Papaipema circumlucens - (Smith, 1899)
Hop Borer
Taxonomic Status: Accepted
Unique Identifier: ELEMENT_GLOBAL.2.117438
Element Code: IILEYC0280
Informal Taxonomy: Animals, Invertebrates - Insects - Butterflies and Moths - Papaipema Moths
 
Kingdom Phylum Class Order Family Genus
Animalia Mandibulata Insecta Lepidoptera Noctuidae Papaipema
Genus Size: D - Medium to large genus (21+ species)
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Concept Reference
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Concept Reference: Hodges, R.W. et al., eds. 1983. Check List of the Lepidoptera of America North of Mexico. E.W. Classey Limited and The Wedge Entomological Research Foundation, London. 284 pp.
Concept Reference Code: B83HOD01HQUS
Name Used in Concept Reference: Papaipema circumlucens
Taxonomic Comments: The name circumlucens has been applied to other species in older literature. Ths species it is now applied to has also been called P. humuli and P. ochroptena.
Conservation Status
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NatureServe Status

Global Status: G3G4
Global Status Last Reviewed: 16Feb2010
Global Status Last Changed: 03Sep2004
Rounded Global Status: G3 - Vulnerable
Reasons: This moth has declined or disappearance east of Ohio with the last eastern collections probably in the 1970s in Litchfield Co., Connecticut. P. circumlucens is not considered common in many, if any, parts of the range but there is insufficient information to really assess current status. The species is not imminently imperiled or highly rare in parts of the Midwest, but it is historic, and probably extirpated, from a substantial part of its range to the east, and it is not clear this species is secure any place. A major unknown is whether or not the non-native and locally invasive Japanese hops will prove to be a foodplant, especially eastward where native Humulus taxa are rather rare. A combination rank of G3G4 is arrived at by process of elimination, without attempting to factor in impacts (neutral, positive or negative) from cultivated or invasive strains of hops. This moth is generally uncommon to rare but probably not really imperiled westward.
Nation: United States
National Status: N3N4 (03Sep2004)
Nation: Canada
National Status: NU (17Aug2017)

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 Arizona (SNR), Colorado (SNR), Connecticut (SH), Delaware (SH), Illinois (SNR), Michigan (SNR), Missouri (SNR), New Jersey (SH), New Mexico (SNR), Ohio (SNR), Wisconsin (SNR)
Canada Manitoba (SU), Saskatchewan (S3)

Other Statuses

NatureServe Global Conservation Status Factors

Range Extent: 20,000-2,500,000 square km (about 8000-1,000,000 square miles)
Range Extent Comments: Historically a relatively narrow band from Connecticut to Wisconsin, and populations now considered to be be this species (as P. ochroptena) occur in Colorado and probably other Rocky Mountain states. Southern limit in the east either now or historically is unknown but it reached at least northern Delaware and southern Ohio. Reportedly uncommon but documented from five southern counties in Ohio (but as of Rings eta l. (1992) the last record was in 1986). Fairly widespread in Wisconsin. No specimens from Kentucky or elsewhere south of Ohio in the east. Historic in at least Connecticut, New York, Pennsylvania, New Jersey and Delaware.

Number of Occurrences: Unknown

Population Size: Unknown

Number of Occurrences with Good Viability/Integrity: Unknown

Overall Threat Impact: High
Overall Threat Impact Comments: Threats include development or other modifications of habitat, habitat fragmentation, and in some places potentially exotic plants, especially on floodplains. It is uncertain to what extent deer browse hops, but most likely high deer populations are a threat in many places. For example, native hops are seldom seen any more in the Northeast with deer a likely explanation. Probably a major problem is fragmentation such that local extirpations cannot be balanced by local colonization. On the other hand if this species adapts to Japanese hops as a foodplant in may start increasing as this invasive becomes more common.

Short-term Trend: Unknown
Short-term Trend Comments: It is not clear that this species is still declining. Status in and west of Ohi not well known.

Long-term Trend: Decline of 50-90%
Long-term Trend Comments: Seems likely to have diappeared or become very much rarer east of Ohio and obviously has lost much to most habitat even in regions where it still occurs.

Environmental Specificity: Unknown
Environmental Specificity Comments: It is quite possible but not assumed that other factors besides sufficient foodplant are important to this species.

