Plebejus melissa samuelis - (Nabokov, 1944)
Karner Blue
Other English Common Names: Karner Blue Butterfly, Karner blue butterfly
Synonym(s): Lycaeides melissa samuelis Nabokov, 1944
Taxonomic Status: Accepted
Related ITIS Name(s): Plebejus melissa samuelis (Nabokov, 1944) (TSN 778920)
Unique Identifier: ELEMENT_GLOBAL.2.120997
Element Code: IILEPG5021
Informal Taxonomy: Animals, Invertebrates - Insects - Butterflies and Moths - Butterflies and Skippers
 
Kingdom Phylum Class Order Family Genus
Animalia Mandibulata Insecta Lepidoptera Lycaenidae Plebejus
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Concept Reference
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Concept Reference: Nabokov, V. 1944. The Nearctic formes of Lycaeides Hüb. (Lycaenidae, Lepidoptera). Psyche 50(3/4): 97-99. [Sep.-Dec. 1943, 1944].
Concept Reference Code: A44NAB01EHUS
Name Used in Concept Reference: Lycaeides melissa samuelis
Taxonomic Comments: Several biological differences and the lack of intergrades suggest this is a separate species. However, all literature to date (December, 2011) treats it as subspecies. Also results of recent electrophoretic work by Laurence Packer and colleagues are most consistent with (but do not prove) subspecies status. There is some DNA evidence suggestive of differences between eastern populations and those farther west, but it does not seem likely taxonomic separation will prove warranted. Native populations of the eastern type survive now only in New York's Hudson Valley, but stock from there has been introduced to New Hampshire where the subpecies was native. See Recovery Plan for more details. This species has been put in Lycaeides by many recent authors, but research summarized in Opler and Warren (2002) has resulted in sinking of that genus and several others.
Conservation Status
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NatureServe Status

Global Status: G5T2
Global Status Last Reviewed: 30Mar2008
Global Status Last Changed: 01Sep1998
Rounded Global Status: T2 - Imperiled
Reasons: Major declines were documented in the 1970s and 1980s at most of the larger occurrences, often of 90% or more. The extirpation rate was high for populations that fell below 1000 in the July brood. These declines led to the subspecies being federally listed as Endangered, and it has stabilized substantially as a result. The Karner Blue is extirpated in Maine, New Hampshire (but reintroduced), Massachusetts, Pennsylvania, Ohio (but likely to be reintroduced), Ontario; and also New Jersey and Iowa if it really occurred in those states. However, there are still several substantial protected metapopulation occurrences in Michigan and Wisconsin, at least one in Indiana, and a few potentially viable smaller occurrences in these and other states, notably Minnesota, and at least one large population in New York. The Karner Blue has become management dependent in all parts of its range, and would be unlikely to persist more than a decade or two if management were to cease. This subspecies was usually found in uncommon and declining natural communities such as inland pine barrens and oak savannas, or in artificial habitats like airports where such communities formerly existed.
Nation: United States
National Status: N2 (01Sep1998)
Nation: Canada
National Status: NX (01Sep1998)

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 Illinois (S1), Indiana (S1), Iowa (SNR), Maine (SX), Massachusetts (SX), Michigan (S2), Minnesota (S1), New Hampshire (S1), New York (S1), Ohio (S1), Pennsylvania (SX), Wisconsin (S3)
Canada Ontario (SX)

Other Statuses

U.S. Endangered Species Act (USESA): LE: Listed endangered (14Dec1992)
Comments on USESA: Listed Endangered as Lycaeides melissa samuelis.
U.S. Fish & Wildlife Service Lead Region: R3 - North Central
Canadian Species at Risk Act (SARA) Schedule 1/Annexe 1 Status: XT (05Jun2003)
Committee on the Status of Endangered Wildlife in Canada (COSEWIC): Extirpated (25Apr2010)
Comments on COSEWIC: Reason for designation: Has not been observed since 1991.

Status history: Designated Extirpated in April 1997. Status re-examined and confirmed in May 2000 and April 2010.

IUCN Red List Category: NE - Not evaluated

NatureServe Global Conservation Status Factors

Range Extent: 20,000-200,000 square km (about 8000-80,000 square miles)
Range Extent Comments: Currently found and ranked in MN, WI, IN, NY, MI. Extirpated from IL, OH, ON, PA, MA., NH, but reintorduced in NH. Falsely reported from NC and MB. Significant range confined to Wisconsin and Michigan just isolated dots on a map elsewhere.

