Celastrina ladon - (Cramer, 1780)
Spring Azure
Other English Common Names: Dogwood Azure, Edwards' Azure
Taxonomic Status: Accepted
Related ITIS Name(s): Celastrina ladon (Cramer, 1780) (TSN 777893)
French Common Names: azur printanier
Unique Identifier: ELEMENT_GLOBAL.2.116204
Element Code: IILEPG0010
Informal Taxonomy: Animals, Invertebrates - Insects - Butterflies and Moths - Butterflies and Skippers
 
Kingdom Phylum Class Order Family Genus
Animalia Mandibulata Insecta Lepidoptera Lycaenidae Celastrina
Genus Size: C - Small genus (6-20 species)
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Concept Reference
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Concept Reference: Opler, P. A., and A. D. Warren. 2002. Butterflies of North America. 2. Scientific Names List for Butterfly Species of North America, north of Mexico. C.P. Gillette Museum of Arthropod Diversity, Department of Bioagricultural Sciences and Pest Management, Colorado State University, Fort Collins, Colorado. 79 pp.
Concept Reference Code: B02OPL01EHUS
Name Used in Concept Reference: Celastrina ladon
Taxonomic Comments: Until the 1980s or even by some books into the 2000s this name included several other species such as C. neglecta, C. neglectamajor, C. humulus, C. idella, C. lucia, C. echo etc. See Wright and Pavulaan (1999) for review of the nomenclature. The names pseudargiolus and violacea are synonyms of ladon. Further details by David Wright are in preparation. This is the familiar spring azure of most of the eastern United States except most of the Atlantic coastal plain, and along the Canadian border. Compared to most azures, C. ladon is well understood and uniform in appearance (except in New England see below), biology, and primary foodplant (flowering dogwood) over most of its range. Most populations of C. ladon occur as forms "violacea", many with "marginata" also common and the form "lucia" is usually completely absent or much less than 1% except at the extreme northeast. The male has the by now well-known "unique" scale morphology (see Wright and Pavulaan, 1999) and lacks androconia and is thus easily identified microscopically from spread specimens. This scale trait is almost completely diagnostic except that it also occurs in C. ebenina. Various western taxa (Opler and Warren, 2002) traditionally included in ladon do not share the unique scale morphology or otherwise resemble ladon and there is almost no chance any of them are conspecific with C. ladon. No subspecies of true C. ladon are recognized or anticipated in the future despite the names used in some popular quides, which if followed would place three or four "subspecies" sipping from the same patches of wet sand at the same time in New Jersey and elsewhere. C. ladon is not monophagous on dogwood (Cornus florida) and often uses Viburnum spp. and less often Ilex opaca. Taxonomic recognition is not warranted for such local foodplant strains.

The most common spring azure in northwestern New Jersey through Connecticut, Rhode Island and much of Massachusetts (see Klots,1951 PL 21: 6 and ?7) is an enigma. It has the male wing scales of C. ladon and C. ebenina but looks more like C. lucia and shares foodplants with both.. The "violacaea" form is at best rare, and the "lucia" patched form occurs at a non-trivial frequency with most individuals being of the "marginata" form. According to David Wright these azures are similar to C. lucia in allozymes, but this is not published and needs to be evaluated. Highbush blueberry (Vaccinium corymbosum) is the most commonly used foodplant, but oviposition has been observed (and the voucher collected) on Prunus serotina flowers near Boston. Viburnum spp. are used range-wide and apparently become the primary foodplant at the western end of the range in New Jersey. Viburnum species are also major foodplants for C. ladon. These southern New England azures are not known to use flowering dogwood (Cornus florida), the primary foodplant of C. ladon in much of its range, even in Connecticut where this tree was common well into the 1980s and still occurs. However, it is not known whether the larvae could mature on dogwood and it is possible they do use it. In New Jersey these "New England Azures" occur commonly in the relatively cold northwest of the state south into parts of Warren County. On the Piedmont of adjacent counties to the east and south normal C. ladon is the only member of this group. The "marginata" form occurs in both entities and individuals of this form are apparently inseparable by any characters. Normal New Jersey Piedmont C. ladon occurs more commonly as form "violacea" but there seems to be no other difference. D. Wright has informed Schweitzer that the relative frequencies of these forms can vary substantially from year to year in adjacent eastern Pennsylvania. Possibly the dark forms are induced by cold in C. ladon. At any rate a major shift in the frequency of the color forms alone, even assuming this is at least in part genetically based, does not justify treating these southern New England Azures as a separate species from C. ladon with which they share obviously derived scale morphology. It should be noted that otherwise C. ladon is not known to use blueberry, the main foodplant of more southern populations of C. lucia, as a foodplant anywhere. Schweitzer has sleeved two South Jersey C. ladon on V. corymbosum and obtained no eggs. C. lucia always oviposit freely on the same bushes. This difference does suggest some genetic differentiation. For now these northeastern populations seem best treated as a northeastern variant of C. ladon, with which they share a foodplant and from which "marginata" form specimens apparently cannot be separated. Alternate explanations are that that this entity is an intergrade between C. lucia and C. ladon, probably a relict of hybridization during the Pleistocene since these species remain separate in their modern contact areas; or that the Southern New England Azure is a fully separate species. The difficulty with the former explanation is that that rarely or never hybridize now. This summary by Dale Schweitzer.
Conservation Status
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NatureServe Status