Other NatureServe Conservation Status Information

Distribution
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Global Range: (20,000-2,500,000 square km (about 8000-1,000,000 square miles)) Historically a relatively narrow band from Connecticut to Wisconsin, and populations now considered to be be this species (as P. ochroptena) occur in Colorado and probably other Rocky Mountain states. Southern limit in the east either now or historically is unknown but it reached at least northern Delaware and southern Ohio. Reportedly uncommon but documented from five southern counties in Ohio (but as of Rings eta l. (1992) the last record was in 1986). Fairly widespread in Wisconsin. No specimens from Kentucky or elsewhere south of Ohio in the east. Historic in at least Connecticut, New York, Pennsylvania, New Jersey and Delaware.

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
Endemism: occurs (regularly, as a native taxon) in multiple nations

U.S. & Canada State/Province Distribution
United States AZ, CO, CT, DE, IL, MI, MO, NJ, NM, OH, WI
Canada MB, SK

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
CT Litchfield (09005)*
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
01 Housatonic (01100005)+*
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Non-Migrant: N
Locally Migrant: N
Long Distance Migrant: N
Palustrine Habitat(s): FORESTED WETLAND, Riparian
Terrestrial Habitat(s): Forest - Hardwood, Forest - Mixed
Habitat Comments: As far as known, potentially any habitat where hops occur frequently over a substantial area. Probably most often forested floodplains but not exclusively so.
Food Comments: As far as known the larvae bore only in hops. It is unclear to what extent native versus exotic taxa of hops (Humulus) are used. Humulus lupulus var. lupulus is considered introduced from cultivation. Except at the extreme edges (Colorado, New Jersey, western Connecticut), the range of the moth seems to fit that given in the USDA Plants Profiles for the native H. lupulus. var. pubescens, but the native variety lupuloides also occupies the entire range of the moth and more. Unlike Hydreacia immanis, P. circumlucens was apparently not much of a pest on cultivated hops in the Northeast. It is unknown whether Humulus japonicus, which is becoming locally invasive, will prove to be a foodplant.
Phenology Comments: See Hessel (1954) and Rings (1993). Adults occur earlier than most Papaipema, often before September. Eggs overwinter in this genus and larvae occur in late spring and summer.
Economic Attributes Not yet assessed
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Management Summary
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Biological Research Needs: It would be very useful to learn whether P. circumlucens larvae will occur in Humulus japonicus, which is becoming invasive in some places, notably northern Delaware. It is also worth noting that USDA Plants Profile for common hops indicates three varieties as native in the USA and Canada with one of these being western. Humulus lupulus var. lupulus is considered introduced. Precise foodplant information is needed, this moth possibly require native hops as larval foodplants. Except at the extreme edges (Colorado, New Jersey, western Connecticut), the range of the moth seems to fit that given in the USDA Plants Profiles for H. lupulus. var. pubescens, but variety lupuloides occupies the entire range and more. Unlike Hydreacia immanis, P. circumlucens was apparently not much of a pest on cultivated hops in the Northeast.
Population/Occurrence Delineation
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Group Name: Forest Papaipema

Use Class: Not applicable
Minimum Criteria for an Occurrence: A location supporting a breeding population. Minimally a specimen (exceptionally for some species a photograph or larval burrows in the foodplant) in association with plausible habitat including the foodplant. In most cases a photograph of an individual sitting near a collecting light will not be identifiable to species, especially from and JPEGS due to color distortion, although use of natural light does help. A genuine expert must approve all identifications based on less than an actual specimen and some specimens are difficult and require expert identification. Generally identification of Papaipema feeding damage (burrows, frass, borings) to genus is rather easy but in many cases these cannot be identified to species because more than one species could occur in that plant species. In a few cases location an symptoms on the plant (Hessel, 1954) will distinguish among possible species, e.g. P. nebris and P. maritima in the same sunflowers. Sometimes larval specimens can be identified to species. Collections of an adult not associated with habitats containing the foodplant are not EOs.




Separation Distance for Unsuitable Habitat: 2 km
Separation Distance for Suitable Habitat: 5 km
Date: 01Feb2007
Author: Schweitzer, D.F.

