Mimulus glabratus var. michiganensis
Current Status: Mimulus glabratus var. michiganensis (Michigan monkey-flower), listed as endangered by the Federal government and the State of Michigan, is an Upper Great Lakes endemic known only from 15 extant occurrences in northern Michigan. Twelve occurrences are currently considered viable. Two of the highest quality occurrences are on public land. Few of the remaining occurrences have protection measures in place beyond State and Federal laws. Due to the clonal growth habit and an apparent reliance on dispersal through fragmentation, the number of genetic individuals within colonies and populations is unknown and cannot be reliably estimated.
Habitat Requirements and Limiting Factors: Biological limitations include an extremely low degree of fertility, strict habitat specificity, and a poor capacity for long-range dispersal. With the exception of a single occurrence, sexual reproduction is negligible. Reproduction is almost entirely vegetative in nature and dispersal is achieved primarily through the fragmentation of clonal colonies. Direct habitat destruction has probably eliminated some colonies at both inland and Great Lakes shoreline sites. Hydrological disruptions within or near the taxon's fragile habitat, particularly those that modify water flow or result in a warming of the substrate, constitute a primary threat to this taxon's long-term survival. A lack of formal contact with private landowners unaware of colonies present on their property also constitutes a major threat to survival.
Recoveiy Objective: Reclassification to threatened, followed by delisting.
Recovery Criteria: M. glabratus var. michiganensis will be considered for reclassification from endangered to threatened status when protection is secured for all eight occurrences ranked "A" or "B" (Excellent Occurrence and Good Occurrence, respectively, according to Michigan Natural Features Inventory of The Nature Conservancy ranking criteria). M. glabratus var.
michiganensis will be considered for delisting when protection is secured for all 15 known occurrences. Protection is defined as the collective actions necessary to conserve known occurrences, maintain ecosystem process for the perpetuation of essential habitat, and enable each occurrence to be naturally self-sustaining. New colonies discovered subsequent to the approval of the Recovery Plan, and considered to be viable or restorable, must also be protected in order for delisting to be considered.
2. Determine the specific extent and status of all known and historically documented sites, and conduct inventories in potential areas to identify new occurrences.
3. Conduct biosystematic research to determine the most appropriate taxonomic classification.
4. Conduct demographic, physiological, breeding system, and genetic studies to understand population biology, specific habitat requirements, floral biology, and -enetic variability.
Estimated cost of recovery for FY 1998-2008 (in $1000's): details are found in the
Year --------Task --------1 Task------- 2 Task --------3 Task--------- 4 Total
1998--------- 45 ------------5 -----------10 -------------25 --------------85
1999--------- 44------------ 5 -----------10 -------------20-------------- 79
2000--------- 43------------ 5--------------- 5---------------- 20-------------- 73
2001 ---------10---------- --3 -------------0------------- 10-------------- 23
2002---------- 9------------ 2------------- 0--------------- 5 --------------16
2003---------- 6 ------------0 -------------0 ---------------5----------------- 11
2004 ----------6------------ 0--------------0--------------- 2 ---------------8
2005 ----------5----------------0 -------------0--------------- 2 ---------------7
2006---------- 4 -------------0------------- 0--------------- 2 ---------------6
2007---------- 2------------- 0------------- 0--------------- 2--------------- 4
2008---------- 2 -------------0------------- 0--------------- 2 ---------------4
Total 176 20 25 95 316
Date of Recovery: If recovery criteria are adequately met, reclassification to threatened can be
considered in 2001, delisting can be considered in 2004.
Recovery plans delineate reasonable actions which are believed to be required to recover and/or protect listed species. Plans published by the U.S. Fish and Wildlife Service are sometimes prepared with the assistance of recovery teams, contractors, state agencies, and others. Recovery teams serve as independent advisors to the Service. Plans are reviewed by the public and submitted to additional peer review before they are adopted by the Service. Objectives will be attained and any necessary funds made available subject to budgetary and other constraints affecting the parties involved, as well as the need to address other priorities. Recovery plans do not obligate other parties to undertake specific tasks and may not represent the views or the official positions or approval of any individuals or agencies involved in the plan formulation, other than the U.S. Fish and Wildlife Service. They represent the official position of the U.S Fish and Wildlife Service only after they have been signed by the Regional Director or Director as approved. Approved recovery plans are subject to modification as dictated by new findings, changes in species status, and the completion of recovery tasks. By approving this document, the Regional Director certifies that the data used in its development represents the best scientific and commercial data available at the time it was written. Copies of all documents reviewed in development of the plan are available in the administrative record, located at the East Lansing Field Office.
Literature Citation should read as follows:
U.S. Fish and Wildlife Service. 1997. Recovery Plan for Michigan monkey-flower (Mimulus glabratus var. michiganensis). Ft. Snelling, Minnesota. vii+ 37 pp.
Additional copies may be purchased from:
Fish and Wildlife Reference Service
5430 Grosvenor Lane, Suite I 10
Bethesda, Maryland 20184-2158
800-582-3421 or 301-492-6403
The fee varies for the Plan depending on the number of pages of the Plan.
Preparation of the Recovery Plan (Plan) was completed by Mr. Michael Penskar, Program Botanist, Michigan Natural Features Inventory of The Nature Conservancy. The Plan was enhanced considerably through the careful editing, expertise, and suggestions of the following individuals, whose assistance is gratefully acknowledged: Dr. Leni A. Wilsmann, Michigan Natural Features Inventory, Lansing; Dr. Edward G. Voss, University of Michigan Herbarium, Ann Arbor; Ms. Mary Harding, Michigan Department of Natural Resources, Wildlife Division, Lansing; Mr. Robert Jones, naturalist, Glen Arbor, MI and Ms. Elaine Chittenden, Collections Manager, Beal Gardens, Michigan State University. Special thanks must be given to Mr. William Harrison, U.S. Fish and Wildlife Service (Service) Region 3 office, who guided this recovery plan through several phases of preparation and kindly provided help in every way. Additional editing was provided by Ms. Zella Ellshof of the Service's Region 3 office. Final editing and wordprocessing on the plan were completed by Mr. Michael E. DeCapita, Mr. Mark Hodgkins and Ms. Sally Hopp of the Service, East Lansing, Michigan, Field Office. Extensive research by Dr. Robert K. Vickery, Jr., University of Utah, and investigations by Sandra Beadle and Margaret Bliss during their graduate studies at the University of Michigan have greatly enhanced our knowledge of the Mimulus glabratus complex. The cover and figure illustration of Mimulus glabratus (sensu lato) is from The New Britton and Brown Illustrated Flora of the Northeastem United States and Adjacent Canada (Volume 3, page 214, by H. A. Gleason, Copyright 1952), reprinted with permission from the New York Botanical Garden.
