer to protect the population and its pollinators from outside influences and maintain natural regimes that sustain the habitat in which C. rubifolia occurs.
Management Requirements: Livestock grazing in occupied habitat should be controlled to prevent physical trampling of populations (Kral 1983), erosion and compaction of the soil. Long-term, intensive grazing activities may likely result in the extermination of populations.

Since C. rubifolia typically occurs on steep slopes, logging within areas surrounding the occupied habitat should be selective in order to maintain forest canopy and keep erosion at a minimum. Although thinning of the canopy directly over existing populations may have no lasting effect on the species (Kral 1983), physical disturbance from such actions may exert their toll. It may be best to eliminate logging practices from the immediate C. rubifolia population. Extensive thinning or cutting of the overstory may likely destroy C. rubifolia populations due to destruction of its shaded habitat (Kral 1983).

Monitoring Requirements: Monitoring of extant populations should be initiated (Homoya 1992) to track the status of populations with respect to on-going management regimes. Populations should be monitored on a regular basis to determine the number of individuals present, number of plants successfully flowering and fruiting and successful recruitment into the population. Habitat should be monitored, with emphasis placed on percent canopy cover. An assessment of threats and habitat quality should also be made.

Management Programs: No management programs are known to exist for C. rubifolia at this time.
Monitoring Programs: No monitoring programs are known to be in effect at this time.
Management Research Programs: Ramsey (1993d) has been studying aspects of the genus Cimicifuga for most of his professional life. His research is centered on the systematic, ecological, phytogeographical and chromosomal aspects of the genus.

Contact: Gwynn Ramsey, Department of Biology, Lynchburg College, Lynchburg, VA. Telephone: (804) 522- 8100.

Management Research Needs: The effects of various forest management techniques on this species need to be investigated to learn which are detrimental or beneficial for the species. Further study relating to threats and habitat requirements would help in the protection, management and identification of populations.
Additional topics: Additional common names include bugbane and rattletop. An illustration of the C. rubifolia can be found in Pellmyr (1986) and Ramsey (1993a, 1992).

Range distribution maps for the C. rubifolia can be found in the following sources: North America (Ramsey 1993b); United States (Ramsey 1993b, Kral 1983); Virginia (Harvill et al. 1986).
Population/Occurrence Delineation Not yet assessed
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Population/Occurrence Viability
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U.S. Invasive Species Impact Rank (I-Rank) Not yet assessed
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Authors/Contributors
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NatureServe Conservation Status Factors Edition Date: 26Jan2001
NatureServe Conservation Status Factors Author: Ostlie, W.R.; S.L. Neid (1998); rev. K. McConnell (2001)
Management Information Edition Date: 04Mar1994
Management Information Edition Author: AMBROSE, DONN M. OSTLIE, WAYNE R. PENSKAR, MICHAEL R. SCHUEN, DAVID WALTER
Management Information Acknowledgments: We are indebted to all the botanists, ecologists, information managers and others who took the time to provide the information necessary for the preparation of this and many other Element Stewardship Abstract.
Element Ecology & Life History Edition Date: 04Mar1994
Element Ecology & Life History Author(s): AMBROSE, DONN M. OSTLIE, WAYNE R. PENSKAR, MICHAEL R. SCHUEN, DAVID WALTER

Botanical data developed by NatureServe and its network of natural heritage programs (see Local Programs), The North Carolina Botanical Garden, and other contributors and cooperators (see Sources).

References
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  • Chester, E.W. 1975. Range extensions and first reports for some Tennessee vascular plants. Castanea 40:56-62.

  • Cook, R.A. 1993. The population biology and demography of Cimicifuga rubifolia Kearney and the genetic relationships among North American Cimicifuga species. Ph.D. dissertation, University of Tennessee, Knoxville. 161 pp.

  • Eames, A. 1961. Morphology of the angiosperms. McGraw-Hill, New York.

  • Flora of North America Editorial Committee. 1997. Flora of North America north of Mexico. Vol. 3. Magnoliophyta: Magnoliidae and Hamamelidae. Oxford Univ. Press, New York. xxiii + 590 pp.

  • Flora of North America Editorial Committee. 1997. Flora of North America, vol. 3 Magnoliophyta: Magnoliidae and Hamamelidae. Oxford University Press, New York.

  • Gleason, H.A. and A. Cronquist. 1991. Manual of vascular plants of northeastern United States and adjacent Canada. New York Botanical Garden, Bronx, NY. 910 pp.

  • Gleason, H.A., and A. Cronquist. 1991. Manual of vascular plants of northeastern United States and adjacent Canada. New York Botanical Garden, Bronx, New York. 910 pp.