Use Class: Not applicable
Minimum Criteria for an Occurrence: A location supporting a breeding population. Minimally a specimen (exceptionally for some species a photograph or larval burrows in the foodplant) in association with plausible habitat including the foodplant. In mmost cases a photograph of an individual sitting near a collecting light will not be identifiable to species, especially from and JPEGS due to color distortion, although use of natural light does help. A genuine expert must approve all identifications based on less than an actual specimen and some specimens are difficult and require expert identification. Generally identification of Papaipema feeding damage (burrows, frass, borings) to genus is rather easy but in many cases these cannot be identified to species because more than one species could occur in that plant species. In a few cases location an symptoms on the plant (Hessel, 1954) will distinguish among possible species, e.g. P. nebris and P. maritima in the same sunflowers. Sometimes larval specimens can be identified to species. Collections of an adult not associated with habitats containing the foodplant are not EOs.

Mapping Guidance: The essential resource is the larval foodplant so in general EOs consist of foodplant patches and to some extent intervening connecting space, for example one would probably map an entire small bog, and not just the pitcher plant patches, as the EO for northern populations of P. APPASIONATA. Within the overall occurrence it may be advisable to map major foodplant patches carefully so that managers will know their exact locations. See habitat and food comments fields for species-specific information on what constitutes habitat when mapping occurrences.
Separation Barriers: None are really known and it is suspected there are none in practice, although nocturnal lighting could become a barrier in extreme cases. Some species at least do occur in fairly brightly lit residential areas.
Separation Distance for Unsuitable Habitat: 2 km
Separation Distance for Suitable Habitat: 5 km
Alternate Separation Procedure: In general when multiple colonies occur in a distinct natural community occurrence, such as a prairie remnant or wetland complex, they should usually regarded as one metapopulation occurrence and so the suitable habitat distance should be used within the general habitat type(s) (forest, marsh, prairie) occupied by the species in that general area even if the foodplants are patchy. Do not do this across major vegetational changes such as prairie to forest unless both types actually sometimes support the foodplant and its borers. Also if the foodplants are highly patchy and the overall habitat not clearcut-consider compromise distances in marginal habitat but not less than twice the unsuitable distance. For widespread forest understory species, even occasionally the rarer ones like P. duplicata, large occurrences still exist and the Specs for "Forest, Woodland and Scrub Noctuidae" are justified.
Separation Justification: Females of Papaipema, and at least some related genera (e.g. Bellura spp., and Spartiniphaga carterae) are occasionally collected two or more kilometers from any potential larval habitat, but for the most part these are sedentary moths that are usually found within 10 meters of foodplant patches. Females apparently are more dispersive than males and tend to do so after laying some eggs at the natal site. So two kilometers while arbitrary seems adequate. Suitable habitat distance is shorter than for most Noctuidae because habitats and populations of these borers tend to be small, and situations do occur where patches a few km from known colonies remain unoccupied for no obvious reason. Sometimes, but not often, some foodplant is unsuitable due to edaphic or other conditions.

P. pterisii, P. sp. 1, P. frigida, P. furcata, P. inquaesita, and S. carterae can have large occurrences where they are ubiquitous over several hundred hectares. In most cases suitable foodplant patches are occupied at least in some years but in any given year some may not support mature larvae. Foodplant patches are often unoccupied or nearly so by immatures for the first season after a fall, winter or spring burn since by far most are killed. However, the same patch may be very good habitat a year later and until the next burn if there are unburned refugia adjacent (see e.g. Panzer, 1998). Similarly for some species such as P. sulphurata and P. stenocelis some, many, or all habitats may be suitable only in certain years depending on water levels and most or all patches are part of the metapopulation EO regardless of occupancy in a given season. In general then extensive suitable habitat will probably be occupied at least over several years if not every year, but it does seem prudent to consider collections more than five kilometers apart as separate occurrences pending more information especially given that patch sizes and therefore deme size can be quite small.

