Celastrina serotina (Lycaenidae: Polyommatinae): a New Butterfly Species from the Northeastern United States and Eastern Canada.

Abstract

A new light-venter Azure species, Celastrina serotina, is described from the northeastern United States and eastern Canada. The flight period of this univoltine spring species is interpolated (allochrony) between its sympatric congeners, C. lucia Auctorum (not Kirby) and C. neglecta (Edwards). In this regard it is similar to C. neglectamajor Opler & Krizek in the southern Appalachians and C. idella Wright & Pavulaan on the New Jersey coastal plain. In a significant portion of its range, C. serotina larvae feed on eriophyid mite-formed galls on the upper surface of Black Cherry Prunus serotina Ehrh. leaves. The larvae also utilize floral primordia of several additional hosts from different plant families. Adults are morphologically distinct from the earlier flying Spring Azures, C. ladon and C. lucia Auctorum, but have a clear resemblance to C. neglectamajor, C. idella, and C. neglecta. Separation from these lighter phenotype species is by size, distribution, habitat, and flight period. We propose the new species may be one of several late spring allochronic species forming a chain or great arc from the southern Appalachians to northeastern United States & eastern Canada, possibly across the northern Great Lakes region to southern Manitoba & Saskatchewan, to the Foothills of the Rocky Mountains in Colorado.

Full text

Volume 6 1 December 2005 Number 6
The Taxonomic Report
OF THE INTERNATIONAL LEPIDOPTERA SURVEY
CELASTRINA SEROTINA (LYCAENIDAE: POLYOMMATINAE):
A NEW BUTTERFLY SPECIES FROM
THE NORTHEASTERN UNITED STATES AND EASTERN CANADA
HARRY PAVULAAN1
P.O. Box 1124, Herndon, VA 20172
AND
DAVID M. WRIGHT
100 Medical Campus Drive, Lansdale, PA 19446
ABSTRACT. A new light-venter Azure species, Celastrina serotina, is described from the northeastern United States
and eastern Canada. The flight period of this univoltine spring species is interpolated (allochrony) between its sympatric
congeners, C. lucia Auctorum (not Kirby) and C. neglecta (Edwards). In this regard it is similar to C. neglectamajor Opler &
Krizek in the southern Appalachians and C. idella Wright & Pavulaan on the New Jersey coastal plain. In a significant portion
of its range, C. serotina larvae feed on eriophyid mite-formed galls on the upper surface of Black Cherry Prunus serotina
Ehrh. leaves. The larvae also utilize floral primordia of several additional hosts from different plant families. Adults are
morphologically distinct from the earlier flying Spring Azures, C. ladon and C. lucia Auctorum, but have a clear resemblance
to C. neglectamajor, C. idella, and C. neglecta. Separation from these lighter phenotype species is by size, distribution, habitat,
and flight period. We propose the new species may be one of several late spring allochronic species forming a chain or great
arc from the southern Appalachians to northeastern United States & eastern Canada, possibly across the northern Great Lakes
region to southern Manitoba & Saskatchewan, to the Foothills of the Rocky Mountains in Colorado.
DISCOVERY OF AN UNUSUAL BUTTERFLY
During Rhode Island field surveys in 1983-84, a conspicuous sequence of Celastrina emergences
was observed (Pavulaan, 1985). The earliest Azure to appear in early to mid-April (C. lucia Auctorum)
consisted of small individuals with dark dusky gray venters, commonly with black discal patches (f.
“lucia”) and/or broadened margins on the hind wings (f. “marginata Edwards”). C. lucia was found
widely distributed throughout the state, and was especially easy to observe in early spring along the sandy
woodland roads of the Great Swamp Management Area, near West Kingston, Washington County. The
Great Swamp population utilized Highbush Blueberry Vaccinium corymbosum L. as its host, and it was
subsequently learned that lucia utilized several species of Vaccinium in Rhode Island. Shrubby plants in
this genus are plentiful in the state, often forming dense thickets in old fields or a solid understory in
forested habitats. Because of its association with these habitats, C. lucia was encountered routinely in
virtually every region of the state in April. The flight reached peak numbers in late April. (See the
account of C. lucia Auctorum (“of authors”) in discussion of Celastrina lucia on p. 14.)
1 Staff Member, The International Lepidoptera Survey, Herndon, VA.

1 2 3
C. lucia Auct. C. serotina C. neglecta
Figs. 1-3. Sympatric Celastrina in northeastern United States (live specimens). ♂♂ Fig. 1. 3 May 1992, Caumsett S.P.,
Suffolk Co., NY. Fig. 2. 14 May 2000, Sayville, Suffolk Co., NY. Fig. 3. 15 July 2005, Jamaica Bay, Queens Co., NY.
Photos by Chris Adams (Fig. 1) and Steve Walters (Figs. 2 & 3).
By early May, lucia individuals showed signs of wear and their numbers began to fall. It was at
this time that a second irruption of Azures appeared. This emergence consisted primarily of very light-
ventered individuals, some appearing almost white on the first day of emergence and contrasting sharply
with the darker coarsely-patterned lucia phenotype. These light-ventered Azures were neither as common
nor as widespread as the earlier lucia flight, and appeared in only a few locations such as Great Swamp.
Individuals with darkened margins were entirely absent from this flight, however a few rare individuals
(1%) displayed a partly-developed ventral hind wing patch, reminiscent of the earlier “lucia” forms. The
initial thought was these were f. “neglectamajor Tutt”, although the biology of neglectamajor was poorly
understood at the time. Klots (1951) wrote, “In some regions occurs a partial second brood (neglecta-
major), large, very bright colored above, white beneath, with clear-cut markings. This is “spotty” in its
occurrence…” Swayed by Klots’ description, the Rhode Island light phenotype was misidentified as C.
neglectamajor by Pavulaan (1985). Reginald Webster (pers. com., 1984) suggested the unique Rhode Is.
butterfly may represent a new species, citing erroneous reports that neglectamajor utilized Viburnum
acerifolium L. (Shapiro, 1966; Opler & Krizek, 1984), a plant that was relatively scarce at Great Swamp.
It was reasonably concluded that V. acerifolium could not support the large population of what was
believed to be neglectamajor at this location. The first evidence of possible host association came when
several females were flushed from a young P. serotina tree in Warwick, R.I. on May 22, 1984, though it
was not immediately obvious that the species was utilizing leaf galls as the primary host.
In July, a third distant Azure flight (C. neglecta) was witnessed in Rhode Island. These butterflies
had pale white venters with faintly seen maculations. Their occurrence was very spotty and collectively
their flight was the least common of the three flights observed. Popularly thought to be the summer form
of a spring species, this notion seemed unfounded because C. neglecta individuals were seen in far fewer
numbers and in locations that often did not correspond with the earlier flights. A female was found
ovipositing on White Meadowsweet Spiraea alba var. latifolia (Ait.) along a utility powerline near
Cranston, Providence Co., on July 17, 1984. Because this host and cherry (Prunus sp.) were members of
the same plant family (Rosaceae), a link seemed possible, but oddly no Azures were found at this location
in May. At this point a strong suspicion emerged that the light-ventered May and July flights were
separated and represented different species. Shortly thereafter, rearing studies established that the light-
ventered May population was univoltine and reproductively separated from summer flights of C.
neglecta. Similar studies also established C. lucia was univoltine, although this was not entirely
unexpected since this Azure had a vast distribution in northern Canada and Alaska where only a single
flight was known. We began an effort to learn more about the biology and distribution of the light-
ventered May Azure.
2