Area of Occupancy: 26-2,500 4-km2 grid cells
Area of Occupancy Comments:  

Number of Occurrences: 6 - 20
Number of Occurrences Comments: Metapopulations: apparently one or two A rank, less than 5 B rank (some marginally so and/or crashing), perhaps 5 or more C ranked, and > 20 D-ranked.

Population Size: 2500 - 10,000 individuals
Population Size Comments: Estimated 1,000-2,000 for Albany NY area (1987-1988, lower in 1989); 2,000 at Alleghan (1989), several thousand at Indiana Dunes National Lakeshore (1992). 14,000 at Saratoga airport in NY (1990), most EO's all <1,000.

Number of Occurrences with Good Viability/Integrity: Few (4-12)

Overall Threat Impact: Medium
Overall Threat Impact Comments: Despite listing some habitats continue to deteriorate, for example due to deer at Albany. Some populations may simply be too small to survive long term. Some of the better populations probably are now relatively secure. Inappropriate fire regimens, loss of habitat to urbanization and pine farms, and out of control deer habeen major cause of local or general decline.

Short-term Trend: Decline of <30% to relatively stable
Short-term Trend Comments: Has stabilized somewhat with federal listing.

Long-term Trend: Decline of 70-90%
Long-term Trend Comments: Severe decline started in 1970's, 1987-1988 droughts finished off numerous remnant demes. Most current EOs (D-ranked) expected to be lost. Much habitat lost before 1970 also. 98% decline since 1979 at best known EO (Albany NY Pine Bush) despite preservation of large area, without management. No chance of survival now without management. Significant decline at Alleghan, MI. Many EOs lost in 1980's.

Environmental Specificity: Narrow. Specialist or community with key requirements common.

Other NatureServe Conservation Status Information

Inventory Needs: Inventory now largely complete. Evaluation of size and defensibility of remaining EOs is now the major need. Status at Ft. McCoy, WI needs investigation - seems to be a major occurrence.

Protection Needs: Restoration of at least 5 metapopulations to A rank specs or solid B rank specs recommended. Albany Pine Bush and Alleghan were A rankable in 1970's; thus top priorities. Indiana Dunes and Ft. McCoy, WI are also important sites. Translocation offers only hope in NH.

Distribution
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Global Range: (20,000-200,000 square km (about 8000-80,000 square miles)) Currently found and ranked in MN, WI, IN, NY, MI. Extirpated from IL, OH, ON, PA, MA., NH, but reintorduced in NH. Falsely reported from NC and MB. Significant range confined to Wisconsin and Michigan just isolated dots on a map elsewhere.

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 IA, IL, IN, MAextirpated, MEextirpated, MI, MN, NH, NY, OH, PAextirpated, WI
Canada ONextirpated