TABLE OF CONTENTS
TABLE OF CONTENTS
A. Background and Description
B. Taxonomy and Origin
F. Threats and Limiting Factors
G. Conservation Measures
H. Strategy of Recovery
A. Objective and Criteria
B. Step-down Outline
C. Narrative Outline for Recovery Actions
D. Literature Cited
III. IMPLEMENTATION SCHEDULE
A. Key to Implementation Schedule Column I
B. Key to Agency Designations in Columns 6 and 7
C. Implementation Schedule
LIST OF FIGURES
1. Illustration of Mimulus glabratus var. michiganensis (Pennell) Fassett (Michigan monkey-flower)
2. Plot of canonical variates based on discriminant analysis of floral measurements of three of Michigan's Mimulus taxa
3. The rangewide distribution of Mimulus glabratus var. michiganensis (Michigan monkey-flower)
LIST OF TABLES
1. The rangewide distribution and status of Mimulus glabratus var. michiganensis (Michigan monkey-flower)
2. Vascular plants and bryophytes commonly associated with Mimulus glabratus var. michiganensis (Michigan monkey-flower)
LIST OF APPENDICES
A. Mean and range values for floral measurements of Mimulus glabratus var. jamesii, M. glabratus var. michiganensis, and M. guttatus
B . Michigan Natural Features Inventory element occurrence ranking criteria
C. Principal Federal and State laws applicable to the protection of Mimulus glabratus var. michiganensis (Michigan monkey-flower)
D. Public comment and peer review
A. Background and Description
Mimulus glabratus var. michiganensis (Pennell) Fassett, (Michigan monkey-flower), a member of the Scrophulariaceae (snapdragon family), is an endemic variety of a widespread and diverse complex of yellow monkey-flowers. The taxon is known from only 15 extant occurrences and is distributed principally within Michigan's Mackinac Straits region in Charlevoix, Cheboygan, Emmet, and Mackinac Counties, with outlying localities to the south in Benzie and Leelanau Counties. It is restricted to co6l, alkaline springs and streams usually associated with Thuja Occidental (Northern white-cedar) swamps along current and post-glacial Great Lakes shorelines.
Based on a specimen collected in Cheboygan County by J. H. Ehlers in 1925, Francis W. Pennell (1935) described the taxon and named it Mimulus glabratus ssp. michiganensis. Norman C. Fassett (1939) subsequently gave the taxon varietal status. Some researchers noted morphological overlap with other taxa, particularly the more common, wide-ranging Mimulus glabratus var.jamesii (James'monkey-flower) (Crispin 1981; Bliss 1983, 1986), but results from studies of floral characters of closely related taxa support maintaining var. michiganensis as a distinct taxonomic entity (Bliss 1983, 1986; Minc 1989) (See Appendix A). Mimulus glabratus var. michiganensis was proposed for listing as an endangered species in 1989 (U.S. Dept. of Interior 1989) and listed in 1990 (U.S. Dept. of Interior 1990) following a status survey (Crispin and Penskar 1989). The taxon is also listed as endangered by the State of Michigan (MDNR 199 1). As a taxon with a moderate degree of threat, a high recovery potential, and possible conflicts with recreational activities, construction or other development projects, M. glabratus var. michiganensis has a Federal recovery priority of 9C'.
Mimulus glabratus var. michiganensis is an aquatic to semi-aquatic perennial plant characterized by its mat-forming, clonal growth habit. The stems, which range to about 40 cm (15.7 in) or more in length, are lax and reclining at their base, rooting freely at lower leaf nodes (Figure 1) to produce numerous additional shoots via stolons. Propagation in this manner often results in the production of clones of up to several hundred stems or more. The broadly ovate to roundish, opposite leaves are inconspicuously to coarsely sharp-toothed and have leafstalks that are usually shorter than the blades. Upward the leaves become somewhat reduced and shorterstalked. Bright yellow, snapdragon-like, tubular flowers are produced from the upper leaf axils, borne on slender pedicels that may be longer than the leaves. Flowering occurs primarily from approximately mid-June to August, extending occasionally into Oittober. The two-lipped flowers range from 16-27 mm (0.6-1.1 in) in length and have an irregularly red-spotted lower lip and tube. The three-lobed, heavily-bearded lower lip forms a wide landing platform for insect pollinators; the upper lip bears two upright lobes. The fruit, which is seldom produced, consists of an oblong, pointed capsule ca. 8-10 mm (0.3-0.4 in) long, containing numerous oval seeds with longitudinal striations (Grant 1924).
'Federal Priority Numbers from 1 to 18 are assigned to each species at the time of its listing. They are based upon "Endangered and Threatened Species Listing and Recovery Priority Guidelines" that were published by the U.S. Fish and Wildlife Service in 1983 (48 Federal Register 43098, September 21, 1983). These criteria deal with degree and immediacy of threat, recovery potential, taxonomic uniqueness, and conflict with development activities. The Federal Priority Number 9 denotes a moderate degree of threat, a high recovery potential, and that the species will not face extinction if recovery is temporarily held off, although there is continual population decline or threat to its habitat. The "C" designation denotes that the species is, or may be, in conflict with construction or other development,projects or other forms of economic activity.
Mimulus glabratus var. michiganensis is most likely to be confused.lwith the more common, wide-ranging Mimulus g labratus var. jamesii (James' monkey-flower, previously known [Voss 1996] as M. glabratus var. fremontii), which can often be distinguished by its usually smaller flowers that range from 8-18 mm (0.3-0.7 in) in length. Despite some overlap in flower size between these varieties (see Appendix A), the consistently different style and pistil lengths (ca. 2.8-4.6 mm (0. 1-0.2 in) and 6.3-9.3 mm (0.25-0.37 in), respectively, in M. glabratus var. jamesii versus 8.1-9.1 mm (0.32-0.35 in) and 15.4-17.8 mm (0.6-0.7 in), respectively, in M. glabratus var. michiganensis) can be used to most reliably separate them, as noted by Bliss (1983, 1986). Spots on the corolla of M. glabratus var.jamesii are usually only on the tube. Vegetative characters of the two varieties have considerable overlap and cannot be used to dependably distinguish them. The leaves of M. glabratus var. jamesii are usually smaller, have shorter stalks, and are less conspicuously toothed. Mimulus guttatus DC (common monkeyflower), a predominantly western species which is found at a single Michigan locality in the western Upper Peninsula, also has some size overlap with M. glabratus var. michiganensis. Mimulus guttatus, however, is a more erect, stouter plant with much larger flowers that may range from 16-45 mm (0.6-1.8 in) long (Minc 1989). Additional distinguishing floral characters include larger, better developed calyx lobes, a wider corolla, and a more strongly spotted throat and floral tube.
B. Taxonomy and Origin
Mimulus, a large, highly variable (i.e., polymorphic) genus of as many as 150 species (Pennell 195 1), was placed in the section Simiolus (within the tribe Gratioleae) by Grant (I 924), who considered Mimulus glabratus and its varieties to be the most widely distributed group in the genus. Mimulus glabratus is found from Quebec to Saskatchewan, ranging south through Mexico and southern Chile, and seven varieties of the species have been named (Grant 1924; Pennell 1935; Fassett 1939; Skottsberg 1953). The varieties represent diploid as well as several polyploid races, which are strongly correlated with well documented geographical, morphological, and other differences, such as allozyme patterns and cytogenetic characteristics (Vickery 1990).
Pennell (1935) recognized M. glabratus var. michiganensis as a subspecies of M. glabratus based on its larger, "unspotted corollas", the more conspicuously toothed, sinuatedentate leaves, and a more erect growth habit. The type specimen was collected from the banks of Niger Creek "near Topinabee," Cheboygan County, in July 1925 (J. H. Ehlers 3240, MICH). Specimens correctly identified as M. glabratus var. michiganensis were also previously collected near Harbor Springs, Emmet County (C. F. Wheeler July 12, 1890, MSC; M. Irwin, July 1892, MSC).
Fassett (1939), who gave M. glabratus var. michiganensis varietal status, essentially a-reed with Pennell's description, though he correctly noted in the field that the corolla tube and throat were usually spotted'. Some researchers have subsequently questioned the validity of recognizing M. glabratus var. michiganensis as a distinct taxon, based on observations of what appeared to be a continuum of morphological diversity with the more common M. glabratus var. jamesii (Crispin 1981; Crispin and Penskar 1989; Bliss 1983, 1986). In one study, Bliss (198 examined the morphology, fertility, and chromosomes of M. glabratus var. jamesii and M. glabratus var. michiganensis to determine their genetic and morphological distinctiveness. Because of significant morphological differences in their flowers and differences in the percentage of viable pollen produced, Bliss determined that the varieties are genetically different and thus are valid taxonomic entities. Although the origin of M. glabratus var. michiganensis could not be deduced from the study, Bliss formulated three plausible hypotheses:
2) Mimulus glabratus var. michiganensis may have originated as the result of hybridization between M. glabratus var. jamesii and M. guttatus (n = 14), the latter a western species discovered in Upper Michigan since Bliss' study. The morphological intermediacy of M. glabratus var. michiganensis strongly suggests this possible relationship.