C. lucia Auct. C. serotina C. neglecta
Figs. 4-6. Sympatric Celastrina in northeastern United States (spread specimens). ♂♂ Fig. 4. 16 April 2005, Connecticut
College Arboretum, New London, CT. Fig. 5. 21 May 2005, same site. Fig. 6. 24 July 2005, same site. (D/V). Photos by David
Wright.
6 5 4
HISTORICAL TREATMENT IN LITERATURE
The North American Celastrina are extremely diverse in their ecology and phenotypes. Whether
viewed as many closely-related species or as one polymorphous continent-wide species, the extraordinary
displays of regional variation provide opportunity for research and a deeper understanding of insect
evolution. This prospect must have allured those who first encountered the notable differences between
our native Azures. In 1841, Edward Doubleday commented, “I may just express my opinion that two
species are confounded under the name pseudargiolus ..”, an insightful view that foreshadowed a cascade
of papers describing new species from North America. In fact, while Doubleday traveled and collected in
the U.S., Kirby named one from Canada in 1837 (lucia), and William H. Edwards soon followed with
three more (neglecta, echo, violacea) from the United States in the 1860s. As new discoveries unfolded
during the last century, more species were continually added to our fauna. Remarkably, the light-ventered
spring Azure of northeastern U.S. and eastern Canada did not go unnoticed. A detailed account of its
appearance in the lepidopterological literature records no less than 50 entries. (See “Synonymy”, p.6.)
In the late 19th century, recognizable references to this unique Azure came mostly from authors
who resided in the species’ range. The great Azure debate of this period was the status of neglecta.
Writing from West Virginia, Edwards (1875) concluded that neglecta might prove to be the summer
brood of spring forms violacea [= ladon] and pseudargiolus (sensu Edwards) [= neglectamajor], and
inferred that in the north neglecta may be the summer brood of lucia inasmuch it was also a spring form.
Scudder (1876) in Massachusetts, Lintner (1875) in New York, & Saunders (1875) in Ontario challenged
Edwards’ concept and reported lucia and neglecta were not united. Scudder (1876) pointed out that
neglecta appeared twice a year in spring and summer, with the earlier neglecta appearance in May (in fact
serotina) being too close on the heals of the lucia to be derived from it. He subsequently concluded that
spring taxa, lucia in April and neglecta in May, were derived from diapausing chrysalids and their flights
were separated by successive eclosions (Scudder 1883, 1889).
The similarity of northern spring neglecta to spring form pseudargiolus [= neglectamajor] in West
Virginia was obvious to Scudder (1883) …“This blue corresponds in all respects excepting size with
Pseudargiolus proper of the south, and small southern individuals agree altogether with larger northern
examples.” Edwards (1883) agreed, “It is plain these Neglecta, flying with Lucia, must have come from
hybernating chrysalids, as truly did Lucia. The history of these Neglecta would appear to be identical with
the history of Pseudargiolus in W.Va. ...they are interpolated in the series just as Pseudargiolus is. In
fact, these early Neglecta would be very small Pseudargiolus.” Edwards (1884) believed that
pseudargiolus in West Virginia had a small second generation in late summer, and confessed, “If these
late butterflies were suppressed, Pseudargiolus would stand alone as a distinct species. So if anywhere to
the northward the winter form is suppressed, [spring] Neglecta would represent the species.” A century
later this proposition has been confirmed & a new Azure of the north has been gradually accepted
(Ferguson,1975; Pavulaan, 1985, 1989, 1993; Iftner et al., 1992; Wright, 1995; Layberry et al., 1996;
Thomas, 1996; Allen, 1997).
3

Celastrina serotina Pavulaan and Wright, new species
Description. Male (Figs. 2,5&7). Dorsal color uniform light blue; some individuals with purplish-blue tint. White insuffusion
between veins on DHW common. Androconia present. Wing fringes white; black checkering minimal to absent. Ventral color
uniform white to light gray-white. Black maculations reduced. Most individuals are lightly marked, similar to form “violacea”,
except in RI, MA, and PA where a few (1%, 3%, 9% respectively) have partially fused maculations on the VHW disc (form
“lucia”). FW length 10.2-15.0 mm (mean 13.5, n = 52). Rhode Island and southern New England males average slightly
larger than those from northern New England, Canada, and higher elevations in southern range (WV, MD, PA). Female (Fig.
8). Dorsal color lustrous metallic light blue; some individuals with purplish tint. Black on DFW costa and outer margin. DHW
with series of submarginal black dots. White insuffusion on DHW common; occasionally also on DFW. Wing fringes white;
black checkering minimal to absent. Ventral color and pattern as in male. Forewing length 10.5-15.0 mm (mean 13.4, n=28).
Rhode Island and southern New England females average slightly larger than those throughout rest of range. Note: The male
genitalia of eastern North American Celastrina species differ from Eurasian C. argiolus L., but they cannot be reliably
distinguished from one another. The sclerotized apical process of the valve bears small teeth, which tend to be largest in
neglectamajor and smallest in neglecta. All remaining taxa, including serotina, are intermediate in this character. No
distinguishing features of female genitalia of eastern North America are known.
Types. Holotype ♂ (Fig. 7): Great Swamp Management Area, near West Kingston, South Kingstown Township, Washington
County, Rhode Island, U.S.A., May 14, 1990, along main access roadway on east side of Great Neck, collected by H.
Pavulaan. Allotype ♀ (Fig. 8): Great Swamp Management Area, near West Kingston, South Kingstown Township, Washington
County, Rhode Island, U.S.A., May 21, 1984, along main access roadway on east side of Great Neck, collected by H.
Pavulaan. Holotype and allotype deposited in The Academy of Natural Sciences of Philadelphia (ANSP), Philadelphia, PA.
Paratypes: n=212. Paratypes deposited in ANSP, Carnegie Museum (Pittsburgh, PA), National Museum of Natural History
(Washington, D.C.), American Museum of Natural History (New York, NY), and collections of authors. Paratypes include: 53
♂♂, 15 ♀♀, Kent & Washington Cos., RI, May 2-May 23, 1983-1990; 76 ♂♂, 14 ♀♀, Suffolk Co., NY, April 18-June 6,
1981-1992; 35 ♂♂, 19 ♀♀, Monroe Co., PA, May 19-June 14, 1992-2005.
Differentiation from similar Celastrina.(Figs. 9-16) Phenotypically, C. serotina is most similar to three white-ventered
congeneric species in eastern North America (neglectamajor, idella, neglecta). Two of these species (neglectamajor, idella)
also fly in interpolated flights between those of their respective congeners. C. neglectamajor and C. serotina in many respects
have nearly identical color and maculation patterns. Neglectamajor (Figs. 9-10) is distinguished from serotina by size, range,
habitat, and larval host. The forewing length of neglectamajor averages 1.5-3.0 mm larger than the forewing of serotina.
Neglectamajor is found mainly in rich deciduous Transition Zone forests in the central & southern Appalachian Mts. and
Ozark Mts. It flies during the month of May following the flight of C. ladon and before the flight of C. neglecta. Its sole host
(Cimicifuga racemosa) is restricted to the same habitats, which prescribes the range of this unique sibling (Pavulaan & Wright,
2000). Serotina and neglectamajor are largely parapatric in distribution, but their ranges overlap in the central Appalachians
in WV, MD, PA, northern NJ, southeastern NY, & southwestern CT. Neither species utilizes larval hosts of the other nor have
adults been seen together in the same localities. C. idella (Fig. 11-12) is likewise distinguished from serotina by size, range,
habitat, and hosts. The forewing length of idella averages 0.5-1.5 mm smaller than the forewing of serotina. Idella
individuals from the type locality in the pine barrens of southern New Jersey show the greatest size discrepancy when
compared to serotina. The primary color difference in this pair exists in the females. Idella females tend to be slightly more
purplish; the discal streak on the hindwing of serotina is surrounded by a faint elongated whitish halo. Idella resides in the
sandy oak-holly forest habitats of the outer Atlantic Coastal Plain from New Jersey to Georgia. At its type locality, idella flies
in May after the flight of C. lucia and before a meager flight of C. neglecta. In the southern portion of its range, the flight is
sandwiched between flights of C. ladon and C. neglecta, with a signigficant overlap with the ladon flight. Its primary host in
southern New Jersey is Ilex glabra, but I. opaca and other holly species are also used throughout its range (Wright &
Pavulaan, 1999). Serotina and idella are allopatric in distribution. Despite the presence of C. lucia and potential hosts (i.e.
Prunus species with galls) in southern New Jersey, serotina has never been taken in this region nor have ovipositions on
cherry galls been witnessed. Idella hosts (I. glabra and I. opaca) are locally common in the Pitch Pine/Scrub Oak barrens of
Long Island, where serotina flies but idella has not been found. Accidental ovipositions by serotina females on I. glabra have
been observed; however, this host ultimately proved to be toxic to neonate serotina larvae. C. neglecta (Figs. 13-16) is fully
sympatric with C. serotina and all other eastern North American species. It ranges extensively over the eastern portion of the
continent from central peninsular Florida to approximately 480 N latitude. It is absent from Newfoundland. It is most easily
distinguished from serotina by voltinism, flight period, and physical appearance. Neglecta is multibrooded, except in the north
where only one brood flies. Summer individuals (Figs. 13-14) have less intense blue on the dorsum of both wings, often with
considerable white insuffusion between veins of the DHW. The hind wing of the female is occasionally nearly white. The
ventral surface is immaculate white with weak maculation pattern. A spring phenotype (Figs. 15-16) also occurs in most of the
central and southern states. (Neglecta tends to occur only in summer in northern states and southern Canada.) The spring flight
begins early, coinciding with the flight of C. ladon, well before the flights of neglectamajor and serotina. Spring neglecta
individuals have a solid blue dorsum without white insuffusion. The venter is light grayish-white, not as gray as ladon. The
dot pattern is prominent.
4