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
IL Lake (17097)*
IN Lagrange (18087), Lake (18089), Porter (18127)
MI Allegan (26005), Ionia (26067), Kent (26081), Lake (26085), Mason (26105), Mecosta (26107), Monroe (26115)*, Montcalm (26117), Muskegon (26121), Newaygo (26123), Oceana (26127)
MN Anoka (27003), Isanti (27059), Winona (27169)
NH Hillsborough (33011)*, Merrimack (33013), Rockingham (33015)*
NY Albany (36001), Erie (36029)*, Genesee (36037)*, Jefferson (36045)*, Kings (36047)*, New York (36061)*, Niagara (36063)*, Oneida (36065)*, Queens (36081)*, Saratoga (36091), Schenectady (36093), Warren (36113)
OH Lucas (39095)
WI Adams (55001), Burnett (55013), Chippewa (55017)*, Clark (55019), Eau Claire (55035), Green Lake (55047), Jackson (55053), Juneau (55057), Marquette (55077), Menominee (55078), Monroe (55081), Oconto (55083), Outagamie (55087), Polk (55095)*, Portage (55097), Sauk (55111), Waupaca (55135), Waushara (55137), Wood (55141)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
01 Contoocook (01070003)+*, Nashua (01070004)+*, Merrimack (01070006)+
02 Hudson-Hoosic (02020003)+, Mohawk (02020004)+, Middle Hudson (02020006)+, Lower Hudson (02030101)+*, Bronx (02030102)+*, Sandy Hook-Staten Island (02030104)+*, Southern Long Island (02030202)+*
04 Oconto (04030104)+, Upper Fox (04030201)+, Wolf (04030202)+, Little Calumet-Galien (04040001)+, Pike-Root (04040002)+*, St. Joseph (04050001)+, Black-Macatawa (04050002)+, Kalamazoo (04050003)+, Lower Grand (04050006)+, Pere Marquette-White (04060101)+, Muskegon (04060102)+, Ottawa-Stony (04100001)+, Raisin (04100002)+*, Lower Maumee (04100009)+, Niagara (04120104)+*, Oak Orchard-Twelvemile (04130001)+*, Oneida (04140202)+*, Chaumont-Perch (04150102)+*, Upper St. Lawrence (04150301)+*, Mettawee River (04150401)+
07 Rum (07010207)+, Upper St. Croix (07030001)+, Namekagon (07030002)+, Lower St. Croix (07030005)+, Buffalo-Whitewater (07040003)+, La Crosse-Pine (07040006)+, Black (07040007)+, Lower Chippewa (07050005)+*, Eau Claire (07050006)+, Castle Rock (07070003)+, Chicago (07120003)+
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Basic Description: Karner Blue Butterfly
General Description: See any butterfly book for eastern North America. The dorsal side of the male is silvery blue or dark blue with narrow black margins. The females are grayish brown dorsally, with irregular bands of orange inside the narrow black border on the upper wings. Both sexes are slate gray on the ventral side with the orange bands showing more regularity, and black spots circled with white. (N92CLO01EHUS)
Diagnostic Characteristics: Within its range the combination of orange marginal spots beneath and lack of a tail on hw is diagnostic. L. m. melissa approaches from the west and is separable by well defined orange orange spots, often fused as a band, above in females, subspecies samuelis rarely has traces of these spots. Habitat also differs. L. idas approaches its range from the north and is very similar--males almost identical. It differs in habitat and also has only one brood in June-early July. Male genitalia also differ.
Non-Migrant: N
Locally Migrant: N
Long Distance Migrant: N
Mobility and Migration Comments: Basically sedentary, but adults do sometimes move at least two kilometers, probably more. Most probably never go more than 100-200 meters from place of emergence. Some evidence that dispersal is most likely where habitat quality is declining or where nectar is scarce.
Terrestrial Habitat(s): Grassland/herbaceous, Old field, Sand/dune, Savanna, Woodland - Conifer
Habitat Comments: In eastern New York and New Hampshire, habitat typically is in sandplain communities, such as grassy openings within very dry, sandy pitch pine/scrub oak barrens. In the Midwest, the habitat is also dry and sandy, including oak savanna and jack pine barrens, and less often dune communities. Within the overall coummunity remnant inhabited by a metapopulation any patch of foodplant in open to semi-shaded setting is likely to be used. Females lay eggs on or near wild lupine plants, and main requirement seems to be thousands of stems of lupine in the short term. Apparently persistence from xerothermic interval to present requires thousands of hectares of suitable community/habitat in which patches of occupied habitat probably shifted over time.
Adult Food Habits: Nectarivore
Immature Food Habits: Herbivore
Food Comments: Wild lupine is the only known larval food plant and is therefore closely tied to the butterfly's ecology and distribution. A variety of other understory plants associated with the habitat serve as nectar sources for the adults. Adult males also drink from moist sand.
Adult Phenology: Diurnal
Immature Phenology: Hibernates/aestivates
Phenology Comments: Apparently always two broods each year. Eggs that have overwintered from the previous summer hatch in April. Near the end of May or early in June, the larva pupate and adult butterflies emerge very late in May and well into June in most years. The adults are typically in flight for the first 10 to 15 days of June, when the wild lupine is in bloom. New eggs are laid on or near the lupine plants, and hatch in about one week, and the larvae feed for about three weeks. They then pupate and the second brood adults appear in the second or third week of July, sometimes earlier in advanced seasons. Adults typically fly into about mid August. Individual adults live an average of about five days, but females at least can occasionally live for two weeks. This time, the eggs are laid among plant litter or on grass blades at the base of the lupines, or on lupine pods or stems, and these eggs do not hatch until the following spring.
Economic Attributes Not yet assessed
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Management Summary
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Stewardship Overview: This may be the most studied butterfly in North America. This subspecies is listed and therefore protected under the US Endangered Species Act. US Fish and Wildlife Service and/or state agency personnel are involved in management of all known occurrences. Anyone involved with managing lands with Karner Blue populations should consult the Recovery Plan, which summarizes what is known or reasonably suspected regarding management and monitoring. However, new information, much of it relating to management, appears every year, making consultation with USFWS or state biologists familiar with this species crucial. A lot of information is available on line.