Global Status: G4G5
Global Status Last Reviewed: 09Apr2010
Global Status Last Changed: 19Dec2008
Rounded Global Status: G4 - Apparently Secure
Reasons: While this is the common spring azure in much of the eastern United States, between dogwood (Cornus florida) mortality due to an exotic fungus, and loss of flowering viburnums to deer, this species is already declining in some parts of its range (David Wright), although it is not of immediate concern except locally in places with both high dogwood mortality and severe browsing of Viburnum by deer. For now this azure appears to be persisting where at least one of these plants remains common and reliably flowering, or in places where alternate foodplants, like holly, are used. Persistence depends on flowering foodplants every year, not just in most years. Given impacts from deer and the dogwood fungus, this butterfly really cannot be considered "demonstrably" secure in a substantial part of its range, but is is "apparently" globally secure. Ironically it might be most secure in the few places (e.g. southern New Jersey) where American holly is a foodplant since deer do not impact this tree. This azure should be monitored in places where the foodplants are being lost to fungus or their flower production is curtailed by deer.
Nation: United States
National Status: N4 (19Dec2008)
Nation: Canada
National Status: N5 (14Jun2016)

U.S. & Canada State/Province Status
United States Alabama (S5), Arizona (SNR), Arkansas (S5), California (SNR), Colorado (S5), Connecticut (S5), Delaware (S5), District of Columbia (SNR), Florida (SU), Georgia (S5), Idaho (SNR), Illinois (S5), Indiana (S5), Iowa (S5), Kansas (S5), Kentucky (S5), Louisiana (S5), Maryland (S5), Massachusetts (S5), Michigan (S5), Minnesota (S5), Mississippi (S5), Missouri (S5), Montana (S5), Nebraska (S5), Nevada (SNR), New Hampshire (S5), New Jersey (S5), New Mexico (SNR), New York (S5), North Carolina (S4S5), North Dakota (SNR), Ohio (S5), Oklahoma (S4), Oregon (SNR), Pennsylvania (S5), Rhode Island (S5), South Carolina (S5), South Dakota (SNR), Tennessee (S5), Texas (SNR), Utah (SNR), Virginia (S5), Washington (S5), West Virginia (S5), Wisconsin (S5), Wyoming (SNR)
Canada Alberta (SU), Labrador (S4), Manitoba (S5), New Brunswick (SNR), Newfoundland Island (SNR), Northwest Territories (SNR), Nova Scotia (S5), Nunavut (SNR), Ontario (SU), Prince Edward Island (S5), Quebec (S4S5), Saskatchewan (S5)

Other Statuses

NatureServe Global Conservation Status Factors

Range Extent: >2,500,000 square km (greater than 1,000,000 square miles)
Range Extent Comments: If the New England Azure is included, the range is approximately Massachusetts west through southern New York, extreme southern Ontario, southern Michigan, and presumably to Wisconisn or Minnesota south certainly to Arkansas and presumably into Texas and definitely Georgia, probably also the Florida panhandle. This species is absent from much of the coastal plain including the core of the New Jersey Pine Barrens, and from some higher ridgetops and probably the colder parts of the Poconos. This is the common spring azure of mainly deciduous forests of the Piedmont, inner coastal plain, lower elevations in the southeastern mountains, and most of the Midwest. Inclusion of populations in New England to northwest New Jersey is tentative. See Taxonomy Comments. No western populations appear to be conspecific based on their appearance and scale morphology.

Number of Occurrences: 81 to >300

Population Size: 10,000 to >1,000,000 individuals

Overall Threat Impact: High
Overall Threat Impact Comments: The loss or reduction of a major foodplant, healthy flowering dogwood (Cornus florida), to an exotic fungus is a threat to this species in some parts of the range and may become so in most of the range. This threat is compounded now in many places where deer have greatly reduced or eliminated the usual alternate foodplants, viburnum flowers (although not necessarily actually killed the plants). As far as known Celastrina ladon should persist where one of the foodplants remains common and flowers successfully every year. This butterfly is declining substantially in the Appalachians (David Wright) and may become extirpated from some counties. It is not presently threatened in most of its range but the dogwood blight could spread. Ironically this azure is apparently stable along Delaware Bay and on the Cape May peninsula in southern New Jersey, one of few areas in its range where it was historically uncommon and local. American holly, which deer do not normally eat, is an important alternate foodplant (along with dogwood) there. Other than dogwood blight fungus and excessive deer browsing, there are no known large-scale threats, although invasive shrubs and vines might also be impacting foodplants.