C. serotina
7 8
C. neglectamajor
9 10
C. idella
11 12
C. neglecta
summer phenotype
13 14
C. neglecta
spring phenotype
15 16
Figs. 7-16. Light-ventered Celastrina in eastern United States. ♂♂, ♀♀ (D/V). Fig. 7. Holotype, 14 May 1990, Great
Swamp, Washington Co., RI. Fig. 8. Allotype, 21 May 1984, same as 7. Fig. 9 & 10. 10 May 1999, Berrys, Clarke Co., VA.
Fig. 11. Holotype, 11 May 1987, Chatsworth, Burlington Co., NJ. Fig. 12. Allotype, 19 May 1990, same as 11. Fig. 13. 23
July 1988, Red Lion, Burlington Co., NJ. Fig. 14. 20 June 1992, same as 11 & 12. Fig. 15 & 16. 5 April 1988, SGL 157,
Bucks Co. PA. Photos by David Wright.
5

Etymology. The new species Latin name serotina is the feminine form of the adjective serotinus [L., from sero (adv.) = late]
meaning “coming late”. In a biological sense, the name implies delayed or late development with an appearance later in the
season. For the common name, although Cherry Gall Azure has appeared in print, we suggest the name Late Spring Azure is
more appropriate, reflecting its distinction from Spring Azure and Summer Azure.
6
Synonymy. Earlier workers most frequently referred to the northeastern May flight as neglecta or violacea. Names used in
literature in bold; not italicized. Polyommatus pseudargiolus “northern species” in Doubleday (1841) [“I may just
express my opinion that two species are confounded under the name of pseudargiolus, one a northern species, with markings
of the under surface very distinct and coarse, the other a southern one, in which all the markings below are beautifully
delicate..”] [EASTERN NORTH AMERICA] [Note: From Doubleday’s description above, one might suppose that his
“northern species” may be referable to lucia (Kirby, 1837). However, see next.]. Polyommatus pseudargiolus “northern
species” in Scudder (1869) [(Letter, Doubleday to Harris, April 30, 1842). “I am now quite convinced that pseudargiolus of
the North is not pseudargiolus of the South. There are two species. To which does the name belong? Boisduval’s plate will
not distinguish them.”] [EASTERN NORTH AMERICA] [Note: The “northern species” to which Doubleday refers
certainly could not have been lucia, since Boisduval & Le Conte’s [1833] plate would have easily distinguished
pseudargiolus from lucia. In a return letter, Harris to Doubleday (Nov. 17, 1842), Harris states, “the name Polyommatus
pseudargiolus must be applied exclusively to the Southern species, if the Northern blue species is distinct from it.
Deutargiolus would be a good name for the Northern species.” Neither Doubleday nor Harris ever published this name and
we consider it to be a nomen nudum. In a footnote to Harris’ last letter, Scudder (1869) states, “It has since been named L.
neglecta by Mr. W.H. Edwards [1862].” Scudder clearly understood that Doubleday was not referring to lucia in the north.
Scudder’s concept of neglecta, which differed from Edwards’, included a spring appearance of neglecta [May], which may
have been what Doubleday reported. Later, Harris (1862) concluded that “northern” pseudargiolus and lucia co-occurred in
Massachusetts.]. Lycaena neglecta in Packard (1869) [“Lycaena Neglecta Edwards is very common about Kalmia and
Rhodora in May, and a new brood appears in June and July.”] [MASSACHUSETTS] Cyaniris sp. ? in Parker (1874) [“like
neglecta, but ... dusky gray beneath”; Mt. Holyoke, June 25th; “violacea”, Amherst, June 1] [MASSACHUSETTS].
Lycaena neglecta in Edwards (1875) [“Mr. Scudder wrote me that in Mass. ‘neglecta, lucia and violacea all appear in
May’, the inference being that one could not be the parent of another.”] [MASSACHUSETTS]. Lycaena neglecta in
Saunders (1875) [“Neglecta is taken every season here [London, Ontario], the first brood usually during latter part of May
and early days of June, and the second brood in July and later.” “In 1862, two specimens on the 14th and one on the 15th of
May.” [ONTARIO]. Lycaena neglecta in Lintner (1875) [“I can not believe that neglecta and lucia will ever be united as
seasonal varieties of the same species.”; “Our observations do not agree with those of Mr. Edwards, giving June as the
earliest appearance of neglecta.”; “In [May] ... neglecta occurred in great abundance.”] [NEW YORK]. Cyaniris violacea
(in part) in Scudder (1876) [“..specimens of Cyaniris in any one locality seem to become larger, more lightly marked
beneath ... as the season advances.”; “[violacea] makes its advent during the first week of May ... and it still remains upon
the wing throughout June.”] [NEW ENGLAND]. Cyaniris neglecta (in part) in Scudder (1876) [“Lucia appear[s] in New
England but once, neglecta twice a year.”; “.. this insect [neglecta] is double-brooded ... earliest males appear at or shortly
before middle of May, but do not become abundant before the last week of the month; the first females appear about ten
days later than the males, but are still rare at the beginning of June ... earliest butterflies of the second brood appear about
the first of July.”; “.. it is extremely uncommon for two such closely allied species as neglecta and Lucia to differ ... in the
number of their broods.”] [NEW ENGLAND]. Cyaniris pseudargiolus f. violacea (in part) in Scudder (1881) [“.. in the
vicinity of Boston .. first appears Lucia .. the next blue butterfly to appear is ... violacea. It makes its advent during the first
week of May and remains on the wing until late June. There is no alternative but to suppose .. this blue butterfly is
dimorphic in the northern part of its range, appearing as an earlier and later form, Lucia and violacea.”] [NEW ENGLAND].
Cyaniris pseudargiolus f. pseudargiolus in Scudder (1881) [“Shortly after the appearance of violacea , or about the middle
of May, we encounter another blue butterfly, too early to have been produced from violacea, just as violacea appeared too
soon to have been produced from Lucia. This blue corresponds in all respects excepting size with Pseudargiolus proper [=
neglectamajor] of the south, and small southern individuals agree altogether with large northern examples.”] [NEW
ENGLAND]. Lycaena pseudargiolus f. neglecta in Edwards (1883) [“It is plain that these [northern spring] Neglecta,
flying with Lucia, must have come from hybernating chrysalids, as truly did the Lucia. The history of these Neglecta would
appear to be identical with the history of Pseudargiolus [= neglectamajor] in W.Va., … they are interpolated in the series
just as Pseudargiolus. In fact, these early Neglecta would be very small Pseudargiolus, though perhaps indistinguishable
from the examples of Neglecta which come ... later.”] [ONTARIO, NEW YORK]. Lycaena pseudargiolus f. neglecta in
Edwards (1884) [“Pseudargiolus [= neglectamajor] is an interpolated spring generation. Its second generation comprises a
part of the few butterflies which fly [later in summer]. If these late butterflies were suppressed, Pseudargiolus would stand
as a distinct species. So, if anywhere to the northward the winter form is suppressed, Neglecta would represent the species.
Prof. Lintner describes Neglecta as appearing in swarms at Centre, N.Y. ... and also in the vicinity of Albany, from middle
of May to middle of June. These myriads of course are from hibernating chrysalids. Neglecta is the winter form at Albany.”
[NEW YORK]. Cyaniris pseudargiolus f. neglecta, “the earlier neglecta” in Scudder (1889) [“… how does neglecta
appear so early?” “The only alternative .. is to regard .. the earlier neglecta as part of the first brood which then becomes
trimorphic.” “The successive apparition of lucia, violacea, and neglecta before July in New England should be looked upon
as the successive eclosion from wintering chrysalids of first one, then a second and finally a third form.”] [NEW