The primary goals should be to maintain enough lupine, nectar, and other resources to sustain the occurrence and if possible to create a "viable population" as defined in the Recovery Plan, which usually would consist of a metapopulation of at least 3000 adults in the larger brood each year. Past observations suggest smaller populations sometimes suddenly crash and fail to recover. Other important issues include connecting remnant colonies into viable metapopulations, allowing for recolonization after fires (prescribed or otherwise), or in some cases implementing an alternate management strategy such as late summer or dormant season mowing. Ideally lupines should be well distributed through the habitat in both very open and partially shaded situations with varying exposures, and it is important that the summer brood has easy access to several different genera of nectar flowers. Some level of monitoring should be implemented. Karner Blues usually occur with other species that are locally, and sometimes globally, as rare or rarer and often occur in remnant or somewhat degraded examples of relatively rare natural communities. These other species need to be considered in management. Schweitzer et al. (2011) provides information on most such species as well as the Karner Blue

Restoration Potential: See Recovery PLan. If habitat is suitable, this species can be translocated (with proper permits).
Preserve Selection & Design Considerations: See Recovery Plan
Management Requirements: See Recovery Plan. Note fire is an appropriate management tool but most or all individuals in the burned area will be killed. If refugia are nearby though recovery is rapid.
Monitoring Requirements: See Recovery Plan. Mark-release-recapture is generally necessary if and only if, an estimate of populations size is needed. Changes can be tracked with simpler census methods.

Management Programs: USFWS and appropriate stat DNR etc.

Monitoring Programs: USFWS and appropriate stat DNR etc.
Biological Research Needs: See Lawrence and Cook report (MI Heritage), consult upcoming ESA (NYFO), also and extensive compilation distributed at 1992 Zanesville conference. Decent (MRR) population estimates should be made at certain sites.
Population/Occurrence Delineation
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Use Class: Not applicable
Minimum Criteria for an Occurrence: A location where Karner Blues occur, or have recently occurred, where there is potential for persistence or regular recurrence. Minimally a place where adults or immatures have been verified in association with sufficient habitat and lupine to sustain a population or (high quality occurrences) a metapopulation. Occurrences ranked higher than D should have the species every year. Occurrences ranked higher than D will usually, and those ranked higher than C will probably always, be metapopulations in some sort of savanna or pine barren context. With metapopulations a patch of habitat with foodplant in the overall habitat matrix is part of the occurrence even if temporarily vacant (e.g. small patches after fires) unless it is known for some reason to be consistently unoccupied. The recommendations of the 2003 US Fish and Wildlife Service Recovery Plan (N03FWS01ehus) should be followed on this point.

Mapping Guidance: Habitat patches are specifically openings or lightly wooded terrain with lupine and nectar where adults and immatures occur. Occasionally these two resources are a few dozen meters apart but they should be mapped together. Most occurrences now, and almost certainly all originally, are metapopulations consisting of multiple more or less discrete habitat patches.
Separation Barriers: It does not seem likely this concept should be invoked in defining occurrences for Karner Blues. There is some discussion of potential barriers in the Recovery Plan but few landscapes are absolutely uncrossable. Features like extensive asphalt, busy four lane highways and dense closed-canopy forest do to some degree impede dispersal.
Separation Distance for Unsuitable Habitat: 4 km
Separation Distance for Suitable Habitat: 8 km
Alternate Separation Procedure: When multiple colonies occur in the same large sandy pine barren or oak savanna landscape separation distances may be violated, especially for practical reasons. However it is suggested separation distances should generally not exceed 8 km. Conceptually at least some of the intervening habitat can be considered to be potentially restorable and probably some was recently suitable and occupied. Populations within such xeric communities were obviously not persistently isolated, even if they are now, and sometimes a realistic conservation and management goal will be to reconnect them. It also seems pointless to consider small isolated demes as separate low quality (D-ranked)occurrences as opposed to assuming a core-satellite model even if this means violating standard separation distances. See Recovery Plan especially Appendix G, regarding such issues. However there is no actual evidence of adults moving more than 6.6 km and movements more than 1500 meters appear to be rare.
Separation Justification: There are many studies (see Recovery Plan Appendix G and several references cited) that clearly show a pattern of high patch occupancy when habitats are close together in metapopulations, inability of isolated colonies to persist, and good short range (up to about 1-2 km) colonization between habitats. This virtually an obligate metapopulation species. Movements of more than about 200-300 meters are infrequent in most places, but important. Schweitzer in New Hampshire found virtually no movement even with over 1000 individuals marked, as have a few other studies with smaller numbers. The King study in Wisconsin in which about 11% moved at least 1,150 meters represents the opposite extreme in documented dispersal and may very well approximate the original conditions for the species. His maximum documented movement was 6.6 km. As discussed in Appendix G it can be reasonably assumed that most studies will underestimate significant movements due to declining probability of recapture with radial distance. Notably some of the longer distances documented were along right of ways or other corridors. It seems clear from the various studies cited in the Recovery Plan that movements are greatly impeded by closed canopy and are greatest in more open situations. Colonies more than one to two kilometers apart in most current landscape contexts probably no longer function as an effective metapopulation, but that was not originally the case and usually will not be with successful recovery efforts.