3) Mimulus glabratus var. michiganensis may have originated as an aneuploid' of M. guttatus, a species considered by Grant (1924) to be the most variable in the genus. Vickery et al. (1968) found aneuploid pollen in M. guttatus, and Vickery (1959) notes that populations with reduced seed set, a characteristic of W. glabratus var.
michiganensis, commonly produce aneuploid pollen.
Bliss (1986) stated that it would not be possible to determine the origin of M. glabratus var. michiganensis without a comparative analysis with M. guttatus. The discovery of a presumed native occurrence of M. guttatus in the western Upper Peninsula, a region where other western disjuncts are well known, provided an opportunity to initiate such an investigation. Michigan Natural Features Inventory ( conducted a morphological study that compared M. guttatus with M. glabratus var. jamesii and M. glabratus var. michiganensis by using the only available Michigan specimens of M. guttatiis augmented with additional live specimens of M. guttatus from Utah. Minc (1989) analyzed floral' morphological data of four sample populations of M. glabratus var. jamesii, four populations of M. glabratus var. michiganensis, and two populations of M. guttatus (see Appendix A for a summary of these data). When used in a canonical discriminant analysis, the data revealed a strong separation of the three taxa (Figure 2), indicating the substantial differences in floral morphology between these taxa. Although most of the mean values for floral characters in M. gut'tatus differed significantly from those of M. glabratus var. michiganensis, the range of variation for most floral characters overlapped with
2As Bliss (1983) observed during her research, corolla spots tend to fade on herbarium specimens, which likely led to Pennell's incorrect observation.
' Aneuploid refers to a chromosome count that is higher or lower by one or more chromosomes, as compared to the normal haploid number; the extra or lost chromosomes occur as a result of cytological irregularities.
' Because Bliss (1983, 1986) found floral characteristics to be more diagnostic and less variable than vegetative ones, only floral characters were analyzed in this study.
Vickery (1 99 1) provided evidence which documents the genetic isolation and morphological distinctiveness of M. glabratus var. michiganensis. In reciprocal crossing experiments, plants from Michigan's Maple River site, the only population known to regularly produce seed, were hybridized with plants representative of other diploid M. glabratus flower varieties known to be at least partially interfertile. Because of the failure to produce F, hybrids among 10 interpopulation crosses, Vickery concluded that M. glabratus var. michiganensis is genetically as well as morphologically distinct, and should be promoted to full species status, pending corroboration of these results with other M. glabratus var. michiganensis populations.
These studies confirm that M. glabratus var. michiganensis is a legitimate taxonomic entity, possibly worthy of recognition as a distinct species. This research also demonstrates that further investigation is required to elucidate the taxon's origin and systematic relationships. Biosystematic research involving isozyme analyses and possibly chloroplast or ribosomal DNA studies, in conjunction with natural history studies, may provide the information needed.
Mimulus glabratus var. michiganensis is known only from 15 extant occurrences in northern Michigan, ranging from Benzie and Leelanau Counties in northwestern Lower Michigan to Mackinac County in the eastern Upper Peninsula (Figure 3). Two additional occurrences are known from historical records. The ma ority of occurrences are clustered within the Mackinac Straits region. Because of the particular difficulty in ascertaining, in meaningful biological terms, precisely what constitutes a "population", MNFI's definition of "occurrences" for M. glabratus var. michiganensis will be used. An occurrence consist of colonies of plants in contiguous to semi-contiguous habitat within arbitrarily defined areas; several occurrences could thus appropriately be considered what are termed "metapopulations"'.
Of the three best occurrences of M. glabratus var. michiganensis (Table 1), McFarlane Woods is largely contained within Sleeping Bear National Lakeshore, and Carp Creek (essentially the western region of Reeses Swamp) is contained within University of Michigan Biological Station (UMBS) property, with the latter occurrence located within designated Biosphere Reserve lands. Colonies within the locality named Reeses Swamp (essentially the eastern region of this extensive cedar swamp) lie primarily on private lands, with some colonies occurring on adjoining UMBS property (See Appendix B for an explanation of MNFI occurrence ranking criteria). A portion of a significant M. glabratus var. michiganensis occurrence at Epoufette Bay is protected within a Michigan Nature Association (MNA) preserve. The remainder, and perhaps the majority of the population, occurs on township and private land.
A metapopulation can be defined as a patchwork of interacting populations (i.e. sub-populations),pver a wide and heterogeneous area of landscape (Sandland et al. 1992).
One private landowner has reportedly destroyed plants on his property despite notification of their Federal and State protected status.
The Harbor Springs site had only been known from an 1892 historical record until it was rediscovered in 1997. This site occurs within the private ownership of the Idylwllde Association, a lakeshore homeowners association. The Association is protecting wetlands on the site with a Michigan Landowner Forest Stewardship Plan (Fuller 1994).
Two historical occurrences, Burt Lake-West Shore and Mullett Lake-West Shore, if rediscovered, would almost certainly exist on private land. A new occurrence along the southeastern shore of Burt Lake was discovered in 1995. It is compfised of several local colonies inhabiting springs on the margin of the lake. A portion of this occurrence is reported to be present within the Seven Springs Nature Preserve, which is owned and managed by the Little Traverse Conservancy (LTC). At least two distinct colonies persist within the Manitou Payment Highbanks site on property of the Sand Products Corporation. Survey of previously unexplored shoreline to the east, with its extensive potential habitat, is very likely to result in the identification of additional colonies. A significant new locality, confirmed in 1990 near the Cut River mouth in Mackinac County, appears to occur on state forest land; scanty population information on this occurrence will necessitate further inventory.
The remaining M. glabratus var. michiganensis occurrences lie on private land, several involving multiple ownerships. The Maple River site and portions of the Platte River occurrence are registered with the Michigan Chapter of The Nature Conservancy (TNC), conveying at least informal protection. An occurrence in Little Sand Bay on Beaver Island occurs at least partially within LTC's Little Sand Bay Nature Preserve. The existence of the Beaver Island Harbor occurrence may be in doubt based on a recent field check (J. Paskus, MNFI, pers. comm.). A second Beaver Island occurrence was persisting at the last report. Although the colony remains highly vulnerable to Lakeshore development and potential hydrological disruptions, no protection arrangements have been made. Newly discovered colonies within a condominium development in the City of Brevort were protected during construction. Detailed plans, prepared in consultation with the U.S. Fish and Wildlife Service and the MDNR Endangered Species Program, were implemented to perpetuate this colony and its vulnerable habitat. Protection plans for the Brevort colony have included fencing around the colony, placement of a viewing boardwalk, and an information kiosk describing this species and its status. Additional protective measures were also implemented to prevent hydrological disturbances and erosion during and following condominium construction.
The status of colonies in other sites is more tenuous. In the Burdickville locality, some colonies are being informally protected through the notification of landowners by a local environmentalist. The occurrence consists of a string of M. glabratus var. michiganensis patches persisting in stepy springs along the largely developed eastern shore of Glen Lake. At Parrott Point on Mullett Lake, an extremely small, very localized colony persists in a small spring on a homeowner's lawn, despite repeated mowing.
Mimulus glabratus var. michiganensis is restricted to cold, alkaline spring seepages and streams, usually in association with northern white cedar (Thuja Occidental) swamps occurring along current or post-glacial Great Lakes shorelines. It frequently occurs in northern white cedar swamps formed in drainages found at the base of relatively steep, morainic slopes and bluffs. In these sites, M. glabratus var. michiganensis generally flourishes best in tree canopy openings, along forest edges, or along streams adjacent to open, meadow-like areas. It flowers most abundantly when growing in full sunlight, although it appears to persist as mostly sterile colonies when growing under heavy tree canopy cover.