Inferred Minimum Extent of Habitat Use (when actual extent is unknown): 1 km
Inferred Minimum Extent Justification: Use this radius only with virtually contiguous habitats with the foodplants widespread. Some known occurrences for P. pterisii, P. sp. 1, and S. carterae are several square kilometers. In by far most cases the inferred extent is the entire contiguous habitat which will usually be a few hundred hectares or less and if the habitat is under 400 hectares assume full occupancy (at least over time if not every year). Note also for riparian habitats this distance is not really a radius but more of a linear distance.
Date: 12Sep2001
Author: Schweitzer, Dale F.
Population/Occurrence Viability
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Group Name: Forest Papaipema

Excellent Viability: While original habitats may sometimes have been fairly contiguous for dozens or hundreds of miles in heavily forested regions like the piedmont hills, Ohio Valley, and Appalachians, it seems reasonable to consider A-rank for apparent metapopulations in 2,000 hectares of more or less of contiguous forest with the foodplant scattered commonly throughout such that patches are less than the separation distances apart at an average density of five substantial patches of vigorous foodplants per hectare. Note that small plants with lower stem diameters much less than that of a pencil are unsuitable for most species, but are good enough for root borers like P. rutila and probably the fern feeders. Foodplant patches may have some exotics mixed in but not to the extent that foodplant vigor is being obviously impacted. The foodplants also must not experience substantial herbivory from deer affecting stems. Such occurrences are probably reasonable approximations at least of former much better than average conditions and seemingly suitable habitats do exist for some of the species. An A or B ranked occurrence is most likely to follow some edaphic situation such as the lower slopes of a long ridge, a valley, or a flood plain and probably will have small gaps without the foodplant. For species that feed on understory herbs such occurrences are most likely to exist in mesic or wetter second growth forest on lands that have not been previously plowed or at least were reforested in the early enough in the twentieth century that native flora moved in before the current mix of exotics. Within reason habitat can be considered to be wooded portions of the least polygon along appropriate topography (e.g. valley, stream corridor, low slopes, ridgetop) that would enclose all major foodplant patches, especially if some plants occur between these patches.

Unlike most Lepidoptera, these borers may occupy substantially less than 100% of available habitat, and it is often unclear whether patch occupancy varies from year to year. So A-rank should not be assigned merely on the basis of abundant foodplant over a large area. Some effort is needed to verify that the Papaipema actually does occur widely and in abundance. AC can be used for extensive habitats with minimal actual observations, or AB if there is a at least enough sampling to show that the population is not highly localized. If A rather than AB or AC is to be applied there should be some evidence that the species is well distributed within the apparent habitat at sufficient density that 500 or more adults are probably produced per year. Several last instar larval burrows per hectare would be sufficient evidence as would several adults per blacklight at three widely separated sample sites. A-rank might also be reasonable for smaller, but still substantial, relatively pristine tracts where exceptionally high numbers of adults are found, for example sites in Virginia where dozens of P. duplicata were collected on each of two nights at a blacklight. Similarly large numbers at lights in extensive habitat suggest some likely A-ranked occurrences for species 1 in the Poconos. However since habitat size and probably microhabitat diversity, are also important in the long term, A-rank is never recommended on the basis of large numbers alone. A minimum of 500 hectares is suggested, but this area need not be strictly contiguous. Habitat size recommendations do not take threats into account directly. Consider potential lethal events such as periodic extreme herbivory in spring or wildfire, among others. Apparently inundation for a few days duration is not particularly lethal to any early stages, but flood scour could be if foodplants are severely damaged or buried. A-rank should also be strongly considered for metapopulations containing two demes worthy of B-rank and at least three more worthy of C, or with three demes that would rank as solid Bs on their own.

Good Viability: These are substantial occurrences that have good to excellent prospects for persistence in their current condition, but that cannot be regarded as among the best historic occurrences. Generally there should be some evidence of at least hundreds of adults occupying hundreds of hectares of good habitat. Consider B or BC-rank for apparent metapopulations in more or less of contiguous forest with the foodplant scattered commonly throughout at least 500 hectares such that patches are less than the separation distances apart at an average density of two substantial patches or 50 stems of vigorous foodplants per hectare. Note that small plants with lower stem diameters much less than that of a pencil are unsuitable for most species, but are good enough for root borers like P. rutila and probably the fern feeders. Foodplant patches may have some exotics mixed in but not to the extent that foodplant vigor is being obviously impacted. The foodplants also must not experience substantial herbivory from deer affecting stems. Observations of adults (especially if more than one per site per night) at blacklights or of several larvae per hectare at several sample sites in 500 hectares of habitat would be reasonably sufficient evidence. Within reason habitat can be considered to be wooded portions of the least polygon along appropriate topography (e.g. valley, stream corridor, low slopes, ridgetop) that would enclose all major foodplant patches, especially if some plants occur between these patches. However, unlike most Lepidoptera, these borers may occupy substantially less than 100% of available habitat, and it is often unclear whether patch occupancy varies from year to year. So B-rank should not be assigned merely on the basis of abundant foodplant over a large area. Some effort is needed to verify that the Papaipema actually does occur widely and that the occurrence regularly produces a few hundred adults. Several last instar larval burrows per hectare would be sufficient evidence as would several adults per blacklight at three widely separated sample sites. BC should be used where observations are minimal but habitat is extensive. B-rank also seems appropriate where number of adults or larvae, that is density, seems exceptionally high but habitat is not extensive. A minimum of 100 hectares is suggested, but consider the potential for lethal events such as periodic extreme herbivory (usuully by deer) while larvae are in the portion being consumed (likely in spring), wildfire, among others. Apparently inundation for a few days duration is not particularly lethal to any early stages, but flood scour could be if foodplants are severely damaged or buried.