  • Harvill, A.M., Jr., T.R. Bradley, C.E. Stevens, T.F. Wieboldt, D.M.E. Ware, and D.W. Ogle. 1986. Atlas of the Virginia flora. Second edition. Virginia Botanical Associates, Farmville. 135 pp.

  • Heinrich, B. 1975. Bee flowers: a hypothesis on flower variety and blooming times. Evolution 29: 325-334.

  • Kartesz, J.T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. 2nd edition. 2 vols. Timber Press, Portland, OR.

  • Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United States, Canada, and Greenland. First edition. In: Kartesz, J.T., and C.A. Meacham. Synthesis of the North American Flora, Version 1.0. North Carolina Botanical Garden, Chapel Hill, N.C.

  • Kearney, T.H. 1897. New or otherwise interesting plants of Eastern Tennessee. Bulletin of the Torrey Botanical Club 24:560-575.

  • Keener, C.S. 1977. Studies in the Ranunculaceae of the Southeastern United States. VI. Sida 7(1): 1-12.

  • Kral, R. 1983c. A report on some rare, threatened, or endangered forest-related vascular plants of the South. USFS Tech. Publ. R8-TP 2, Atlanta, GA. 2 Vol. 1305 pp.

  • Leppick, E.E. 1964. Floral evolution in the Ranunculaceae. Iowa State J. Sci. 39:1-101.

  • Mohlenbrock, R. H. 1975. Guide to the vascular flora of Illinois. Southern Illinois University Press, Carbondale. 494 pp.

  • Mohlenbrock, R.H. 1986. Guide to the vascular flora of Illinois. Southern Illinois University Press, Carbondale and Edwardsville, Illinois. 507 pp.

  • Pellmyr, O. 1986. The pollination ecology of two nectarless Cimicifuga sp. (Ranunculaceae) in North America. Nord.J.Bot. 6(6): 713-723.

  • Ramsey, G.W. 1964. Note on Cimicifuga rubifolia Kearney in Virginia. Rhodora 66: 135.

  • Ramsey, G.W. 1965. A biosystematic study of the genus Cimicifuga (Ranunculaceae). Abstract of Ph.D. dissertation, University of Tennessee. Dissertation Abstracts 26(12). 4pp.

  • Ramsey, G.W. 1986. A biometrical analysis of terminal leaflet characteristics of the North American CIMICIFUGA (Ranunculaceae). Virginia J. Sci. 37(1):3-8.

  • Ramsey, G.W. 1987. Morphological considerations in the North American CIMICIFUGA (Ranunculaceae). Castanea 52(2):129-141.

  • Ramsey, G.W. 1992. Historical review of CIMICIFUGA RUBIFOLIA (prepared for Flora North America Project, December 1992). 4 pp.

  • Ramsey, G.W. 1993a. Professor, Department of Biology, Lynchburg College, Lynchburg, Virginia. Prepared description, herbarium specimen photocopy and floral drawings for Cimicifuga rubifolia. 6 pp. Unpublished.

  • Ramsey, G.W. 1993b. Professor, Department of Biology, Lynchburg College, Lynchburg, Virginia. Range maps and county information for Cimicifuga rubifolia. 3 pp. Unpublished.

  • Ramsey, G.W. 1993c. Professor, Department of Biology, Lynchburg College, Lynchburg, Virginia. Ecological and phytogeographical information for Cimicifuga rubifolia. 3 pp. Unpublished.

  • Ramsey, G.W. and E.W. Chester. 1981. The occurrence of Cimicifuga rubifolia Kearney in the Interior Low Plateaus Province of Tennessee. Castanea 46: 100-101.

  • Small, J.K. 1933. Manual of the southeastern flora. Two volumes. Hafner Publishing Company, New York.

  • Southern Appalachian Species Viability Project. 2002. A partnership between the U.S. Forest Service-Region 8, Natural Heritage Programs in the Southeast, NatureServe, and independent scientists to develop and review data on 1300+ regionally and locally rare species in the Southern Appalachian and Alabama region. Database (Access 97) provided to the U.S. Forest Service by NatureServe, Durham, North Carolina.

  • Weakley, A.S. 2000. Flora of the Carolinas and Virginia: working draft of May 15, 2000. Unpublished draft, The Nature Conservancy, Southern Resource Office.

  • Wofford, B.E., T.S. Patrick, L.R. Philippe and D.H. Webb. 1979. The vascular flora of Savage Gulf, Tennessee. Sida 8(2):135-151.

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