ENGLAND]. Cyaniris pseudargiolus f. neglecta in Scudder (1893) [“..the earliest (form lucia) generally appear about the
middle of April, and in the first week of May the numbers are materially increased by the advent of the form violacea, ..
further accompanied, after the middle of May, by the third form, neglecta .. In June, lucia is rarely seen and the others
disappear one after the other; but in July the second brood proper appears, consisting wholly of neglecta, and continues to
emerge .. even into September.”] Celastrina ladon ladon “late spring form” in Comstock & Comstock (1904) [“The late
spring form … appears from over-wintering chrysalids, but considerably later than the forms described above, a fourth
spring form. This is the largest form of the species … Mr. Scudder does not regard neglecta as distinct from C. ladon ladon
… according to this view neglecta is one of the spring forms as well as the summer form. And we have had difficulty in
separating the two by characteristic of size ...”] [EASTERN U.S.]. Cyaniris ladon (in part) in Engel (1908) [“Var. violacea
.. [to] May 17. Common.”; “Var. neglecta ... April 3- June 9. Common.”] [PENNSYLVANIA]. Lycaena ladon (in part) in
Smith (1910) [“form violacea occurs in April and May”] [NEW JERSEY]. Lycaena ladon (in part) in Davis (1910) [“a
female violacea {May 14}”] [NEW YORK]. Lycaenopsis argiolus pseudargiolus f. vernalis neglecta-major (in part) in
Comstock (1940) [“This form follows the early spring forms and is intermediate in occurrence between them and the first
summer brood.”] [NEW JERSEY]. Lycaenopsis argiolus (in part) in Klots (1951) [“On the average the darker specimens
[lucia, marginata] ... represent earlier specimens, the lighter ones [violacea], later ones.”] [EASTERN U.S.]. Lycaenopsis
argiolus lucia in Ferguson (1954) [“few ...violacea emerging late...localized, occurring in certain areas only.”] [See
Ferguson (1975) below.] [NOVA SCOTIA]. Lycaenopsis pseudargiolus pseudargiolus f. neglecta-major in Clench
(1958) [“The brood sequence of this species in unusually complex. In very early spring, during the latter half of April, the
small dark spring form violacea flies, chiefly in the woods. In early May, this is suddenly replaced by the ..lighter, faster-
flying neglecta-major, which seems to show a preference for more open areas.”] [PENNSYLVANIA].1 Lycaenopsis
pseudargiolus “late spring brood” in Remington (1958) [“..somewhat less numerous late spring brood”; Fig. 2, 4th
column (right), top, plus probably all of 3rd column (photo).] [CONNECTICUT]. Plebejus argiolus pseudargiolus f.
violacea in Forbes (1960) [“Three broods in the middle region, the first successively lucia auct., marginata and violacea in
New England to New Jersey.”] [NEW ENGLAND, NEW JERSEY]. Lycaenopsis argiolus (in part) in Shapiro (1966) [“a
few are [spring] form violacea resembling the summer brood beneath”; “violacea increases in frequency as the spring
emergence progresses.”] [More applicable to C. idella Wright & Pavulaan, 1999] [PENNSYLVANIA, NEW JERSEY].
Lycaenopsis argiolus (in part) in Shapiro (1974) [“A very complex species, or perhaps group of sibling species.”; “spring
form violacea”] [NEW YORK]. Celastrina pseudargiolus in Ferguson (1975) [“adults ... larger, paler, and fly in June,
about a month later than the smaller form [lucia] but not quite late enough to be a second generation .... long suspected that
these are two species.”] [NOVA SCOTIA]. Celastrina lucia in Chew & Robbins (1984) [“occasionally oviposit on galls on
their normal foodplants.”] [NORTH AMERICA]. Celastrina ladon “late spring form” in Opler & Krizek (1984) [“The
Appalachian Blue [neglectamajor] is most similar to the late-spring form of the Spring Azure, but it is significantly larger.”]
[EASTERN U.S.]. Celastrina ladon in Leblanc (1985) [“du sud du Quebec ... 2 pics d’abondance…un en mai-debut juin,
l’autre en fin juin-juillet “] [QUEBEC]. Celastrina neglecta-major Tutt ? in Pavulaan (1985) [“evidence suggests .... a
sibling species”] [RHODE ISLAND]. Celastrina ladon form “violacea type-II" in Pavulaan (1989) [“…two spring
flights overlap their flight period during May, and there may be two different species …females have been observed to
oviposit on these galls [wild cherry, P.serotina] in other states, and the larvae readily feed upon them.”] [RHODE
ISLAND]. Celastrina sp. "spring phenotype" in Iftner et al. (1992) [“… research suggests that several sibling species are
included within the C. ladon complex”; Pl. 24, row 6 (photo)] [OHIO]. Celastrina argiolus in Opler & Malikul (1992)
[“...we know there are at least three [species], and very probably there are at least several other sibling host-plant
specialists.”; “There may be as many as three species covered by our current concepts [Spring Azure].”] [EASTERN
NORTH AMERICA]. Celastrina ladon in Glassberg (1993) [“A bewildering complex of species and forms. Although
recent authors have begun treating [neglectamajor and nigra (= ebenina)] as distinct species, there are almost certainly other
species—In our area there may be at least three species ...”; Plate 11, no. 4, Spring Azure, form violacea, 5/9/90, Blue Hills
Reservation (photo)] [MASSACHUSETTS, EASTERN U.S.]. Celastrina ladon "Pine Azure (in part)” in Walter (1993)
[NEW YORK]. Celastrina ladon form "violacea type-II" in Pavulaan (1993) [“second spring flight … Larvae feed on
leaf-surface galls of wild cherry.”] [RHODE ISLAND]. Celastrina ladon race “violacea” type II in Pratt et al. (1994)
[“violacea” .. type II feeds on leaf galls of Prunus serotina, induced by the eriophyid mite Phytoptus cerasicrumena Walsh
… Larvae of “violacea” II populations are also known to feed locally on Viburnum and Aralia flowers … host specialization
may have selected for [adult eclosion] asynchrony .. [a] mechanism inducing genetic isolation ..”] [EASTERN NORTH
AMERICA]. Celastrina ladon “Late Spring Azure” (in part) in Pavulaan & Wright (1994) [“The main host on Long
Island and in New England is black cherry .. the larvae feed on the red leaf galls which infest the cherries. Late Spring
Azures utilize several additional hosts in other areas ..”] [NEW YORK, NEW ENGLAND]. Celastrina sp. “Cherry Gall
Azure” (in part) in Wright (1995) [“… these azures fly between the flights of Spring Azures and Summer Azures, from
mid-May to mid-June … The main host from West Virginia through Pennsylvania to southern New England is Black Cherry
(Prunus serotina) leaf galls.”] [EASTERN NO. AMERICA]. Canadian [Celastrina] species yet undescribed in Layberry
(1996). [“It flies in between the flight seasons of Spring and Summer Azures .. the provisional common name is the Cherry
Gall Azure.” [NOVA SCOTIA, PRINCE EDWARD ISLAND, QUEBEC, ONTARIO]. Cherry Gall Azure (no scientific
name) in Thomas (1996) [“A third species that is as yet undescribed …occurs in New Brunswick. It is single-brooded and
7