B-rank should also be strongly considered for metapopulations containing at least three demes worthy of C if these would add up to minimal B-rank criteria for probable population size and the least polygon enclosing all of them would contain 100 hectares, or even with many very small demes that would meet these criteria.

Fair Viability: These are more or less typical isolated occurrences by current standards that might persist for at least a few decades given current conditions. Often these will be on a few hectares with somewhat isolated large patches of the foodplant in landscapes fragmented by deforestation, invasive understory shrubs and vines or out of control deer. Generally there should be some evidence suggestive of a few dozen adults most years on at least five hectares of habitat, although occasional bad years with 10-20 adults should not disqualify an occurrence for a C. Collection records over more than a ten year period, even two eleven years apart, would be strong evidence in support of C (rather than D) if the habitat extent and quality has not obviously declined in the last 20 years since that would suggest current, rather than past, conditions are adequate for persistence. Generally there should be at least five hectares of habitat as a buffer against stochastic events, human disturbance or intense localized predation in some places some years--which tends to vary from season to season. Consider CD rather than C if the foodplants are confined an area of less than 10 hectares and there is no actual evidence of persistence. Higher density and smaller habitat size could also qualify if the habitats are not prone to severe disturbances or (e.g. flypoison) deer avoid eating the foodplant and there is some evidence of persistence at comparable extent and quality. Localized extirpations occur with Papaipema for many reasons (Hessel, 1954, Decker, 1934, Schweitzer personal observations) and many habitat are unoccupied in fragmented landscapes. However females, and probably less so males, definitely disperse based on light trap collections out of habitat. Such females do find unoccupied foodplants--Schweitzer has observed transient colonies of at least four species in apparently isolated patches occupying less than 20-100 square meters. If recolonization has been observed before or seems quite likely given overall landscape conditions, an occasional year when no adults are produced should not cause an occurrence to be dismissed as of no conservation concern (D). Foodplant stands that commonly produce dozens of adults but occasionally produce none could be regionally significant. Assessing this requires some knowledge of the overall landscape, but generally if a recurrent, but nonpersistent, occurrence often produces 20-50 adults, consider C or CD based on landscape context. An isolated occurrence that often falls below 20 adults should not be ranked higher than CD unless there is convincing evidence that it has survived more than 20 years in that condition, that is that it is not a remnant of a recently larger occurrence. Also consider whether there are enough such colonies in the vicinity that they should be pooled as a metapopulation (probably of C-rank) for ranking purposes.

Poor Viability: These are occurrences that are not likely to persist and are unlikely to be recolonized once extirpation occurs, including transient colonies that cannot be reasonably pooled with other colonies as a metapopulation. Populations that occupy less than a hectare should be ranked D unless strong mitigating factors suggest CD, and all populations that are usually less than 20 adults per year should be ranked D.