Northern white cedar is an important and usually dominant overstory tree, conunonly occurring with Abies balsamea (balsam fir), Betula papyrifera (paper birch), and occasionally Larix laricina (larch or tamarack). In some sites, such as along the Mackinac County shoreline, M. glabratus var. michiganensi's occurs in seeps and ravines bordered by upland hardwood forest, dominated by species such as paper birch, Acer rubrum (red maple), Acer saccharum (sugar maple), and Fagus grandifolia (beech). Caltha palustris (marSh-marigold), Impatiens capensis Oewelweed), and Nasturtium officinale (watercress) are nearly constant herbaceous associates. Myosotis scorpioides (forget-me-not), although non-native, is often present. A list of vascular plants and bryophytes commonly associated with M. glabratus var. michiganensis is presented in Table 2.
Beadle (1990), in an extensive rangewide survey, and Bliss (1982), quantified several of the parameters characterizing M. glabratus var. michiganensis habitat, elucidating several of the specific abiotic requirements for establishment and growth. Beadle found that colonies nearly always occurred in muck-covered sand (Tawas soil series) in cold, flowing water that ranged in temperature from 8.7 0 to 16.6 0 C (47.6 0 - 61.9 0 F). Beadle also found that M. glabratus var. michiganensis occurs in an exceptionally narrow pH range, which over numerous study sites ranged only from 7.66 to 8.2 1. This range is very comparable to that indicated for M. glabratus var. michiganensis habitat on Glen Lake, where pH has been reported as ranging from 7.8 to 8.4 (Jones 1991). Conductivity readings, indicative of nutrient content, were predictably high, ranging from 190 gmhos at Brevort to well over 300 pmhos at most other sites. Selected water sample tests indicated high concentrations of ammonium (NH3 ), nitrate (NO3 ), and phosphorus(P').
At one site, Beadle also conducted field observations with a portable infrared gas analyzer to determine photosynthesis and transpiration rates, an experiment that resulted in apparently significant findings. Beadle compared the photosynthesis and transpiration rates of M. glabratus var. michiganensis plants growing in shallow water versus more terrestrial plants located on an adjacent stream bank. She found that the partly immersed plants had approximately twice the photosynthesis and transpiration rates of non-immersed plants. Beadle noted that the higher recorded photosynthesis rate was similar to the middle of the range of values recorded for crop plants exhibiting C3 photosynthesis. Oxygenation, though not measured by Beadle, was also suggested as an abiotic parameter that may be significant. These data collectively suggest that optimal habitat conditions for M. glabratus var. michiganensis are comprised of a combination of moderate to high light availability, cool substrate, and high nutrient availability within a narrow pH range, demonstrating the relatively specialized habitat requirements.
Habitat requirements, though not definitively known, clearly indicate the relatively restrictive conditions for M. glabratus var. michiganensis establishment and growth, as outlined above. The localized nature of available habitat is thus a strong limiting factor within the currently known range of this monkey-flower. Other significant biological characteristics also serve to severely limit the growth, dispersal, and establishment of M. glabratus var.
Relatively little is known of the population biology of M. glabratus var. michiganensis owing, in part, to a lack of long-term monitoring of this taxon. However, Bliss'study (1983, 1986) of reproductive biology provides information on significant life history characteristics. Bliss found that for 1 1 populations o M. glabratus var. michiganensis the proportion of viable pollen was only 2.8 percent (± 2.6 percent), and that there was considerable variation among sample sites. Only a single population, the Maple River site, produced an appreciable amount of viable pollen, averaging 29 percent (± 5 percent), whereas all other sites averaged just 0. 18 percent (± 0.09 percent). Bliss'data for seed set paralleled the pollen data: all shoots at the Maple River site had some seed set, in contrast to an average seed set of only 3.4 percent for all other populations. Bliss concluded that M. glabratus var. michiganensis is near y totally dependent on vegetative propagation. Because of its dependence on this form of reproduction, dispersal is greatly limited and likely occurs only locally through the fragmentation of clonal colonies.
Even when Bliss attempted to cross different populations (occurrences) of M. glabratus var. michiganensis, no seed set was achieved. The reason for M. glabratus var. michiganensis's low production of viable pollen and consequent sterility is thus unknown, but it may be due to cytological abnormalities that typically result in interpopulational crossing barriers within the Mimulus glabratus complex (Vickery 1969; Tai and Vickery 1970). The fertility of the Maple River occurrence is therefore particularly notable. Beadle (1990) has collected and successfully germinated seed collected from this site, as has Vickery (1991). The Maple River site thus may be the only occurrence with significant long-range seed dispersal potential, possibly via water or animal vectors.
Neither Bliss (1983) nor Beadle (1990) observed pollinators, whose role is thus far unknown. Pennell (1935) described nectar-producing Mimulus flowers as being highly adapted to bee pollination, owing to their broad, bearded palates (i.e. the lower petals) that serve as suitable landing platforms. Vickery (199 1) noted that M. glabratus plants are characteristically self-compatible and readily self-pollinate, producing seed frequently in this fashion.
Little population monitoring of M. glabratus var. michiganensis colonies has been conducted, and thus the demography of populations cannot be characterized. S. Beadle (1990, pers. comm.), however, has observed colonies of the taxon through summer and winter seasons, reporting that in the fall colonies die back and become more or less dormant in streams and springs where water flow and temperature stay relatively constant, with colonies re-initiating growth in the spring.
The greatest threat to M. glabratus var. michiganensis is direct destruction and modification of the species'essential habitat. Development, both inland and along Great Lakes shores, has probably resulted in local extinctions such as at Mullet Lake and portions of Burt Lake. Apparently, potential habitat remains near these historical locations, but they lack the northern white cedar swamps that were cleared and drained for residential areas and seasonal homes. However, collection information is vague for several of these historical occurrences. The 1997 Harbor Springs rediscovery indicates that extant colonies may still remain at or near their original collection sites. Because much of the remaining potential habitat is relatively inaccessible, persistent exploration is necessary before extinction can be presumed for colonies from these localities.
Other than direct habitat destruction, hydrological disruptions of any type, particularly .those that result in eliminating or warming a water source, unquestionably constitute the next most serious threat to the M. glabratus var. michiganensis. Although the occurrences of M. glabratus var. michiganensis colonies in sites such as Parrott Point and Burdickville demonstrate that plants can persist where groundwater flow has been somewhat altered, this may represent far less than optimal habitat conditions. On Burt Lake, Mullett Lake, and at Burdickville prior to development, M. glabratus var. michiganensis probably occurred much more abundantly.
Perhaps one of the strongest current threats to known occurrences of M. glabratus var. michiganensis is the lack of knowledge of their existence and significance. Because the majority of occurrences lie within private landholdings, colonies are likely to be threatened by those who do not know that M. glabratus var. michiganensis occurs on their property or what this species requires to survive. Several local occurrences are threatened because the landowners have not been formally contacted about the presence of a protected plant on their property. Despite the need for increased enforcement, which in one case was necessary to achieve future protection, notification and education are ultimately the best tools and perhaps the only hope for preventing the steady attrition of significant local colonies.
Lastly, an increased interest in this taxon by the scientific community has the potential to result in overcollection and possible contamination of colonies during experimental manipulations. At present, this kind of activity is not a significant threat, but it may become so in the future and should be carefully monitored and regulated. Over the past few years, collectors and institutions, such as arboreta, have become highly interested in propagating and displaying live specimens of endangered species. This may also place additional collection pressure on M. glabratus var. michiganensis colonies and should likewise be closely monitored and regulated to determine impacts.