flies in June. Any freshly-emerged azure butterfly flying in early-to-mid June is likely to be this species.”] [NEW
BRUNSWICK]. Celastrina ladon ladon race “violacea” Type II in Allen (1997) [“A separate sibling flight of blues
appears between the flights of .. lucia .. and neglecta at higher elevations .. flight parallels that of the Appalachian Blue
(Celastrina neglectamajor) .. univoltine in West Virginia.” (photo)] [WEST VIRGINIA]. Celastrina sp. “Cherry Gall
Azure” in Gochfeld & Burger (1997) [“This entity is slightly smaller than the very similar Appalachian Azures which
would be flying at the same time.”] [NOVA SCOTIA, QUEBEC, ONTARIO, MAINE, MASSACHUSETTS, NEW YORK,
PENNSYLVANIA, WEST VIRGINIA, NEW JERSEY, OHIO. S.E. MICHIGAN]. Celastrina n.sp. (unnamed new
species) CHERRY GALL AZURE in Wright (1998) [“Originally discovered on black cherry Prunus serotina leaf galls
created by leaf mites. Now known to be multiphagous utilizing other hosts in allochronic flight window (between ladon and
neglecta flights). Univoltine.”] [OHIO]. Celastrina ladon “violacea” type II in Scott & Wright (1998) [“This taxon [C.
humulus] is named as a species here because its closest relative in eastern U.S., namely ‘violacea’ II (of Pratt et al. 1994)
whose larvae eat Prunus serotina galls, is being named as a species.” Celastrina sp. “Cherry Gall Azure” in Layberry et
al. (1998) [“This species looks more like a Spring Azure than a Summer Azure but tends to be slightly paler, with more
white dusting over the blue than in the Spring Azure.” “Adults … in areas where cherry leaves are infested with mite galls
after the flight of the Spring Azure has waned.” (photo) ] [NOVA SCOTIA, PRINCE EDWARD ISLAND, QUEBEC,
ONTARIO]. Celastrina complex “Cherry-gall Azure” in Pelikan (1998) [“It was not until May 12, when azures [C. lucia]
had been flying for a full month, that I first encountered a ‘violacea’ … But on May 16 the relative abundance of the form
‘violacea’ underwent a meteoric rise. This second flight (apparently consisting exclusively of the form ‘violacea’) continued
widespread into June. Azures flying from mid-May into early June were probably Cherry-gall Azures, a newly-defined
species.…” (Martha’s Vineyard)] [MASSACHUSETTS]. Celastrina ladon unnamed ssp. “Cherry Gall” Spring Azure
in Glassberg (1999) [“Flies between broods of Spring and Summer azures”; Plate 23, no. 6, 6/14/93, Mt. Greylock, MA
(photo)] [EASTERN NORTH AMERICA]. Celastrina “ensemble innommé” (sp.) in Handfield (1999) [“Selon Wright
(1995) (ensemble nommé ‘Cherry Gall Azure’)”] [QUEBEC, ONTARIO]. Celastrina undescribed species in Wright &
Pavulaan (1999) [“The species associated with mite-induced leaf galls of cherry trees Prunus serotina Ehrh. and P.
virginiana L. occurs further north {than C. idella} (photo)] [NEW JERSEY, PENNSYLVANIA, CANADA].
[U]ndescribed cherry-gall feeding taxon in Pavulaan & Wright (2000) [“Subsequent research by the present authors
shows that this Rhode Island insect is not Appalachian neglectamajor, but a distinctly different undescribed species that
feeds on cherry galls (Pavulaan and Wright, in prep.)”] [RHODE ISLAND]. Cherry Gall Azure (species unnamed) in
Pavulaan (2002) [“… emerges in early May (fresh) and flies into early June (worn) … caterpillars feed on the red leaf galls
of Black Cherry trees in most of the northeast but some colonies have switched to other hosts in some areas.”]
[CONNECTICUT]. Spring Azure (C. ladon ladon) in Dirig (2002) [“.. a male Spring Azure offers rare view of its shining
blue topside.” (photo)] [NEW YORK]. Celastrina ladon “Spring Azure Cherry Gall population” in Allen et al. (2005)
[“The named populations of Spring Azure, adapted to particular hosts or groups of hosts, may eventually be considered to
be anything from full species to less than subspecies.” (larval photo)] [PENNSYLVANIA]. Celastrina ladon species
complex in Wagner (2005) [“The Spring Azure is a complex of several sibling species. The most remarkable entity in this
confusing array may be the undescribed Cherry Gall Azure, whose larvae consume the eriophyid mite nipple galls that occur
on cherry leaves.”] [EASTERN NORTH AMERICA]. Celastrina sp. Cherry Gall Azure in Cech & Tudor (2005) [“A
confusing segregate, not yet systematically described … Best noted for the fact that in many populations caterpillars feed on
leaf galls, especially those of cherries. Single-brooded, the Cherry Gall flies slightly later than Northern or Spring Azure at
any given location (overall mid-May to early June).” (photo)] [NEW ENGLAND, NEW YORK, PENNSYLVANIA].
1 In Carnegie Museum files, there is an unpublished manuscript by H. K. Clench (approx. 1974 - 78) titled, "Celastrina pseudargiolus form neglectamajor Tutt
and its types (Lycaenidae)." This manuscript has the following enlightened description: "An interesting and little known fact is that neglectamajor exists in two
forms. They have different facies, and different ranges, but preserve the same phenological attributes (appearing towards the end of the first brood of
pseudargiolus). Form (1) true neglectamajor. Large... Form (2) Small. un [underside] with markings much stronger, barely less intense than in pseudargiolus
form "violacea", but the ground usually quite white. This is the only form of neglectamajor that I have found at Powdermill Nature Reserve (Westmoreland
Co., Penna.). There it consistently appears near the end of the spring brood of pseudargiolus (from May into June). "
Flight period: One brood per year. Follows flight of C. lucia Auct.; precedes flight of C. neglecta. CANADA: Mid-May to
mid-June in eastern Ontario. June to early July in Ouebec and Maritime provinces. UNITED STATES: Late May through
June in northern New England and northern New York. Mid-May to mid-June in so. New England south to Virginia and West
Virginia. CONNECTICUT: May 2 - June 16 (May 23rd mean, n = 40). MASSACHUSETTS: May 3 - June 26 (May 27th mean,
n = 118). RHODE ISLAND: May 2 - June 22 (May 20th mean, n = 698). PENNSYLVANIA: May 16 - June 28 (May 29th
mean, n = 600). The annual flight period typically lasts 31/2 to 4 weeks.
Phenology: Flight phenogram (Fig. 17) of southern New England C. serotina population demonstrates the interpolated flight
of C. serotina relative to its sympatric congeners C. lucia Auct. and C. neglecta. Flights of the three species appear as
successive irruptions in the spring. In general, C. serotina emerges about three to four weeks later than C. lucia Auct., and
three and a half to four weeks before C. neglecta. The phenological pattern is analogous to C. neglectamajor in Appalachians,
which flies between flights of C. ladon and C. neglecta.
8

0
50
100
150
200
250
300
350
neglecta
serotina
lucia
Fig. 17
April May June July
Fig. 17. Flight phenogram of sympatric Celastrina species in southern New England. Adults in six-day intervals.
Data from Massachusetts Butterfly Atlas Project (1986-1990), Connecticut Butterfly Atlas Project (1995-1999), Rhode
Island surveys (1984-2004), museum & private collections (n = 2417).
Adult Nectar Sources: C. serotina males are known to frequent damp earth along dirt roadways, especially early in their
emergence period. Females are not known to frequent damp earth. Adults have been observed on blossoms or flowers of
Black Cherry P. serotina, Pussy Willow Salix discolor Muhl., Chokeberry (Aronia sp.), Daisy Fleabane Erigeron annuus (L.)
Pers., Red Clover Trifolium pratense L., and Yellow Mustard Barbarea vulgaris R. Br.
Larval Hosts: ROSACEAE: In a significant portion of its range, from southern New England to southwestern Ontario &
south to Appalachian plateaus of West Virginia-Maryland-Pennsylvania, C. serotina larvae eat elongate red galls formed on
the upper surface of Prunus serotina leaves (Fig. 19). The galls are induced by the wormlike eriophyid mite Phytoptus
cerasicrumena Walsh (Eriophyidae). Individual galls often start as small greenish bumps on the surface of young leaves, then
acquire red pigment (anthocyanin) as they mature. They may extend up to about 8 mm in length when fully formed, attached to
the leaf by a slender stalk. The galls tend to crowd the leaf surface and may often cover entire leaves, generally causing them
to curl into distorted shapes. The interior of the galls is hollow, sheltering the mites. Young larvae feed by boring a hole in the
side and extending their heads into the gall. Older larvae will devour the entire gall. Larvae inadvertently consume mites
residing within the galls. Although not primary carnivory, this is the second documented carnivorous butterfly caterpillar in
North America after Feniseca tarquinius (Fab.). The larvae are also occasionally found on leaf galls of Chokecherry P.
virginiana L., formed by the same mite. Very small galls are present on leaves of Pin Cherry P. pensylvanica L., but no larvae
have been discovered on this host to date. Rarely, C. serotina females have been observed ovipositing on the floral buds of P.
serotina in mid-May (Middlesex Co., MA, fide Schweitzer). A single mature 4th instar larva was found on floral buds of
Spiraea alba var. latifolia. [Note: Other host families may serve as secondary hosts in habitats with high gall density, or as
primary hosts in habitats with low or absent gall numbers. Larvae and eggs have been collected on the following families.]
CAPRIFOLIACEAE: Nannyberry Viburnum lentago L., Possumhaw V. nudum L., Northern Bush Honeysuckle Diervilla
lonicera Mill. ARALIACEAE: Bristly Sarsaparilla Aralia hispida Vent. (Ferguson, 1975; Layberry et al. 1998).
RHAMNACEAE: NJ Tea Ceanothus americanus L. CORNACEAE. Alternate-leaved Dogwood Cornus alternifolia L., Red
Osier Cornus stolonifera Michx. [Note: Some hosts may also serve as host for C. neglecta, following the flight of C. serotina.]
Several C. serotina females were reported in close association with Wild Sarsaparilla Aralia nudicaulis L. in May, but no
ovipositions were observed (Worcester Co., MA, fide Ziegler). Wagner & Mellichamp (1978) collected Celastrina eggs and
first instar larvae in Washtenaw Co., MI, on two of the aforementioned hosts (V. lentago, C. stolonifera) in May. They reared
these to maturity, but did not state if pupae diapaused. Accidental Hosts: Rare ovipositions were witnessed in May on Crown
Vetch Coronilla varia L (Monroe Co. PA) and Inkberry Ilex glabra L. (Suffolk Co., NY). In both instances, the host proved to
be toxic and 1st instars did not survive. Other Celastrina larvae on galls: Cherry gall feeding is not exclusive to C. serotina.
Larvae of several other Celastrina species have occasionally been discovered in nature consuming galls, consistent with their
polyphagous habits. Documented cases are spotty, but may be more widespread wherever leaf galls occur. Records: C. lucia.
PA: Franklin & Monroe Cos.; eastern MA (R. Robbins, pers. com.). C. ladon. PA: Bucks Co. C. neglecta. PA: Monroe,
Montgomery & Schuylkill Cos.; VA: Madison Co.; WV: Marshall
9