While a 2 km separation distance could certainly be justified by the data for this species, 4 km, which is still well below the largest recorded movement, is suggested as more practical for the purpose of defining separate occurrences in that is it likely to keep remnants of the same original metapopulation as conceptually one (perhaps restorable) occurrence. Also intermediate habitat patches probably or definitely have occurred in the foreseeable past connecting such proximate colonies and might again with management. While some gene flow will probably occasionally or rarely occur between populations 4 km apart it is important to stress that this distance may be too great to maintain good metapopulation dynamics.

The suitable habitat distance should be applied mainly in the case of what USFWS describes as a "living corridor" (Recovery Plan p. G-81). These are situations such as right of ways, roadsides etc. that have some lupine on which eggs and larvae occasionally occur and sufficient nectar to facilitate movements. Such places cannot support Karner Blues on their own but have sufficient resources to facilitate use and inter-site movement. As USFWS points out even if individuals do not traverse the entire distance in one generation, their progeny that mature in this corridor are likely to do so. The distance selected is twice the unsuitable habitat distance. This distance is slightly greater than the maximum individual movement actually documented (6.6 km) but much less that twice that distance. This concept may also be used between discrete core population centers separated by woodland, scrub or barrens with only scattered lupine and nectar.




Inferred Minimum Extent of Habitat Use (when actual extent is unknown): 1 km
Inferred Minimum Extent Justification: This is usually moot since usually extent will be known and need not be inferred. The inferred extent is the actual habitat patch which is usually less than 100 hectares and rarely 1 km in any direction, or in some cases would be used to infer presence in proximate patches if a metapopulation is actually known to occupy the area. If the actual occupied habitat patch is more than 1 km across, then the entire extent is still inferred habitat. Patches are either unoccupied or vacant, not partially occupied, over any reasonable time scale
Date: 29Nov2004
Author: Schweitzer, Dale F.
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: 19May2005
NatureServe Conservation Status Factors Author: Schweitzer, D.F.
Management Information Edition Date: 08Mar2013
Management Information Edition Author: Schweitzer, Dale F.
Element Ecology & Life History Edition Date: 06Jan1993
Element Ecology & Life History Author(s): SCHWEITZER, D. F.

Zoological data developed by NatureServe and its network of natural heritage programs (see Local Programs) and other contributors and cooperators (see Sources).

References
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  • Bess, J.A. 1989. Status of the Karner Blue butterfly, Lycaeides melissa samuelis Nabokov, in the Manistee National Forest. Michigan Natural Features Inventory, Natural Heritage Program, Wildlife Division. 20 pp. + appendices.

  • Bess, James. 2005. A Report on the Remnant-Dependent Insects of the Coastal Zone Natural Area Remnants in Northwest Indiana. 23 pp..

  • Bleser, C. A. 1994. Karner Blue Butterfly survey, management and monitoring activities in Wisconsin: 1990-Spring 1992. Pages 153-162 in D. A. Andow, R. J. Baker, and C. P. Lane, editors. Karner Blue Butterfly: a symbol of a vanishing landscape. Miscellaneous Publication 84-1994, Minnesota Agricultural Experiment Station, University of Minnesota, St. Paul, Minnesota.

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  • Carson, P.J. 1996. Draft. Status report on the Karner Blue butterfly Lycaeides melissa samuelis in Canada. COSEWIC - Committee on the Status of Endangered Wildlife in Canada. 24 pp.

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  • Committee on the Status of Endangered Wildlife in Canada (COSEWIC). 2010a. COSEWIC Assessment Results, April 2010. Online. Available: http://www.cosewic.gc.ca/.

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  • Cuthrell, David L. 1990. Status of the Karner Blue Butterfly, Lycaeides melissa samuelis Nabokov, in Minnesota 1990. Funded by the MN DNR, Section of Wildlife, Natural Heritage and Nongame Research Program, the US Fish & Wildlife Service, and The Nature Conservancy. Results in unpublished report.