Principal Federal and State laws directly relevant to the protection of M. glabratus var. michiganensis and its habitat are listed in Appendix C.
G. Conservation Measures
Conservation measures include scientific studies, legal protection, and activities that affect the conservation of the species.
The most recent study on M. glabratus var. michiganensis was conducted by Beadle (1990) on ecological and physiological requirements affecting its distribution.
Conservation measures provided to M. glabratus var. michiganensis as a listed endangered species under the Federal Endangered Species Act of 1973, as amended (Act), include recognition, recovery actions, requirements for Federal protection, and prohibitions against certain practices. Recognition through listing encourages and results in conservation actions by Federal, State, and private agencies, groups, and individuals. The Act provides for possible voluntary land acquisition and cooperation with the States and requires that recovery actions be carried out for all listed species. The protection required of Federal agencies and the prohibition against certain activities involving listed plants are discussed, in part, below.
Section 9(a)(2) of the Act and its implementing regulations found at 50 CFR 17 1 set forth prohibitions and exceptions that apply to endangered plants. Prohibited activities applicable to any person subject to the jurisdiction of the United States are as follows: 1) import or export; 2) transport in interstate or foreign commerce in the course of a commercial activity; 3) sell or offer for sale in interstate or foreign conunerce; or 4) to remove and reduce to possession from areas under Federal jurisdiction; 5) maliciously damage or destroy this species on any area under Federal jurisdiction; or 6) remove, cut, dig up, damage, or destroy this species on any other area in knowing violation of any State law or regulation or in the course of any violation of a State criminal trespass law. "Plant" means any member of the plant kingdom, including seeds, roots, and other parts. Certain exceptions apply to agents of the U.S. Fish and Wildlife Service (Service), other Federal land management agencies, and State conservation agencies [50 CFR 17.61(c)(2)]. In addition, State conservation agencies which are party to a Cooperative Agreement with the Service may conduct certain prohibitied activities with endangered plants for conservation purposes [50 CFR 17.61(c)(4)].
Section 10 of the Act and its implementing regulations provide for the issuance of permits to carry out otherwise prohibited activities under certain circumstances. Permits are available for scientific purposes or to enhance the propagation or survival of the species in the wild. In some instances, permits may be issued to relieve undue economic hardship that would be suffered if such relief were not available. It is anticipated that few trade permits would ever be sought or issued, because this species is not common in cultivation or in the wild. A section 10 permit was granted in 1995 to the Hiawatha National Forest of the U.S. Forest Service for the collection of voucher specimens necessary for a forest-wide survey. In addition, activities carried out by a state conservation agency may include many recovery research projects that are identified in this plan, in accordance with authorities described above [(50 CFR 17.61(c)(4)]. Requests for copies of the regulations on endangered plants and inquiries regarding them may be addressed to the U.S. Fish and Wildlife Service, Ecological Services Operations, 1 Federal Drive, Fort Snelling, Minnesota 55111-4056. Copies of Federal laws and regulations are also available electronically at http://www.thomas.loc and related sites.
Section 7 of the Act requires Federal agencies to consult with the Service prior to authorizing, funding, or carrying out activities that may affect listed species. Section 7 also requires that these agencies use their authorities to further the conservation of listed species. There has not been any formal Section 7 consultation for M. glabratus var. michiganensis. The National Park Service, Army Corps of Engineers, and the U.S. Forest Service have conducted informal consultations for this species.
Section 6 of the Act provides for Cooperative Agreements between the Service and state agencies for conservation of listed species. Currentlv, the Service has partially funded a landowner contact program since 1992 as well as status surveys in 1994 and 1996 through a Federal grant program administered under Section 6(d) of the Act. These activities have been carried out by the Wildlife Division of the Michigan Department of Natural Resources.
Rules for protection of listed plants in the National Forests are in the U.S. Forest Service Manual Title 2600--Wildlife, Fish, and Sensitive Plant Habitat Management, Chapter 2670-Threatened, Endangered and Sensitive Plants and Animals. The U.S. Forest Service must abide by the Endangered Species Act and the National Environmental Protection Act in managing national forests. Horseshoe Bay Wilderness Area within Hiawatha National Forest was established by the Wilderness Act and yields an untouched, protected area. The National Forest Management Act of 1976 mandates that a Management Plan be written for each National Forest. The Hiawatha National Forest Land and Resource Management Plan (U.S.Dept.of Agriculture 1986) includes management actions to be performed for endangered, threatened, and special concern plants, including M. glabratus var. michiganensis.
3. State Protections
Mimulus glabraius var. michiganensis was listed as a threatened species by the Michigan Department of Natural Resources in 1976. It was listed pursuant to Michigan's Endangered Species Act (Public Act 203 of 1974), now Part 365, Endangered Species Protection, of the Natural Resources and Environmental Protection Act (NREPA), PA 451 of 1994. The plant was elevated to endangered species status in 1991 pursuant to Federal listing as endangered. Part 365 of NREPA prohibits the taking of endangered species on both public and private lands without a permit. Part 303 of NREPA also provides for the preservation, management, protection, and use of certain wetland habitats. The law lists habitat for threatened and endangered wildlife species as a criteria to be considered in the administration of the Public Act.
In May, 1994, seven Federal agencies, including the U.S. Fish and Wildlife Service, U.S. Forest Service and National Park Service signed the Federa Native P ant Memorandum of Understanding to establish the Federal Native Plant Conservation Conu-nittee. The committee is to identify conservation needs for native plants and coordinate implementation of programs to address those needs. Since the original signing by seven agencies, additional agencies have joined the agreement.
In September 1994, 18 Federal agencies signed a Memorandum of Understanding (MOU) on the Implementation of the Endangered Species Act. In this MOU, the cooperators agreed to work together to (1) conserve Federally listed species; (2) use existing Federal authorities and programs to further the purposes of the Act; and (3) to improve the efficiency and effectiveness of consultation under Section 7 of the Act. Conservation Agreements are specifically mentioned as a tool to implement this MOU.
Recovery of M. glabratus var. michiganensis is based on three fundamental components: (1) the protection of the relatively few known extant occurrences, (2) field surveys to better determine the status of this taxon, and (3) biosystematic research, including long-term monitoring, to ascertain the most appropriate taxonomic placement of M. glabratus var. michiganensis, and to understand its basic natural history so that it may be properly managed.
There are very few known occurrences of the M. glabratus var. michiganensis and its biology is poorly understood. Although additional significant occurrences can be expected, intensive site inventories have thus far resulted in the identification of few new occurrences Therefore, to insure its survival, the immediate recovery priority is to protect all extant "populations" and their immediate habitat. One large, exemplary occurrence is predominantly contained and protected within a National Lakeshore. Another occurrence lies within a University of Michigan biosphere reserve. Portions of three additional occurrences lie within small nature preserves. The remainder, which are the majority of the extant occurrences, occur primarily on private land where they remain highly vulnerable to both direct and indirect impacts. The conservation of most known occurrences of M. glabratus var. michiganensis must occur through landowner notification to implement voluntary protection programs. Voluntary programs may include landowner contact and education, non-binding registry agreements with private or governmental agencies, and formal, recorded conservation easements. Fee title purchase from willing sellers and ownership by a private conservation organization, or a state or federal agency, is an option for the protection of some sites.
Artificial propagation and reintroductions of the M. Glabratus var. michiganensis are not currently being considered. Prior to consideration of these actions, more productive tasks such as protection of extant occurrences and searches for new occurrences should be exhausted. If new information indicates a pressing need for reintroduction, it should only be considered when there is a complete understanding of the original extirpation, original threats have been eliminated, adequate protection of the sites is guaranteed, and a monitoring program is in place.