& Ohio Cos.; and IN: Howard & Monroe Cos. Rearing: Of 125 wild larvae collected on cherry galls in nature (MD, NY, PA,
RI), 117 developed successfully and pupated in the lab. C. neglectamajor will not oviposit on leaf galls in nature. When young
neglectamajor larvae are switched from their natural host Black Cohosh Cimicifuga racemosa (L.) to cherry galls in captivity,
they will readily accept them but produce dwarfed adults (Pavulaan & Wright, 2002).
Immature Stages. Egg (Fig. 20) white, similar to other Celastrina; placed on cherry galls or floral buds of hosts. Four larval
instars. First instar light green; bores into floral buds. Mature larva (4th instar) variable, usually light green to creamy white,
lightly contrasted with rosy mid-dorsal stripe, dorsal white chevrons, dark green & brown subdorsal blotches, maroon
prothoracic shield, and thin lateral white line. Generally lighter and less boldly marked than larvae of C. lucia Auct. Some
larvae dark red or brown with white chevrons, or occasionally wine red (Fig. 21), matching color of galls. Myrmecophilous
mature larvae with dorsal nectary organ on abdominal segment A7 and eversible tubercles on A8. Larvae attended by worker
ants of Formica subsericea Say, Lasius alienus (Foerster), Camponotus noveboracensis (Fitch), and C. pennsylvanicus
(DeGeer). Hymenopterous parasitoids recovered during rearing include a red-eyed and a blacked-eyed trichogrammatid
species (Trichogramma minitum & T. parkeri) from eggs, a small wasp Cotesia cyaniridis (Riley) from mature larvae, and a
larger wasp Anisobas luzernensis (Bradley) from pupae in the spring following overwintering diapause. The tachinid fly
Aplomyia theclarum (Scudder) was also recovered from mature larvae. Pupa light brown (7.7-9.0 mm, n=17), obtect. Pupation
dates, June 20-July 17 in Monroe Co., PA. Pupae diapause. Of 117 pupae formed ex larvae found on cherry galls, 116
diapaused. The single non-diapausing pupa in retrospect may have been C. neglecta.
Habitat. At Great Swamp, Washington Co., Rhode Island (type locality), 90-150 ft. elevation, C. serotina is most commonly
found along woodland roads of a “neck” of upland mixed deciduous hardwood forest surrounded by wetlands. The forest on
Great Neck consists primarily of Red Maple Acer rubrum L., several species of oaks (Quercus sp.), a mid-level understory of
American Holly Ilex opaca Ait., and a lower understory of blueberries and huckleberries (Vaccinium sp.). Black Cherry
Prunus serotina trees, uniformly and heavily-infested with eriophyid leaf galls, are frequent along the woodland roads, forest
edges, and sandy open, scrubby areas. The butterflies are seldom found within the wetlands except as transitory strays.
Throughout eastern Long Island (Suffolk Co.), in southeastern New York, C. serotina is common and widespread in a variety
of habitats. The host Prunus serotina is widespread in natural habitats on Long Island's acid sandy soil (deciduous forests,
Pitch Pine/Scrub Oak barrens), as well as a rather profuse pioneering species (“weed tree”) in old-field habitats and in virtually
any area affected by human activity including heavily urbanized and industrialized settings. The majority of these Long Island
trees are heavily infested with Phytoptus mites. During the first week of the C. serotina emergence, adults are primarily
restricted to forest-associated habitats and always near the host trees. As the flight season progresses, adults (mainly females)
stray out into the surrounding region, and can be found in virtually any habitat. The adults have been seen as strays in
suburban yards, neighborhood parks, schoolyards, commercial districts and industrial parks. It is not surprising to encounter
females in any wild or neglected open space. Some host trees in old-field habitats eventually succumb to an overgrowth of
invasive Common Greenbriar Smilax rotundifolia L. C. serotina occurs throughout the Pocono Mountains of northeastern
Pennsylvania. This area is a glaciated rugged highland plateau, 1800-2100 ft. elevation, with rocky soil, nutrient-poor bogs,
and an extensive forest cover dominated by a Beech-Maple forest. Black Birch Betula lenta L., Yellow Birch B.
alleghaniensis Britton, Quaking Aspen Populus tremuloides Michx., Bigtooth Aspen P. grandidentata Michx., Chokecherry
Prunus virginiana, and Black Cherry P. serotina are frequent associates of the Beech-Maple forest. Eastern Hemlocks Tsuga
canadensis (L.) are scattered throughout forest, especially in shaded river ravines. Serviceberries Amelanchier sp. and
Mountain Laurel Kalmia latifolia L. provide the understory. The wetlands are dominated by Highbush Blueberry Vaccinium
coyrmbosum and Great Laurel Rhododendron maximum L. The Pocono region has a large population of C. lucia Auct., whose
emergence begins initially in the wetlands in mid-late April and then “explodes” in the surrounding forest in early May. The
earlier lucia females oviposit on blueberry floral buds, while the later woodland females lay eggs on cherry flower buds.
(There is speculation that more than one species under the name lucia occurs in this region.) C. serotina does not begin to fly
until late May when the woodland lucia is markedly worn and its numbers have radically declined. Shortly after the flight
begins, large aggregations of fresh serotina males are seen at mud puddles in streamside trails and on gameland & state forest
roads. Females are observed exploring host trees. They selectively oviposit on leaf galls, even though a few cherry flowers
may still be unopened. They also utilize floral primordia of Nannyberry Viburnum lentago and shrubby dogwood species (C.
alternifolia, C. stolonifera). An interesting parallel has been observed between the spring Celastrina emergences of the region
and a sympatric pair of small day-flying moths. The delicate bluish spring geometrid Lomographa semiclarata (Wlk.) begins
flight in early-mid May. At the end of May and early June, as its numbers decline, a second whiter species L. vestaliata (Gn.)
begins to fly. Both Lomographa species utilize cherry and their staggered flights are reminiscent of the irruptive emergences
of different Celastrina species in the same woodland habitat. In higher elevations of northern Virginia & West Virginia, C.
serotina has been observed in relatively open mixed hardwood forest and associated open places where the understory host P.
serotina trees reach 25 ft. One studied location is the ridgetop forest to the south & east of Big Meadows Recreation Area, on
top of the Blue Ridge, 3200-3600 ft. elevation, in Shenandoah National Park (Madison and Page Cos., VA). C. serotina
emerges in mid-May about three weeks later than C. ladon. It flies with similar-looking, light-ventered siblings C.
neglectamajor and C. neglecta, which both begin to emerge in late-May, making accurate identification either by sight or
binoculars very difficult, if not impossible. To compound matters, where C. serotina larvae feed on gall-infested leaves, later-
emerging C. neglecta females will occasionally oviposit on the same leaf galls. However, in all instances, reared immatures of
gall-feeding C. neglecta do not diapause.
10