  • Dirig, R. 1994. Historical notes on Wild Lupine and the Karner Blue Butterfly at the Albany Pine Bush, New York. Pages 23-36 in D. A. Andow, R. J. Baker, and C. P. Lane, editors. Karner Blue Butterfly: a symbol of a vanishing landscape. Miscellaneous Publication 84-1994, Minnesota Agricultural Experiment Station, University of Minnesota, St. Paul, Minnesota.

  • Eberlie, W.J.D. 1987. Request for protection of the Karner Blue butterfly and the Frosted Elfin butterfly under the Endangered Species Act. Unpublished, letter to Vincent G. Kerrio, Minister of Natural Resources, dated Nov. 20, 1987. 3 pp.

  • Ewert, D.N. and H.E. Ballard, Jr. 1990. Distribution and management of an indicator species of Oak Barrens, the Karner Blue butterfly (Lycaeide melissa samuelis): pre-treatment population studies. Report for The Nature Conservancy, Michigan Chapter, dated November 27, 1990. 10 pp.

  • Gall, L. F. 1985. Measuring the Size of Lepidopteran Populations. Journal of Research on the Lepidoptera. 24(2):97-116.

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  • Herms, C. P., D. G. McCullough, L. S. Bauer, R. A. Haack, D. L. Miller, and N. R. Dubois. 1997. Susceptibility of the endangered Karner Blue Butterfly (Lepidoptera: Lycaenidae) to Bacillus thuringiensis var. kurstaki used for gypsy moth suppression in Michigan. The Great Lakes Entomologist 30:125-141.

  • Herms, C.P., D.G. McCullough, D.L. Miller, L.S. Bauer and R.A. Haak. 1996. Laboratory rearing of Lycaeides melissa samuelis (Lepidoptera: Lycaenidae), an endangered butterfly in Michigan. The Great Lakes Entomologist 29(2): 63-75.

  • Hirsch, L.P. 1978. Status of the Karner Blue Butterfly in New York (Lycaeides melissa samuelis). Proposed Rulemaking for the Department of the Interior, Fish and Wildlife Service, Endangered and Threatened Wildlife and Plants. 6 pp.

  • Huber, R. L. 1981. An updated checklist of Minnesota butterflies. Minnesota Entomological Association Newsletter 14(3):15-25.

  • Iftner, D. C., J. A. Shuey, and J. V. Calhoun. 1992. Butterflies and Skippers of Ohio. Ohio Biological Survey Bulletin. New Series, Vol. 9, no. 1, xii + 212 pp., 40 color plates.

  • Konecny, A. 1986. A Status report on the Karner blue butterfly (Lycaeides melissa samuelis Nabokov) in Canada - DRAFT. Prepared for Nongame Program, Wildflie Branch, OMNR. 48 pp.

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  • Lane, C. P. 1992. The status of the karner blue butterfly (Lycaeides melissa samuelis: Lycaenidae) and its associated plant resources in Minnesota 1991. Final report to U.S. Fish and Wildlife Service, Minnesota Department of Natural Resources, and The Nature Conservancy. 23 pp.

  • Lane, C. P. 1999. Benefits of heterogeneous habitat: oviposition preference and immature performance of Lycaeides melissa samuelis Nabokov (Lepidoptera: Lycaenidae). Dissertation, University of Minnesota, St. Paul, Minnesota. 185 pp.

  • Lane, C., and R. Dana. 1994. The status of the Karner Blue Butterfly in Minnesota. Pages 113-122 in D. A. Andow, R. J. Baker, and C. P. Lane, editors. Karner Blue Butterfly: a symbol of a vanishing landscape. Miscellaneous Publication 84-1994, Minnesota Agricultural Experiment Station, University of Minnesota, St. Paul, Minnesota.

  • Lawrence, W.S. and A.C. Cook. 1989. The status and management of Karner Blue (Lycaeides melissa samuelis) populations in the Allegan State Game Area, Michigan. A report subitted to The Nature Conservancy, Michigan Field Office. 58 pp. + appendices pp.

  • Magdich, M. 1989. The Karner Blue. Newsletter of the Division of Natural Areas & Preserves, Ohio Department of Natural Resources, Vol. 11, No. 3: 1-2.

  • Malicky, H. 1970. New aspects on the association between lycaenid larvae (Lycaenidae) and ants (Formicidae, Hymenoptera). Journal of the Lepidopterists' Society 24:190-202.

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