The fundamental recovery objective for M. glabratus var. michiganensis is to secure long-term protection for all of its 15 known occurrences as well as viable or restorable occurrences discovered subsequent to the preparation of this recovery plan, or newly identified extant colonies in historical sites. Viable sites contain the number of individuals and area of essential habitat necessary for a self-sustaining population. Protection is defined as the collective actions necessary to conserve known occurrences, maintain ecosystem processes for the perpetuation of essential habitat, and enable each occurrence to be naturally self-sustaining. M. glabratus var. michiganensis will be considered for reclassification from endangered to threatened status when protection is secured for all eight occurrences ranked "A" or "B" ("Excellent Occurrence" and "Good Occurrence", respectively, see Appendix B). When all known Michigan occurrences are sufficiently protected, delisting can be considered.
The goals of the recovery plan can be accomplished by (1) long-term protection for all known existing occurrences, with primary emphasis on the preservation of essential habitat, (2) field surveys for new occurrences and to determine the specific status of recently discovered and historical sites, (3) biosystematic research to determine the most appropriate taxonomic classification, and (4) demographic, physiological, breeding system, and genetic studies to understand population biology, specific habitat requirements, floral biology, and genetic variability, and long-term post delisting viability monitoring.
12. Education of land managers and the public.
122. Interpretation of scientific research and communication of relevant findings to land managers and the public.
123. Preparation and dissemination of specific guidelines for the management of M. glabratus var. michiganensis occurrences.
13. Development of management plans for occurrences on Federal, State, and local public lands.
14. Registry agreements, conservation easements, and other contractual arrangements for site and colony protection.
15. Land purchase.
16. Implementation of protection strategies for all identified sites.
16-4. Burt Lake-West Shore (historical occurrence).
16-5. Burt Lake Southeast
16-6. Carp Creek.
16-7. Cut River West.
16-8. Epoufette Bay.
16-9. Harbor Springs.
16-10. Little Sand Bay.
16-11. Manitou Payment Highbanks.
16-12. Maple River.
16-14. Mullett Lake-West (historical occurrence).
16-15. Parrot Point (Mullett Lake-Southeast).
16-16. Platte River.
16-17. Reeses Swamp.
2. Conduct field surveys for new occurrences and to determine the specific status of recently discovered and historical sites.
22. Known extant sites.
23. New occurrences.
3. Conduct comparative biosystematic research of M. glabratus and M. guttatus to determine the most appropriate taxonomic classification.
4. Conduct demographic, physiological, breeding system, and genetic studies to monitor and understand population biology, specific habitat requirements, floral biology, and genetic variability.
42. Physiology studies.
422. Greenhouse research.
43. Breeding system biology.
44. Genetic research.
45. Long-term monitoring.
C. Narrative Outline for Recovery Actions
The perpetuation of M. glabratus var. michiganensis depends on achieving long-term protection for its specialized habitat so that the few known occurrences can be selfsustaining. This can be best attained through the following recovery actions.
Landowner notification or contact consists, at a minimum, of communicating with all landowners, both public and private, on whose property M. glabratus var. michiganensis occurs. Landowner notification is currently carried out by MNFI with the eventual goal of turning over the program to local land trusts. Initial contact is made by mail and/or telephone with follow up contact made in person. Notification consists of the confin-nation that a Federal and State endangered species occurs within an ownership. This notification must include prepared educational materials that clearly describe and illustrate the M. glabratus var. michiganensis in non-technical terms, providing information on the rarity of the species, why it is rare, what it requires to be self-sustaining, and why it should be protected. Integral to the notification process is the necessity of apprising landowners of the legal protection afforded M. glabratus var. michiganensis under provisions of both the Federal and State Endangered Species Acts. Notification should entail contacting adjacent landowners whose property provides contiguous and potential habitat. Notification may be a continuing process, requiring the transmittal of new information relevant to protection as provided by scientific research.
12. Education of land managers and the public - Both short and long-term protection and management of M. glabratus var. michiganensis are dependent upon the dissemination of understandable information to land managers and the public, especially those who require non-technical information on important elements of Great Lakes biological diversity. Education can be accomplished in the following ways.
122. Intelpretation of scientific research and communication of relevant findings to land managers and the public - As research provides important information, these findings require interpretation and transfer to all distributed materials, and particularly to public land managers and private owners directly responsible for maintaining M. glabratus var. michiganensis colonies and their habitat.
123. Preparation and dissemination of specific guidelines for the management of M. -labratus var. michiqanenst's occurrences - In addition to notification and general educational materials concerning M. glabratus var. michiganensis, most public agencies and private land owners will require specific instructions and guidelines for preparing management plans.
13. Development of management plans for occurrences on Federal, State, and local public lands - The protection of occurrences on public lands, including Federal, State, and local (county, township, or city) holdings, should be guided by specific management plans. Clear management plans are particularly important for public agencies and governmental units, owing to the expected changes in personnel and the need for familiarity and consistency in management policies and actions. For local government units, management plans may consist of the enactment of zoning provisions or other plans that allow for the perpetuation of M. glabratus var. michiganensis habitat and occurrences. Sites that are at least partially within public ownership, and for which management plans are appropriate, include McFarlane Woods (National Park Service), Carp Creek (UM Biological Station), Cut River West (State Forest) and Epoufette Bay (city and/or township).
14. Regist[y agreements, conservation easements, and other contractual arrangements for site and colony protection - Public and private registry agreements, consisting of legal, non-binding contracts, are proactive alternatives that can provide short-term protection for occurrences. Because M. glabratus var. michiganensis occurrences often lie within several private ownerships, registry is a desirable approach that provides short-term protection. It may ultimately lead to long-term protection through eventual donation, purchase, natural area designation, or other means. The registry program of TNC, complementing the landowner contact/registry program of the Michigan Natural Heritage Program (Wildlife Division, Michigan DNR), should place a high priority on protecting M. glabratus var. michiganensis. Conservation easements, pursuant to provisions of Part 21, Conservation and Historic Preservation Easement, NREPA, or other formal, legally-binding contracts, also provide a viable approach to protection.
15. Land purchase - The above recovery actions are all based primarily on the perpetuation of essential habitat of the M. glabratus var. michiganensis. Land purchases from voluntary sellers are another definitive method of protecting the plant's essential habitat. Two programs in Michigan have good potential for incorporating critical sites under state ownership. Purchases under the Michigan Natural Resources Trust Fund are nominated by citizens or government officials and administered by the state or local government organizations. Part 21, Land Exchange Facilitation Fund, of NREPA allows the MDNR to sell state land and use the proceeds to purchase other land. Land purchase by TNC, the LTC, or other statewide and local land conservation organizations, are also recommended approaches.
16. Implementation of protection strategies for all identified sites - Recovery actions are recommended for the following sites with extant and historical occurrences.
16-14. Mullett Lake-West (historical occurrence) - A specimen was collected here in 1925 and has been tracked as a historical site. A recent search (Beadle 1990) identified potential habitat but no extant populations. The area has been poorly explored due to access difficulty. Continued systematic inventory is needed. This is a site for potential reintroductions if it is first carefully surveyed and evaluated.
16-15. Parrot Point (Mullett Lake-Southeast) - Reduced to an extremely localized colony on the lawn of a lakeshore resident, this clump persists, apparently in spite of mowing. This occurrence should be given high priority for immediate landowner contact and possible registry. Any potential habitat in the general area strongly merits inventory.
16-16. Platte River - Survevs in 1990 found small colonies more widely ranging than known previously and many occurring on private property. A brief downstream survey conducted at the same time did not result in the identification of additional colonies, although this inventory was conducted over less than a 400 in (0.25 mi) stretch. Detailed surveys of the general area are necessary, including potential areas within the Platte River drainage near and beyond this branch. Two owners are currently registered with the Michigan Chapter of TNC. Additional registry, as appropriate, should be sought.