20 19
21 18
Figs. 18-21. Cherry galls and Celastrina serotina immatures. Fig. 18. Old growth field with Black Cherry (Prunus serotina)
saplings (arrow) (Dauphin Co., PA). Fig. 19. Eriophyid mite galls on upper surface of Black Cherry leaves (Monroe Co., PA).
Fig. 20. Egg of C. serotina on Black Cherry gall. (Monroe Co., PA) Fig. 21. Mature 4th instar larva of C. serotina consuming
Black Cherry leaf gall (Monroe Co., PA). Photos by David Wright.
C. serotina can stray widely and otherwise occur in a diverse array of habitats, attesting to it’s abundance in some regions. We
examined the voucher specimens and records of the Massachusetts Butterfly Atlas Project (1986-1990) and Connecticut
Butterfly Atlas Project (1995-1999) to index the habitat types from which this species has been recorded. They are as follows:
black spruce bog; heath lands; mature scrub habitats including thickets, brambles and scrub marsh; oak woods; pitch pine-
scrub oak barrens; a very broad range of deciduous & coniferous woods with species of oak, beech, birch, maple, hemlock,
pines, and black spruce intermixed to varying degrees; wood edges; power line utility right-of-ways; rocky hilltops or
mountain summits; various disturbed areas such as abandoned farms, fields, hillsides, gravel pits; a variety of open areas
ranging from open fields, wet & dry meadows, “old field” (second-growth) successional habitats of varying stages; suburban
lawn and garden areas; pasture. These descriptions detail a sizeable range of wooded and open habitat types, providing a sense
of the resourcefulness of this species.
Frequent Associates. C. serotina is occasionally found in woods, but more commonly along edges of woodlands and open
sunny second-growth habitats with Papilio glaucus L. (so. New England & Long Is., NY), P. canadensis Rothschild & Jordan
(no. New England, southeastern Canada), P. canadensis x glaucus (hybrid zone, no. PA.), P. appalachiensis Pavulaan &
Wright (so. PA, WV), P. rapae (L.), Colias philodice Godart, C. eurytheme Boisduval, Lycaena phlaeas (L.), Callophrys
(Mitoura) gryneus (Hübner), C. (Deciduphagus) augustinus (Westwood), C. (Deciduphagus) henrici (Grote & Robinson), C.
(Incisalia) niphon (Hübner), Chlosyne harrisii (Scudder), Phyciodes tharos (Drury), P. cocyta (Cramer), Nymphalis antiopa
(L.), L. arthemis arthemis (Drury), L. a. astyanax (Fab.), Megisto cymela (Cramer) (type 1), Coenonympha tullia (Müller),
Erynnis icelus (Scudder & Burgess), E. brizo (Boisduval & Le Conte), E. juvenalis (Fab.), E. horatius (Scudder & Burgess),
Poanes hobomok (Harris), and Amblyscirtes hegon (Scudder).
11

Range. The range of C. serotina (Fig. 22) coincides with the range of C. lucia Auct., which it follows in flight. Prime
densities occur in northeast Pennsylvania, New England, and eastern Canada. Should be looked for in Michigan and other
Great Lake states. More local records should turn up following careful search of proper habitat. Look for a new allochronic
flight after the flight of lucia. We have examined and verified specimens and reports from the following locations:
CANADA. NEW BRUNSWICK: CHARLOTTE CO.: “southern county”; GLOUCESTER CO.: Allardville, Daly Point,
Nigadoo, Petit-Rocher; KINGS CO.: Belleisle; MADAWASKA CO.: St. Jacques; NORTHUMBERLAND CO.: Holtville;
SAINT JOHN CO.: Saint Martins; WESTMORLAND CO.: Cap-Pelé, Moncton, Sackville, Shediack; YORK CO.: Charters
Settlement, Tower Lake, University of New Brunswick woodlot. NOVA SCOTIA: ANNAPOLIS CO.: Lequille; CAPE
BRETON CO.: Locality not given; COLCHESTER CO.: Truro; DIGBY CO.: Digby, Beamans Mountain; HALIFAX CO.:
Prospect Bay; HANTS CO.: Mount Uniacke; INVERNESS CO.: Cape Breton Highlands National Park; KINGS CO.: Black
Rock; LUNENBURG CO.: Petite Rivière; SHELBURNE CO.: Locality not given; VICTORIA CO.: Baddeck, Beinn Bhreagh,
Cape Breton Highlands National Park; YARMOUTH CO.: Argyle. ONTARIO: BRUCE CO.: MacGregor Point Provincial
Park, Port Elgin; DUFFERIN CO.: Luther Lake; DURHAM (Regional Municipality): Ajax; FRONTENAC CO.: Fermoy,
Thousand Islands; GREY CO.: Owen Sound; HALIBURTON CO.: Kinmount, Miner’s Bay, near Norland; HALTON CO.:
Campbellville; KAWARTHA LAKES (City, formerly Victoria Co.): Norland; LENNOX & ADDINGTON CO.: Cloyne;
MIDDLESEX CO.: London; MUSKOKA CO.: Bracebridge; NORTHUMBERLAND CO.: Northumberland Forest, Baltimore
OTTAWA (City, formerly Ottawa-Carleton Regional Municipality): Bells Corners, Constance Bay, Fitzroy Harbour, Kanata,
Ramsayville; PETERBOROUGH CO.: Warsaw Caves; SIMCOE CO.: Lefroy, Midhurst; TORONTO (City, formerly
Metropolitan Toronto): Saint Michael’s, Toronto; WELLINGTON CO.: Arkell, Guelph, Rockwood Conservation Area;
YORK (Regional Municipality): Brown Hill, Koffler Scientific Reserve at Joker’s Hill (U. of Toronto) west of Newmarket,
Vivian Forest. PRINCE EDWARD ISLAND: KINGS CO.: Pooles Corner; QUEENS CO.: Brackley Beach, Dalvay by the
Sea. QUEBEC: BELLECHASSE (Regional County Municipality): Saint Charles, Saint Raphael; LES COLLINES-DE-
L’OUTAOUAIS (Regional County Municipality): Chelsea; LES LAURENTIDES (Regional County Municpality): Parc du
Mont-Tremblant; LEVIS (City, separate municipality): Saint Jean Chrysostome, Saint Redempteur; LA JACQUES-CARTIER
(Regional County Municipality): Stoneham; LA CÔTE-DE-GASPÉ (Regional County Municipality, formerly Gaspé-Est Co.
in part): Forillon National Park; MONTMAGNY (Regional County Municipality): Montmagny; MONTREAL (City, separate
municipality): Montreal; QUEBEC (City, separate municipality): Quebec; RIMOUSKI-NEIGETTE (Regional County
Municipality): Macpes, Parc du Bic, Pointe-au-Pere, Saint Fabien.
UNITED STATES. CONNECTICUT: FAIRFIELD CO.: Sherman; HARTFORD CO.: Berlin, East Granby, Enfield, Rocky
Hill, Windsor; LITCHFIELD CO.: Canaan, Litchfield, Mohawk Mountain State Park, Nepaug, Norfolk, Salisbury;
MIDDLESEX CO.: East Haddam, Essex, Killingworth, Middletown, Middlesex, Old Saybrook; NEW HAVEN CO.:
Branford, 2 miles W of Clinton, East Haven, North Haven ; NEW LONDON CO.: Colchester, Connecticut College Arboretum
(New London), East Lyme, Franklin, Groton, Montville Center, New London, Old Lyme, Salem; TOLLAND CO.: Coventry,
Union; WINDHAM CO.: Plainfield, Scotland, Sterling, Thompson. MAINE: CUMBERLAND CO.: Brunswick; FRANKLIN
CO.: Locality not given; HANCOCK CO.: Bar Harbor, Southwest Harbor; KENNEBEC CO.: Augusta, Oakland; LINCOLN
CO.: Newagen, Boothbay; OXFORD CO.: Magalloway; PENOBSCOT CO.: Chester, Enfield, Lincoln, Lowell, Orono,
Passadumkeag; WALDO CO.: Isleboro; YORK CO.: Shapleigh. MARYLAND: GARRETT CO.: Swallow Falls State Park;
ALLEGANY CO.: near Frostburg. MASSACHUSETTS: BARNSTABLE CO.: Barnstable, Eastham, Harwich, Mashpee,
Truro, West Falmouth, Yarmouth; BERKSHIRE CO.: Ashley Falls, Becket, Egremont, Florida, Hinsdale, Lenox, Monterey,
Mount Greylock, New Ashfield, North Egremont, Sandisfield, Savoy, Sheffield, South Egremont, Washington, West
Stockbridge, Williamstown; BRISTOL CO.: Dartmouth, New Bedford, North Easton, Raynham, Westport; DUKES CO.:
Chilmark, Edgartown, Naushon Island, West Tisbury; ESSEX CO.: Ipswich, Manchester, Marblehead, Peabody, Rockport,
Salisbury; FRANKLIN CO.: Ashfield, Heath, Montague Barrens, Monroe, Northfield, Quabbin (Resevoir Gate #15), Rowe,
Shelburne; HAMPDEN CO.: Granville, Hampden, Holyoke, Wales; HAMPSHIRE CO.: Amherst, Belcherton, Mount
Holyoke, Northampton, Plainfield; MIDDLESEX CO.: Ashby, Billerica, Chelmsford, Concord, Dracut, Lexington, Medford,
Middlesex Fells Reservation, Stoneham, Waltham; NANTUCKET CO.: Siasconset; NORFOLK CO.: Blue Hills Reservation,
Medfield, Walpole, Wellesley, Westwood, Wollaston; PLYMOUTH CO.: Hingham, Lakeville, Marion, Marshfield,
Middleboro, Miles Standish State Forest, Plymouth, Scituate; SUFFOLK CO.: Boston, Hyde Park, Stony Brook Reservation;
WORCESTER CO.: Ashburnham, Douglas, Dudley, Gilbertville, North Brookfield, Petersham, Princeton, Sturbridge,
Uxbridge, West Brookfield, West Springfield, Westminster. NEW HAMPSHIRE: CARROLL CO.: Albany, Effingham;
CHESHIRE CO.: Marlow, Roxbury. COOS CO.: Crawfords Purchase, Pinkhams Grant, Second College Grant, Scott Bog,
Thompson & Meserves Purchase; GRAFTON CO.: Benton, Dorchester, Glencliff, Hanover, Lebanon; HILLSBOROUGH
CO.: Deering, Francestown, Goffstown, Greenville, Hancock, Mason, Pelham, Peterboro, Sharon, Windsor; MERRIMACK
CO.: Boscawen, Bow, Concord, Epsom, Hopkinton, Pembroke, Webster; ROCKINGHAM CO.: Newmarket, North Ampton,
Northwood, Portsmouth, Rye; STAFFORD CO.: Durham, Middleton, Rollinsford; SULLIVAN CO.: Croyden. NEW
JERSEY: SUSSEX CO.: Delaware Water Gap National Recreation Area, High Point State Park, Mashipacong Pond. NEW
YORK: ALBANY CO.: Albany, Colonie; Centre, Karner;CATTARAUGUS CO.: Allegany State Park; CHAUTAUQUA CO.:
Boutwell State Forest, Stockton State Forest; ESSEX CO.: Keene Valley; ALLEGANY CO.: near Wellsville; GREENE CO.:
Tannersville; MONROE CO.:
12

