endemic to a single nation
Regularity: Regularly occurring
Type of Residency: Year-round
Global Range: (5000-200,000 square km (about 2000-80,000 square miles)) The orginal range was much larger and more discontinuous than the current range which is a small narrow area of shale ridges from south-central Pennsylvania into Virginia and isolated occurrences in Ohio.The maps in popular guides tend to overstate the original range which actually was in several disjunct parts. The core of the range was the shale region from about Huntington County, Pennsylvania southwest though northeastern West Virginia and adjacent Virginia including a bit of far western Maryland. These populations reached east, not exclusively on shale, to Lancaster and Dauphin Counties, Pennsylvania in the 1950s and Washington, D.C. in the 1880s (specimen from Rock Creek Park in Academy of Natural Sciences). There are no gaps more than two counties wide from Dauphin County and D.C. to extreme southwest Virginia and (LeGran and howard, 2008) adjacent North Carolina. There were also a few seemingly isolated records in the early 1970s for central New York which could have been an extension of the main range. There were disjunct well known populations in the trap rock glades region and surrounding areas in northern New Jersey (specimens from seven counties, Iftner and Wright, 1996) and immediately adjacent New York into the late 1950s. There are or were several apparently disjunct populations in southern Ohio. Consult Shapiro (1974) for New York, Iftner et al. (1993) for Ohio, and Allen (1997) for West Virginia. A separate and distinctive group of populations in northern Michigan has traditionally also been included with this taxon but see taxonomy field. Minnesota populations should be referred to the taxon freija. The pre-1863 Long Island record was probably in error for a nearby part of New Jersey or New York since no likely edaphic formations occur on Long Island. The Cook Co., Illinois dot shown by Parshall (2002) is very dubious and has generally been ignored. This also seems to date from around 1900 or earlier and collectors were often sloppy with data in those days. Information on the three reports along Lake Erie in New York and Ohio (Parshall, 2002, Shapiro, 1974) is unavailable. Aside from omitting the seven New Jersey counties, the map in Parshall (2002) seems to be the most complete compilation. The Kentucky record was probably false.
The flight period is close to diagnostic by itself, but to be certain field marks should be observed. Either a specimen or a photograph should be easily separable from Pyrgus communis by the more olive shades beneath, the duskier hairier appearance and smaller and fewer white spots above, for example the lack of the two rows of well-defined small marginal spots on the forewing. Also the white spots on the hindwing above occupy much less of the total surface.and the second spot from rear on postmedian band of forewing is strongly offset inward. Opler and Krizek (1984) and Glassberg (1993), provide especially good photographs of upperside. See these or Brock and Kaufman (2003) or virtually any other butterfly book that illustrates P. communis and either P. wyandot or the similar P. centaureae. Flying individuals probably should not be reported as valid records although it is unlikely the variable P. communis would be flying as early as P. wyandot. Either follow it until it lands or net it if a definite identification of that individual is needed.
Comments: Habitat is openings within jack pine or pine-oak barrens in Michigan. For typical P. wyandot habitat was trap rock glades and associated woodland in northeastern New Jersey and adjacent New York. Typical Appalachian habitat is/was shale barrens, pastures and powerlines on south to west facing shale slopes, always with much bare rock or soil. Key features were plentiful Potentilla canadensis, the larval foodplant, and nectar flowers, and also some source of moisture such as a streamlet at the base of the hill, or even deep wheel ruts. See Schweitzer (1989) for more detailed discussion of habitat. Habitat could formerly easily be spotted from a car using the features in Schweitzer (1989). Adults seldom occur more than about 30 meters from woods and sometimes occur in the woods. Shale barrens and other habitats tend to be surrounded by scrubby oak woodland or forest with some to a lot of Virginia pine. Oaks in such xeric habitats leaf out late, leaving no canopy for much of the flight season making these woods less of a barrier to adults than one would expect. When the trees are bare adults can be found in the forest occasionally. The exact nature of the habitats near Ithaca New York is less certain. It appears they were successional stages in reforestation (Robert Dirig email to D. Schweitzer, 15 February 2005; limited earlier observations by Schweitzer. That brings up the question of where the original colonizers for this and other rare species came from One can see from the geological and butterfly distribution maps of Shapiro (1974) that the area has much shale and also had Glaucopsyche lygadamus lygdamus (=nittanyensis), another characteristic Appalachian shale inhabitant. Some Virgina habitats also appear to be successional and are not classic shale barrens (S. Roble). Colonies occasionally occurred as high as about 760 meters in West Virginia and the current North Carolina populations are slightly over 900 meter, but most colonies are at lower elevations and some older ones were below 100 meters.
Non-Migrant: No. All populations of this species make significant seasonal migrations.
Locally Migrant: No. No populations of this species make local extended movements (generally less than 200 km) at particular times of the year (e.g., to breeding or wintering grounds, to hibernation sites).