16-17. Reeses Swamp - While only a relatively small portion of this occurrence exists on University of Michigan Biosphere Reserve land, the majority of the population occurs in multiple private ownerships. TNC registry is strongly recommended, and possibly the pursuit of conservation easements, to ensure long term habitat protection.
2. Conduct field surveys for new occurrences, status determinations for recently discovered sites, and historical sites.
Relatively few M. glabratus var. michiganetisis sites are known, and of these, several are disturbed. However, significant potential habitat, which merits surveys, exists both within and beyond its current delineated range. Although recent field investigations have resulted in few new sites, they are highly important discoveries (e.g. McFarlane Woods) and have given direction to future surveys. Continued inventory is strongly recommended, as are detailed assessments of known sites and historical occurrences.
22. Known extant sites - Detailed mapping of colonies in several known extant occurrences is still needed to determine the complete local distribution and status. Mapping could result in the potential revision of element occurrence ranks.
23. New occurrences - Conducting inventories in northern Lower Michigan and the eastern Upper Peninsula, focusing on current and post-glacial Great Lakes shorelines, may result in the identification of significant new colonies.
3. Conduct biosystematic research to determine the most appropriate taxonomic classification.
Bliss (1983, 1986) and Minc (1989) confirmed the validity of recognizing M. glabratus var. michiganensis as a distinct variety of M. glabratus, as established by Fassett (1939). However, significant questions remain concerning possible recognition at the species level, which, if identified as valid, would place greater urgency upon protection. Research should be conducted to address the fundamental question of relationship and taxonomic placement, particularly because of recent findings by Vickery (1990, 1991) that emphasize significant evidence for recognition above the rank of variety. Biosystematic research, involving thorough comparative studies of M. glabratus var. jamesii, M. glabratus var. michiganensis and M. guttatus, are recommended. This research should be in conjunction with isozyme and/or chloroplast DNA analyses to determine the taxonomic standing and origin of var. michiganensis. It is suggested that reciprocal crossing experiments, such as those conducted by Vickery (1991), be continued to corroborate a potential assignment of specific standing.
There are currently no long-term monitoring studies of M. glabratus var. michiganensis and there exists little knowledge of population dynamics. Beadle (1990) gathered data on ecological and physiological requirements from wide-ranging field studies of most occurrences, supplementing these data with observations of geenhouse-culture specimens. Long-term demographic monitoring is an immediate priority, in addition to continued ecological physiology studies, to clarify specific biological and abiotic requirements. These studies must be linked with research of the breeding system, which, at present, is poorly understood. Lastly, studies (e.g. isozyme analyses) to determine genetic variability would assist in understanding population structure (e.g. the number of genetic individuals within colonies) and dynamics.
42. Physiology studies - Because of the narrow restriction of M. glabratus var. michiganensis to very specific and vulnerable habitat, an understanding of physiological requirements is critical for protection and management. Both field and greenhouse research are necessary for quantifying significant physiological parameters.
422. Greenhouse research - Physiological studies are incomplete without complementary investigations of inherent parameters (temperature, nutrients, oxygen and light) determined through experiments and observations of greenhouse-cultured specimens.
43. Breeding system biology - Mimulus glabratus var. michiganensis reproduces primarily by vegetative propagation (Bliss 1986), yet is markedly different in fertility in at least one site where viable seed is produced (Beadle 1990). The breeding system is poorly known and requires comprehensive study, comprising detailed research on vegetative and floral biology to determine pollinators, ability to self pollinate, the role of sexual versus asexual reproduction, and other factors relevant to natural propagation.
44. Genetic research - Because this taxon persists largely through vegetative propagation, the number of genetically distinct individuals within sites, and thus population structure and dynamics, is unknown. Genetic research would focus on the degree of variability within and among occurrences. This information would provide critically important data relevant to protection and long-term management such as the number of genetic individuals within populations and the degree of heterozygosity and homozygosity.
45. Long-term monitoring - Long-term monitoring needs to continue beyond the potential delisting date to confirm the vitality and viability of populations. Monitoring should continue on selected sites for a minimum of five years to ensure that recovery activities have been successful and no further actions will be required.
D. Literature Cited
Beadle, S. J. 1990. A study of the ecological and physiological requirements affecting the distribution of Mimulus glabratus var. michiganensis,
the Michigan monkey-flower. Unpublished report. Submitted to the Michigan Department of Natural Resources, Wildlife Division, Nongame program, Lansing, Michigan. Unnumbered + appendices.
Bliss, M. 1982. The taxonomic status of Mimulus glabratus var. michiganensis (Pennell) Fassett. Unpublished report. 16 pp.
Bliss, M. 1983. A comparative study of the two varieties of Mimulus glabratus (Scrophulariaceae) in Michigan.
M.S. Thesis, University of Michigan, Ann Arbor, Michigan. 58 pp.
Bliss, M. 1986. The morphology, fertility and chromosomes of Mimulus glabratus var. michiganensis and M. glabratus
var.fremontii (Scrophulariaceae). American Midland Naturalist 116:125-131.
Crispin, S. R. 198 1. Mimulus glabratus var. michiganensis (Pennell) Fassett, Michigan monkeyflower (Scrophulariaceae)
in Michigan. Unpublished report. Submitted to U.S. Fish & Wildlife Service, Ft. Snelling, Minnesota. 6 PP.
Crispin, S. R.' and M. R. Penskar. 1989. Rangewide status survey of Mimulus glabratus var. michiganensis (Pennell)
Fassett, the Michigan monkey-flower. Unpublished report. Submitted to U.S. Fish & Wildlife Service, Ft. Snelling, Minnesota. 14 pp + appendices.
Fassett, N. C. 1939. Notes from the herbarium of the University of Wisconsin. Rhodora 41: 524-525.
Fuller, D 1994. Michigan landowner forest stewardship plan for Idylwilde Association. Tip-ofthe-Mitt Watershed Council. Conway, Mich. pages unnumbered.
Gleason, H. A. 1952. The New Britton and Brown Illustrated Flora of the Northeastern United States and Adjacent Canada. New York Botanical Garden, Bronx, New York. 3 volumes.
Grant, A. L. 1924. A monograph of the genus Mimulus. Annals of the Missouri Botanical Garden 11:99-389.
Jones, B. 1991. Comments on the Technical-Agency Draft of Federal Recovery Plan for Michigan monkey-flower.
September 26, 1991 letter to U.S. Fish and Wildlife Service, Region 3 Office. 2 pp.
Lewis, H. 1958. Rapid evolution of Clarkia. Evolution 12:319-336.
Michigan Dept. of Natural Resources (MDNR). 1991. Part 365, Endangered Species Protection, of the Natural Resources and
Environmental Protection Act, 1994, PA 45 1. MCL Sections 324.36501 to 324.36507. Special Plant List dated November 14, 1991.
Minc, L. D. 1989. A morphometric comparison of Mimulus glabratus var. michiganensis, M. glabratus var. fremontii, and M. guttatus
based on floral characters. Unpublished report. Michigan Natural Features Inventory, Lansing, Michigan. 20 pp.
Pennell, F. W. 1935. The Scrophulariaceae of eastern temperate North America. Monograph of the Academy of Natural Science of Philadelphia 1: 1-650.
Pennell, F. W. 195 1. Scrophulariaceae. In: L. Abrams (ed.). Illustrated flora of the Pacific States. Stanford University Press,
Stanford, California. Volume 3, pp. 686-859.
Sandland, 0. T., K. Hindar, & A. H. D. Brown. 1992. Conservation of Biodiversity for Sustainable Development.
Scandinavian University Press. Oslo 6, Norway. 324 pp.