Short-term Trend: Decline of 10-30%

Long-term Trend: Decline of <30% to increase of 25%

Other NatureServe Conservation Status Information

Inventory Needs: Need to determine identity and limits of sibling species or subspecies.

Distribution
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Global Range: (>2,500,000 square km (greater than 1,000,000 square miles)) If the New England Azure is included, the range is approximately Massachusetts west through southern New York, extreme southern Ontario, southern Michigan, and presumably to Wisconisn or Minnesota south certainly to Arkansas and presumably into Texas and definitely Georgia, probably also the Florida panhandle. This species is absent from much of the coastal plain including the core of the New Jersey Pine Barrens, and from some higher ridgetops and probably the colder parts of the Poconos. This is the common spring azure of mainly deciduous forests of the Piedmont, inner coastal plain, lower elevations in the southeastern mountains, and most of the Midwest. Inclusion of populations in New England to northwest New Jersey is tentative. See Taxonomy Comments. No western populations appear to be conspecific based on their appearance and scale morphology.

U.S. States and Canadian Provinces
Color legend for Distribution Map
Endemism: occurs (regularly, as a native taxon) in multiple nations

U.S. & Canada State/Province Distribution
United States AL, AR, AZ, CA, CO, CT, DC, DE, FL, GA, IA, ID, IL, IN, KS, KY, LA, MA, MD, MI, MN, MO, MS, MT, NC, ND, NE, NH, NJ, NM, NV, NY, OH, OK, OR, PA, RI, SC, SD, TN, TX, UT, VA, WA, WI, WV, WY
Canada AB, LB, MB, NB, NF, NS, NT, NU, ON, PE, QC, SK

Range Map
No map available.


U.S. Distribution by County Help
State County Name (FIPS Code)
FL Okaloosa (12091), Walton (12131), Washington (12133)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
03 St. Andrew-St. Joseph Bays (03140101)+, Choctawhatchee Bay (03140102)+, Yellow (03140103)+
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Basic Description: Butterfly, Lycaenidae.
Non-Migrant: N
Locally Migrant: N
Long Distance Migrant: N
Palustrine Habitat(s): FORESTED WETLAND
Terrestrial Habitat(s): Forest - Hardwood, Forest - Mixed, Woodland - Hardwood, Woodland - Mixed
Habitat Comments: This is mostly a forest butterfly that is typically seen along paths or on flowering shrubs often well into the woods.. In most of the range any sort of deciduous or mixed deciduous-pine forest with the locally utilized foodplants in the understory or subcanopy is habitat. Adults may occur along edges, but they do not venture more than a few meters into fields or open habitats and they are rarely seen in yards or gardens.
Food Comments: The primary larval foodplant is usually flowering dogwood (Cornus florida), but species of Viburnum are also very widely used. In southern New Jersey American holly (IIlex opaca) is apparently a significant foodplant, although this azure is not often found there in the absence of dogwood. Populations from northwestern New Jersey, southeastern New York, Connecticut, Rhode Island, and Massachusetts that are tentatively included in this species feed on highbush blueberry (Vaccinium corymbosum) and Viburnum spp., apparently both if they are available, and less often on Prunus serotina flowers. With all foodplants, larvae feed mostly on the flowers.



Adult Phenology: Diurnal
Immature Phenology: Hibernates/aestivates
Phenology Comments: There is always only one brood in spring, mostly in April or early May in much of the range. Where they overlap. C. ladon starts later than C. lucia (sometimes more than a month later in southern New Jersey), about the same time as C. idella, and earlier than C. serotina. However any of these that are locally sympatric will definitely overlap in phenology. The larval stage takes about a month, and except in the northeast corner of the range, most or all larvae probably complete feeding in May or very early June. All populations have one generation annually and spend most of the year as pupae.
Economic Attributes Not yet assessed
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Management Summary Not yet assessed
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Population/Occurrence Delineation
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Group Name: Celastrina Species (Azures)