Locally Migrant: No. No populations of this species make annual migrations of over 200 km.
Adults apparently are very dispersive within ridge systems, especially along powerlines. Allen and Schweitzer marked several in West Virginia, none were present next day, new ones were.
Comments: Larvae feed on wild strawberry in Michigan, apparently exclusively on POTENTILLA CANADENSIS elsewhere. A variety of spring wild flowers are used for nectar, including pussy toes (ANTENNARIA), birds foot violet, and PHLOX SUBULATA.
Number of Occurrences
Note: For many non-migratory species, occurrences are roughly equivalent to populations.
Estimated Number of Occurrences: 1 - 80
Comments: Parshall (2002) reports it to be extant in six counties total in Pennsylvania, Maryland, West Virginia, Virginia, and Ohio. Probably each represents a single occurrence but this is not certain for Virginia. He also reports the Michigan entity as extant in two adjacent counties. It is very probable but not certain that <20 occurrences are extant.
250 - 10,000 individuals
Comments: Seldom common. Ten per hour would be exceptional. With fewer than ten known extant populations, it is quite possible there are fewer than 1000 adults rangewide most years now. However adults do seem to move around and effective population sizes may be larger than they appear.
Life History and Behavior
Comments: Adults present April (rarely late March) to early May in Appalachia: formerly late April-mid May in New Jersey and New York; May in Michigan. Larval period is approx. 100 days (in West Virginia, T. Allen) beginning at end of adult season. Larvae remain on foodplant all hours. Adult activity period is quite restricted on cool days.
National NatureServe Conservation Status
Rounded National Status Rank: N1 - Critically Imperiled
NatureServe Conservation Status
Rounded Global Status Rank: G1 - Critically Imperiled
Reasons: Parshall (2002) and other recent efforts have found this species extant in only about 13% of counties with records in the 1990s. There are only about six or slightly more known extant occurrences. It thus seems likely there really are under 20, possibly only one or two, extant viable occurrences, but some recoverable occurrences. This is probably among the three most imperiled butterfly or skipper species in North America and is imminently threatened in some areas, extirpated in most others, and it does not appear secure anywhere. It is lready extirpated from a major portion of range (New York-New Jersey and apparently most of its more eastern range within Pennsylvania). Very few occurrences documented in or before the mid 1980s remained extant after half a decade of gypsy moth spray programs. Many to most metapopulations in core of range are apparently extinct. The status of this species could change very quickly in either direction depending on degree of gypsy moth spraying in the near future. There might still be 10-20 viable or recoverable occurrences, but far fewer are now known, and recent decline and/or threats are severe in most of range. If Michigan populations prove to be distinctive, then both taxa should probably get the same rank as imperiled. Threats are similar, and the Michigan entity is now possibly extirpated from ten of the twelve counties where it occurred (Parshall, 2002).
Environmental Specificity: Very narrow. Specialist or community with key requirements scarce.
Comments: One of the most specialized skippers east of the prairies. While the foodplant is very common, even as a lawn weed, on most substrates, this skipper was restricted or nearly so to a narrow range of very hot rock outcrop habitats. By far most populations were on shale barrens or sparse woodlands (Pennsylvania to Virginia) or traprock glades (new Jersey immediately adjacent New York) or on right of ways, very open woods, or disturbances on south or west facing slopes of these substrates. Furthermore populations were always very close to woods even if the foodplants extended far out in the open. See Schweitzer (1989) report for details. Habitats were consistent enough though in 1985-1986 that presence could often be predicted from looking at a site from a kilometer or two away!
Other Considerations: The reasons for the lack of US Endangered Species listing for this species are unknown, but apparently do not reflect current knowledge including status survey results. It is actually not certain there are any extant long-term viable occurrences remaining.
Global Short Term Trend: Unknown
Comments: At present it is possible this species is recovering some from losses around 15-20 years ago. This species is so reduced now that any trend would be hard to notice and a major decline would end in extirpation in most or all parts of range.
Global Long Term Trend: Decline of >90%
Comments: As mapped by Parshall (2002) and with the seven New Jersey counties added and allowing for dubious records, the historic range of typical wyandot included about 45 counties some with multiple populations. The Michigan entity is reported from 14 counties. The current ranges include six and two counties. As far as known the species was basically stable until about 1958-1960 and especially 1987-1992. Catastrophic decline about 1987 to 1995 due to Dimilin and probably Btk spraying directed at gypsy moth, in combination with drought. A similar decline to extirpation occurred in New Jersey about 1958-1960 in conjunction with large scale DDT spraying of habitats also for gypsy moth which peaked about 1957. Most known populations in Maryland and West Virginia were eradicated soon after 1986, including apparently all of those referred to by Schweitzer (1989). It is unclear what happened to the central New York populations and these may not have ever really been established. Apparently the few specimens were all collected in a period of less than five years in a generally well studied area. It is nearly certain these populations are extirpated. As far as known the sites were all unnaturally disturbed areas succeeding back to forest. This species was common in Virginia in the early 20th century, and now is among the most imperiled skippers in the US.