Skottsberg, C. 1953. The natural history of the Juan Fernandez and Easter Islands, Volume 2, part 6, p 784. Almquist and Wiksells, Uppsala, Sweden.
Tai, W., and R. K. Vickery, Jr. 1970. Cytogenetic relationships of key diploid members of the Mimulus glabratus complex (Scrophulariaceae)
Tai, W., and R. K. Vickery, Jr. 1972. Unusual cytological pattern in n-ticrosporogensis and pollen development of evolutionary significance
in the Mimulus glabratus complex (Scrophulariaceae). American Journal of Botany 59:488-493.
U.S. Department of Agriculture, Forest Service. 1986. Land and Resource Management Plan, Hiawatha National Forest.
U.S. Department of the Interior, Fish & Wildlife Service. 1989. Endangered and threatened wildlife and plants; endangered status
proposed for Mimulus glabratus var. michiganensis (Michigan monkey-flower). Federal Register 54(189):40454-40458.
U.S. Department of the Interior, Fish & Wildlife Service' 1990 ' Endangered and threatened wildlife and plants;
determination of endangered status for Mimulus glabratus var. michiganensis (Michigan monkey-flower). Federal Register 55(120):25596-25599.
Vickery, R. K., Jr. 1959. Barriers to gene exchange within Mimulus guttatus (Scrophulariaceae). Evolution 13:300-310.
Vickery, R. K., Jr. 1969. Crossing barriers in Mimulus. Journal of Genetics 44 (Suppl. 1):325336.
Vickery, R. K., Jr. 1978. Case studies in the evolution of species complexes in Mimulus. In: M. K. Hecht' W. C. Steere, and B. Wallace, editors.
Evolutionary Biology. Plenum Publishing Corporation, New York, New York. Volume 1 1, pp 405-507.
Vickery, R. K., Jr. 1990. Close correspondence of allozyme groups to geographic races in the Mimulus glabratus complex (Scrophulariaceae). Systematic
Vickery, R. K., Jr. 1991. Crossing relationships of Mimulus glabratus var. michiganensis (Scrophulariaceae). American Midland Naturalist 125:368-37 1.
Vickery, R. K., Jr., K. W. Crook, D. W. Lindsay, M. M. Mia, and W. Tai. 1968. Chromosome counts in section Simiolus of the genus Mimulus
(Scrophulariaceae) VII. New numbers for M. guttatus, M. cupreus, and M. tilingii. Madroflo 19:211-218.
Voss, E.G. 1996. Michigan Flora. Part III Dicots (Pyrolaceae-Compositae). Cranbrook Institute
111. IMPLEMENTATION SCHEDULE
The Implementation Schedule that follows outlines actions and estimated costs for the recovery program. It is a guide for meeting the objectives discussed in Part II of this Plan. Actions are subdivided into tasks. This schedule indicates task priorities, task numbers, task descriptions, duration of tasks, the responsible agencies, and lastly, estimated costs. These actions, when accomplished, should bring about the recovery of the species and protect its essential habitat. The estimated funding needs for all parties anticipated to be involved in recovery are identified. Part M reflects the total estimated costs for the recovery of this species. The estimated recovery costs for the I 1 -year program are $3 16,000; $ 1 0,000 of this estimated cost is allocated to a minimum of five years of monitoring following a potential delisting date of 2004, to assess the adequacy of recovery actions and determine if there will be cause to consider re-listing. Funding is indicated beyond the potential delisting year of 2004 for some subtasks such as landowner contact and public education because these activities will be necessary even after delisting.
The costs presented are the estimates of the author and the Service, based on experience with costs of similar work. They are not based on budgets prepared for individual tasks or subtasks. Actual costs may be higher or lower than costs indicated in the Implementation Schedule.
A. Key to Implementation Schedule Column 1
Task priorities are set according to the following standards:
Priority 1: An action that must be taken to prevent extinction or to prevent the species from declining irreversibly in the foreseeable future.
Priority 2: An action that must be taken to prevent a significant decline in species population/habitat quality, or some other significant negative impact short of extinction.
Priority 3: All other actions necessary to meet the recovery objectives.
B. Key to Agency Designations in Columns 6 and 7
FWS ------U.S. Fish and Wildlife Service
TE --------Endangered Species Division, FWS
MDNR ---Michigan Department of Natural Resources
NPS ------National Park Service (Department of Interior)
MNFI ----Michigan Natural Features Inventory
TNC -----The Nature Conservancy
C. Implementation schedule for Mimulus glabratus var. michiganensis (Michigan monkey-flower) Recovery Actions
APPENDIX C. PRINCIPLE FEDERAL AND STATE LAWS APPLICABLE TO THE PROTECTION OF MIMULUS GLABRATUS VAR. MICHIGANENSIS AND ITS HABITAT
Endangered Species Act of 1973, as amended (I 6 U.S.C. 153 1-1544). Regulations, in part, at 50 CFR 17 and 50 CFR 402.
Federal Water Pollution Control Act of 1948, as amended ("Clean Water Act") (33 U.S.C. 125 11376). Regulations at 33 CFR 320-338.
National Environmental Policv Act of 1969, as amended (42 U.S.C. 4321-4347).
Part 17, Michigan Environmental Protection, of the Natural Resources and Environmental Protection Act, 1994 PA 451, as amended. MCL Sections 324.1701 - 324.1706.
Part 19, Natural Resources Trust Fund, of the Natural Resources and Environmental Protection Act, 1994 PA 451, as amended. MCL Sections 324.1901 - 324.19 1 0.
Part 21, Land Exchange Facilitation Fund, of the Natural Resources and Environmental Protection Act, 1994 PA 451, as amended. MCL Sections 324.2130 --324.2136.
Part 21, Conservation and Historic Preservation Easement, of the Natural Resources and Environmental Protection act, 1994 PA 451, as amended. MCL Sections 324.2140 324.2144.
Part 301, Inland Lakes and Streams Protection, of the Natural Resources and Environmental Protection Act, 1994 PA 451, as amended. MCL Sections 324.30101 - 324.30113
Part 303, Wetland Protection, of the Natural Resources and Environmental Protection Act, 1994 PA 451, as amended. MCL Sections 324.30301 - 324.30323.
Part 323, Shorelands Protection and Management, of the Natural Resources and Environmental Protection Act, 1994 PA 451, as amended. MCL Sections 324.32501 - 324.32515.
Part 353, Sand Dune Protection and Management, of the Natural Resources and Environmental Protection Act, 1994 PA 451, as amended. MCL Sections 324.35301 - 324.35326.
Part 365, Endangered Species Protection, of the Natural Resources and Environmental Protection Act, 1994 PA 451, as amended. MCL Sections 324.36501 - 324.36507.
APPENDIX D. PEER REVIEW AND PUBLIC COMMENT
Development of this recovery plan began shortly after listing in July, 1990, before the Service issued its July 1, 1994, policy on peer review. A draft recovery plan for Mimulus glabratus var. michiganensis (Michigan monkey flower) was made available for public review and comment for 30 days on June 10, 1991 (56 FR 26693). The administrative record for public review and comment on this plan is maintained by the Service Regional Office in Twin Cities, MN.
Although 20 or more reviewers, representing a wide spectrum of agencies and private individuals, received copies of the combined technical/agency draft plan, the reviewers listed here consist only of those who submitted written comments to the U.S. Fish and Wildlife Service. Comments and opinions received are not summarized separately in this plan, but have been considered. and incorporated when appropriate into the Final Recovery Plan.
1420 Providence Hwy.
Norwood MA 02062
Mr. Robert Jones
Glen Arbor, Michigan 49636
Dr. Robert K. Vickery, Jr.
Department of Biology
University of Utah
Salt Lake City, Utah 84112
Dr. Edward G. Voss
North University Building
University of Michigan
Ann Arbor, Michigan 48109-1057