Use Class: Not applicable
Minimum Criteria for an Occurrence: A location, generally a woodland or forest, where the species occurs, or has occurred, where there is potential for persistence or continued recurrence. Minimally a suitable habitat with the larval foodplant where the species has been verified by specimens or photographs. In some situations sight records will suffice but these are not recommended except for summer records for C. NEGLECTA and should be used only with the approval of an expert highly familiar with the local fauna. Not all specimens and certainly not all photographs are presently reliably identifiable for some taxa in some places, so in some cases a series may be needed.
Mapping Guidance: Occurrence boundaries will normally closely match the limits of the foodplants except in summer for C. NEGLECTA. This is especially helpful for taxa like C. NIGRA, NEGLECTAMAJOR and HUMULUS that feed on understory plants that are somewhat to highly localized. With taxa feeding on co-dominant subcanopy or shrub layer plants like flowering dogwood, holly, highbush blueberries, viburnums etc. boundaries may be very hard to define and thus arbitrary. Consult habitat and foodplant comments fields for species-specific information on what constitutes suitable habitat when mapping occurrences for individual species.
Multiple colonies within a single patch of forest should nearly always be treated as a metapopulation.

Separation Barriers: There are no actual data, but the forest and woodland species seem unusually reluctant to fly more than about 15 meters into open habitats. So there may be cases where croplands, paved areas etc. should be treated as barriers and not merely unsuitable. Two lane roads are definitely not barriers if forest occurs on both sides.
Separation Distance for Unsuitable Habitat: 2 km
Separation Distance for Suitable Habitat: 10 km
Alternate Separation Procedure: Within continuous forest use the suitable habitat distance unless there are gaps with no foodplant of at least half the suitable habitat distance.
Separation Justification: No real data are known. However on warm sunny days the spring woodland taxa such as LADON, LUCIA, IDELLA are easily observed flying widely throughout any available habitat as long as their common foodplants are present at all in the subcanopy or shrub layer. Under cooler conditions they concentrate in protected sunny areas and can give the false impression of small localized occurrences or that they stay along paths. In places like southern New Jersey and some parts of the Appalachians these azures are so ubiquitous in the woods that EOs are virtually undefinable. One can easily directly observe that occurrences cover several square kilometers if the foodplants do. As far as known the species such as NEGLECTAMAJOR and NIGRA that feed on more localized forest understory plants stay within a kilometer and usually much less of them. The woodland and forest species almost never move even 100 meters into open fields or residential areas, although they do move across powerlines and roads with woods on both sides. For example Schweitzer has never seen C. IDELLA or C. LADON LUCIA in his yard in 13 years despite presence of the foodplant for both within less than a kme and some of the best occurrences globally of the former within about five kilometers. Probably larger distances are appropriate for C. NEGLECTA which readily enters and breeds in towns and will cross larger fields, but for now no separate Specs are suggested for that very common species. Instead be more liberal in applying suitable habitat distance with this species. It is less clear what Specs should be for species 2 (cherry gall azure) but it seems to occur along forest edges and more widely in open woodlands such as barrens or outcrops. For now use the generic Specs
Inferred Minimum Extent of Habitat Use (when actual extent is unknown): 1 km
Inferred Minimum Extent Justification: Some occurrences of C. NEGLECTA, LADON LADON, C. LADON LUCIA, and in New Jersey also IDELLA, can easily be observed to extend over several kilometers in at least one direction. At least in Schweitzer's experience, occurrences under 50-100 hectares are unusual but occurrences of 200-500 hectares are rather routine for these four taxa. However some arbitrary limit is needed in places where the habitat and foodplant are contiguous for large distances. This figure is recommended with large habitats until additional sampling establishes the correct (generally larger) extent. With smaller habitats (up to 400 hectares) assume full occupancy.
Date: 25Jul2001
Author: Schweitzer, D. 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: 19Dec2008
NatureServe Conservation Status Factors Author: Schweitzer, D. F.
Element Ecology & Life History Edition Date: 27Jan2009
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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  • Layberry, R.A., P.W. Hall, and J.D. LaFontaine. 1998. The Butterflies of Canada. University of Toronto Press: Toronto, Canada. 280 pp. + color plates.

  • Opler, P. A., and A. D. Warren. 2002. Butterflies of North America. 2. Scientific Names List for Butterfly Species of North America, north of Mexico. C.P. Gillette Museum of Arthropod Diversity, Department of Bioagricultural Sciences and Pest Management, Colorado State University, Fort Collins, Colorado. 79 pp.

  • Pelham, J. P. 2008. A catalogue of the butterflies of the United States and Canada with a complete bibliography of the descriptive and systematic literature. The Journal of Research on the Lepidoptera. Volume 40. 658 pp.

  • Wright, D.M., and H. Pavulaan. 1999. Celastrina idella (Lycaenidae, Polyommatinae) a new butterfly species from the Atlantic coastal plain. The Taxonomic Report of the International Lepidoptera Survey 1(9): 11 pp.

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