Degree of Threat: Very high - high
Comments: That this species was reduced largely by gypsy moth spraying is now widely accepted and obviously correct, although there probably were other factors. Gypsy moth spraying eliminated most known Appalachian populations from about 1985-1992, and all known ones in New Jersey about 1958-1960. A few more applications of Dimilin or BTK in key areas could finish off this species. There also appears to be a change in the abundance of host plants on some West Virginia historic sites with Potentilla canadensis being represented by far fewer plants. This may reflect droughts. Numbers are now so low that even collectors may constitute a threat to remaining colonies, although collecting was not a factor in the crash. Likewise minor fluctuations could cause colonies to "wink out". Low numbers and fragmentation greatly increase this threat, and this species probably cannot survive unless some metapopulation function is restored. Broadcast herbiciding of powerlines would also be a very potent threat considering that powerline corridors were major habitats in the 1980s and will almost certainly be important if this species ever recovers. They appear to be better dispersal corridors than any kind of natural feature. The most likely cause of the loss of the central New York populations was succession (Robert Dirig, email to Schweitzer 15 February 2005; also limited observations by Schweitzer in the 1980s). Succession may also be a threat in Virginia. Forestry, fire suppression, and gypsy moth spraying are threats to the unique Michigan populations of this complex.
Restoration Potential: Restoration potential is good, at least in its more suitable habitats. A restoration or recovery effort would include protecting these sites from the use of pesticides. Control of gypsy moths should be done using host specific biological methods. Habitat management may be required to insure a suitable amount of nectar and host plants within the community.
Preserve Selection and Design Considerations: This species requires shale barren habitat in the Appalachians. Shale barrens are semi-open shale slopes with sparse herbaceous vegetation and low density overstory of mainly red, scarlet, and black oak, scrub oak, and pine. Open areas created by shale slopes, log roads or power and gas rights-of-way must be in close proximity to more densely wooded areas. These dry, shale slopes should favor the natural growth of Canada cinquefoil as well as a variety of spring plants such as spring beauty (CLAYTONIA spp.), phlox (PHYOX SUBULATA) and birdsfoot violet (VIOLA PEDATA).
Size of an element site is uncertain. The species is quite mobile and may require 50 acres (20 ha) or more for population maintenance, with suitable breeding habitat scattered throughout.
Management Requirements: A natural area which meets the habitat requirements should maintain itself as a shale barren. Management of these natural areas would involve their protection against destruction from outside elements and from spraying. Grassy roads and right-of-ways can be managed by maintaining a suitable plant community structure with Canada cinquefoil as primary component. In Michigan, habitat for this species requires periodic burning in patches to maintain the open vegetation structure.
Management Programs: At the present time there are no known management programs for this taxon in either the Appalachians or in Michigan.
Monitoring Programs: Monitoring has taken place in recent years through annual surveys by Natural Heritage personnel in the Appalachian region from New York south to Virginia. In West Virginia, surveys have been conducted on historical sites to determine the presence or absence of the Grizzled Skipper, and its abundance.
Management Research Programs: There is no current research being conducted. A life history study was completed in West Virginia in 1988 (Allen, 1993).
Management Research Needs: A more intensive effort is needed to locate populations, and to provide immediate protection from gypsy moth spraying or other disturbances. Research efforts can then be concentrated on determining the size of suitable habitat required to maintain a viable population. An investigation of whether or not the numbers of host plants can be manipulated to benefit the skipper is also needed. The effects of precipitation and soil acidity on the growth of POTENTILLA CANADENSIS should be examined. The amount of POTENTILLA on historic sites in West Virginia has been declining over the last few years and may be partly responsible for declining populations.
Biological Research Needs: Need to determine if Michigan-Minnesota populations are the same taxon and also how these populations are effected by fire. Need to determine sensitivity to Btk, although this is probably very high. Feasibility of reintorductions especailly in West Virginia needs investigation.
Comments: A review of previous work done by Forbes (1974) and a review of museum specimens is needed to complete the taxonomic classification of the species. Electrophoretic studies may also help determine the taxonomic status of the Michigan and Appalachian populations.
Global Protection: Unknown whether any occurrences are appropriately protected and managed
Needs: Protect all EOs in New York if found again. Appalachian and long term populations in Pennsylvania need protection from spraying. Examine possible causes of host plant decline and ways to enhance growth of host plant. Michigan needs large barrens tract with several populations. Encourage listing under the US Endangered Species Act or pre-listing recovery by USFWS.