Justification: Quantification of parameters is impossible, but it is known that many to most small habitats are unoccupied at least east of western Virginia and Ohio, and that most of these species have become uncommon to historic over large areas in the Northeast, obviously reflecting at least some combination of habitat fragmentation and reduction of foodplant vigor (and often abundance) by out of control deer and invasive exotic shrubs and vines and perhaps even herbs like garlic mustard. It is unlikely these are the only factors. In particular egg predation by exotic earthworms is a suspected, but unsubstantiated, explanation for some absences in Connecticut and Delaware, where the habitat is still substantial and in the latter area historic presence is documented (Jones, 1928) and foodplants are still in good condition on some protected lands. It appears most of these species have become scarcer since the 1970s, the fern feeders and flypoison borer being apparent exceptions (which very strongly suggests deer as a major factor). EORANKSPECs attempt to define criteria that would give an occurrence some chance of persisting (C) and to separate out possible high quality occurrences (BC or higher). Many observers have witnessed local extirpations of small colonies, often for the reasons suggested by Hessel but sometimes without apparent cause, but clusters of individually small colonies do commonly persist. The A and B ranks include criteria that should allow such populations to persist for the long term and to maintain their current quality including genetic variability. They are more or less based on an assumption of one adult or five patches of foodplant per hectare. However there is no general agreement on what minimum numbers would be required so 200 adults per year (about the lowest such suggested number) is used. Realistically occurrences that might rank higher than C will probably not be completely isolated and so will receive some gene flow. Since full habitat occupancy cannot be assumed, A and B ranks should be based on some real documentation beyond mere presence, but BC or AC would be appropriate based only on extensive habitat extent and abundant foodplants. While these criteria are somewhat arbitrary they should result in the better extant occurrences being separated out as at least BC rank even without a lot of information. The ranks A, AB, and B probably cannot often be used because there will not usually be enough information--that is one collection of adults does not establish extent of occupied habitat if the foodplant is widespread in the area. However it is not difficult to establish the criteria suggested, especially based on larvae, but several well placed blacklights would also usually suffice. If this is not done, it is still probably reasonable to assume that presence plus extensive habitat indicates at least a C, so ranks like BC etc. are probably justified. However with poor knowledge of threats, it is possible that an on-going decline could invalidate this, some monitoring is strongly suggestedFor now though the default assumption with documented presence and substantial habitat is that the occurrence is not a D. Only time will tell, and for now C ranked occurrences need to be protected to the extent feasible.

Key for Ranking Species Element Occurrences Using the Generic Approach (2008).
Date: 01Feb2007
Author: Schweitzer, D.F.
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: 16Feb2010
NatureServe Conservation Status Factors Author: Schweitzer, D.F.
Element Ecology & Life History Edition Date: 16Feb2010
Element Ecology & Life History Author(s): Schweitzer, Dale F.

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

References
Help
  • Covell, C. V., Jr. 1999. The butterflies and moths (Lepidoptera) of Kentucky: An annotated checklist. Kentucky State Nature Preserves Commission, Scientific and Technical Series Number 6, Frankfort, Kentucky. 220 pp.

  • Ferge, L. A., and G. J. Balogh. 2000. Checklist of Wisconsin Moths (Superfamilies Drepanoidea, Geometroidea, Mimmallonoidea, Bombycoidea, Sphingoidea, and Noctuiodea). Contributions in Biology and Geology of the Milwaukee Public Museum No. 93. Milwaukee, Wisconsin. 55 pp. and one color plate.

  • General Status, Environment Canada. 2014. Manitoba moth species list and ranks as recommended by expert.

  • Hessel, Sidney, A. 1954. A guide to collecting the plant-boring larvae of the genus Papaipema (Noctuidae). Lepid. News. 8: pp. 57-63. Note a corrected version is included as Appendix D of the Lepidopterists' Society's basic techniques manual (Winter, 2000).

  • Hodges, R.W. et al., eds. 1983. Check List of the Lepidoptera of America North of Mexico. E.W. Classey Limited and The Wedge Entomological Research Foundation, London. 284 pp.

  • Lafontaine, J.D. and B. C. Schmidt. 2010. Annotated check list of the Noctuoidea (Insecta, Lepidoptera) of North America north of Mexico. ZooKeys 40:1-239.

  • Rings, R. W., E. H. Metzler, F. J. Arnold, and D. H. Harris. 1992. The Owlet Moths of Ohio (Order Lepidoptera, family Noctuidae). Ohio Biol. Surv. Bull. New Series, Vol. 9, no. 2, vi. + 219 pp., 16 color plates.

  • Schweitzer, D.F. 1999. Papaipema moths with emphasis on prairie species. Element Management record, Current version date: 1999-12-01. IN Natureserve 2004. NatureServe Central Databases. Arlington, Virginia. U.S.A.

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