Relevance to Humans and Ecosystems
Stewardship Overview: Immediate management issues are maintaining the few habitats that still support this species and protecting them from gypsy moth spraying. Any past history of gypsy moth spraying was a reliable indication that this species would be absent even before its general collapse (Schweitzer, 1989). Gypsy Moth spraying is not compatible with continued existence of populations of this species. It is not known if BTK poses less risk than chemical biocides, but cirumstantial evidence suggests little difference with this species.
In some cases foodplant cinquefoils may need augmentation after droughts. A longer term goal is restoring metapopulation dynamics that almost certainly existed until the late 1980s by restoring population clusters that were sprayed out. Other possible issues could be controlling herbivory by deer affecting either the foodplant or nectar flowers, and eliminating exotic weeds. On right of ways broadcast herbiciding needs to be eliminated and the area maintained by either dormant season mowing or by careful selective ground applications of herbicides aimed at sprouts or shrubs. A few sites were relatively lightly grazed pastures. If any populations still exist in such situations presumably the status quo should be the goal. At a few sites in Virginia and perhaps elsewhere where the primary habitat is not a powerline, pasture, or shale barren, forest succession may be an issue. Details of how such habitat would be managed should be worked out by persons familiar with the sites. The possibility of using logging as a management tool should be considered. These recommendations are for typical P. wyandot.
For the Michigan entity, gypsy moth spraying is also a critical issue. Deer herbivory may be more of an issue with this entity since wild strawberry grows taller and may be more conspicuous to deer than prostrate cinquefoils. Otherwise management may be rather different since habitats are openings in or sparse pine, oak, or mixed barrens or woodlands which probably were originally maintained by occasional fires. However there is less occupied habitat now and as with most butterflies or skippers substantial survival within a burned area would be unlikely unless there were a lot of skips. Probably some level of prescribed burning is needed. If the population is using right of ways see recommendations above and consider whether it could be expanded by opening up adjacent woodlands.
Names and Taxonomy
Comments: Most authors have treated this as a subspecies of the circumboreal P. centaureae despite substantial differences. Genitalia differences noted by several authors (Shapiro, 1974; Klots, 1951; Forbes, 1960), adult maculation differences, as well as larval color (Allen, 1997), greatly different habitat, are cited by some recent works to justify treatment as a separate species as proposed by Shapiro (1974) and indirectly by Forbes (1960). Assuming the genitalia differences given explicitly by Forbes are reliable, these combined with both adult maculation and larval color differences would be more than enough to justify full species status by the normal practices in insect taxonomy. Genitalia differences alone are routinely used to justify species versus subspecies status. Forbes (1960), a notably conservative taxonomist with a very good record of getting species concepts right, left wyandot as a subspecies of centaureae but considered freija sufficiently different from it in genitalia as to warrant species status. It is difficult to conclude from Forbes' discussion of the genitalia and other differences that wyandot and freija are conspecific and apparently there is no disagreement now that freija and centaureae are conspecific. There has been no other relevant revisionary study since Lindsey, Bell and Williams (1930) who listed it under centaureae without comment and Forbes (1960) who compared wyandot only to freija and neither to European centaureae. Therefore, inertia favors subspecies status in the absence of indisputable evidence to the contrary-strong contrary evidence notwithstanding. Opler and Warren (2002) "tentatively" included wyandot within P. centaureae.
For now Pyrgus wyandot is retained in this database (as in previous versions) as a species distinct from Pyrgus centaureae freija largely on the basis of the genitalia differences between wyandot and freija, which as far as known hold reliably as reported and are supported by the other lesser differences. If the genitalia differences do not hold or mDNA proved very similar to other taxa, then the case for full species would be weakened considerably. Until a more careful published analysis is published, Forbes' account is accepted as the most recent taxonomic treatment and the universal treatment by subsequent authors of freija and centaureae as conspecific is also accepted. Also one would not expect a boreal-low arctic/alpine species in some of the hottest microhabitats in eastern North America--a fair description of a typical P. wyandot site in much of its former range. This is definitely a low elevation (rarely if ever >900 meters) barrens, not subalpine, taxon. The foodplant genera used in nature by this complex (Potentilla, Fragaria, Rubus) do not justify any particular taxonomic status. These genera are closely related and the exact species involved are all ground cover plants of similar growth form. The Q is mainly in deference to the tentative placement as a subspecies by the most recent published treatment (Opler and Warren, 2002). D. F. Schweitzer.
Whether as a full species or subspecies the circumscription of this taxon is also problematic until the Michigan entity is clarified. Michigan populations differ from typical wyandot in foodplant and habitat as well as adult appearance and larval color (pinkish brown, not green, Allen, 1997). It is very likely that Michigan populations are not the taxon wyandot. All other United States records for Pyrgus centaureae from Ohio south and east refer to P. wyandot.
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