Fi
g
. 22
Rochester; NASSAU CO.: Bethpage State Park; ONEIDA CO.: Trenton Falls; QUEENS CO.: Corona; ST. LAWRENCE CO.:
Madrid, Morristown; SCHENECTADY CO.: Schenectady, Niskayuna; SCHUYLER CO.: Cayuta Lake; SUFFOLK CO.:
Babylon, Bay Shore, Brentwood, Brookhaven, Commack, Deer Park, East Islip (Heckscher State Park), East Moriches,
Eastport, Edgewood, Great River, Manorville, Middle Island, North Babylon (Belmont Lake State Park), Oakdale (Connetquot
State Park), Riverhead, Sarnoff Preserve, Sayville, Smithtown, Tiana (Munn’s Pond), West Islip, Yaphank, Wyandanch;
SULLIVAN CO.: Catskill Township (Catskill Mts.); TOMPKINS CO.: Ithaca, McLean Bogs Preserve, Monkey Run Road;
ULSTER CO.: New Paltz, Oliverea, Wittenberg Mountain. OHIO: COLUMBIANA CO.: Beaver Creek State Park.
PENNSYLVANIA: ALLEGHENY CO.: Pittsburgh; ARMSTRONG CO.: Elderton; BEAVER CO.: Locality not given;
BRADFORD CO.:Barclay; CARBON CO.: Split Rock; CENTRE CO.: State College; CLARION CO.: Clarion;
CLEARFIELD CO.: Shawville; COLUMBIA CO.: Bloomsburg; CRAWFORD CO.: Little Cooley; DAUPHIN CO.: Rt. 325
(3 miles SW of Orwin); ELK CO.: Allegheny National Forest (Buzzard Swamp); FOREST CO.: Allegheny National Forest
(Buzzard Swamp); INDIANA CO.: Clarksburg; LACKAWANNA CO.: Choke Creek (near Thornhurst); LUZERNE CO.:
Mountain Springs Road, Ricketts Glen State Park, Stoddartsville; MONROE CO.: Pocono Lake, Pocono Pines, State Game
Lands #127 ; MONTOUR CO.: Danville; PIKE CO.: Sugar Creek near Carlton Hill; SCHUYLKILL CO.: near Tamaqua (Owl
Creek); SOMERSET CO.: Seven Springs; SULLIVAN CO.: Lopez, near Ricketts Glen State Park (Rt. 487);
SUSQUEHANNA CO.: Montrose; WAYNE CO.: Gouldsboro; WESTMORELAND CO.: Jones Mills, Powdermill Nature
Reserve, Bushy Run Battlefield (5 mi. NE of Irwin), Greensburg; WYOMING CO.: Mountain Springs Road. RHODE
ISLAND: BRISTOL CO.: Barrington, Bristol, Warren ; KENT CO.: Coventry, Greene, Warwick, West Greenwich, West
Warwick; NEWPORT CO.: Jamestown, Little Compton, Middletown, Portsmouth; PROVIDENCE CO.: Burrillville,
Cumberland, East Providence, Foster, Johnston, North Smithfield, Scituate; WASHINGTON CO.: Charlestown, Exeter,
Hopkinton, Middlebridge, Narragansett, New Shoreham, Richmond, Shannock, South Kingstown, West Kingston (Great
Swamp Management Area), Westerly. VERMONT: BENNINGTON CO.: Beartown, Equinox Mountain, Manchester; ESSEX
CO.: Victory Township; GRAND ISLE CO.: Grand Isle (West Shore Road); WASHINGTON CO: Montpelier. VIRGINIA:
MADISON CO.: Big Meadows (Shenandoah National Park); PAGE CO.: Tanners Ridge (Shenandoah National Park);
RAPPAHANNOCK CO.: Sperryville (Shenandoah National Park). WEST VIRGNIA: HAMPSHIRE CO.: Ice Mountain;
HARDY CO.: River Road east of Moorefield; PENDLETON CO.: 1.5 mile NW of Upper Tract (Briggs Run Forest Road);
POCAHONTAS CO.: Cass; RANDOLPH CO.: Spruce Knob; TUCKER CO.: Blackwater Falls State Park.
13

A SHORT DISCUSSION OF CELASTRINA LUCIA
25 24 23
C. lucia (Kirby) C. lucia Auctorum C. serotina f. “lucia”
Figs. 23-25. Comparison of North American lucia’s. ♂♂ (D/V) Fig. 23. 1 June 1982, Whitehorse, Yukon. Fig. 24. 8 May
1993, State Game Lands #127, Monroe County, PA. Fig. 25. 29 May 1998, same site as Fig. 24. Photos by David Wright.
Presently, the name lucia comprises multiple meanings:
(1) William Kirby (1837) named Polyommatus lucia from material collected at Cumberland House,
Saskatchewan, 540 N latitude, during Franklin’s expedition to western Canada in 1825-27. This
population is widespread throughout the northern boreal forest where Labrador Tea Ledum
groenlandicum Oeder (Ericaceae) is a frequent larval host. Adults are small and very pale blue above
(Fig. 23), sometimes tending to greenish. The venter is dark gray with coarse brown markings. Following
its discovery, lucia Kirby had been treated for an extended period as a subarctic subspecies of a single
continent-wide species (ladon). Recent reappraisals by Wright (1998), Nielsen (1999), Ochlenschlager &
Huber (2002), and Opler & Warren (2002) have returned C. lucia (Kirby) to a full species. The precise
relationships between lucia populations at Cumberland House (Saskatchewan) and those in western North
America, Yukon-Alaska, and eastern North America have not been worked out.
(2) In eastern North America, lucia was loosely considered to be the northern population of the first
spring brood of the nominate eastern subspecies (ladon). It was pointed out by Klots (1952) and Forbes
(1960) this was not lucia Kirby, rather a highly polymorphic larger entity, solid blue above and quite
variable below (Fig 24). Pratt et al. (1994) designated it ssp. lucia Auctorum (“of authors”) and extended
its range into the central Appalachians. Lucia Auct. differs significantly from ladon in scale morphology
and ecology (Wright & Pavulaan, 1999). It is a separate species, but its relationship with lucia Kirby and
other lucia populations needs further study. The newly-described species (serotina) is closely linked with
eastern lucia Auctorum. We propose there may be a chain or great arc of allochronic late spring siblings
from southern Appalachians, north through northeastern U.S. & eastern Canada, west through the Great
Lakes to the Manitoba & Saskatchewan, and south to the Foot Hills of the Rockies in Colorado. In the
central continent, argentata Fletcher and humulus Scott & Wright should be examined in this light.
(3) Males from southern New England possess the unique scale of ladon males, but in all other respects
they are lucia Auct. This is believed to be an example of single character introgression during a warmer
postglacial period (Hypisthermal) when southern deciduous forests invaded southern New England
(Wright, 1971). Because of the introgressed trait, wings of males from this region (Fig. 4) contrast with
males elsewhere (Fig. 24). By good fortune, this circumstance allowed us to separate lucia Auct. and
serotina in southern New England not only by flight period but also by a precise morphological analysis.
We found no introgression of the unique scale trait into serotina, boosting our confidence in its status.
(4) The name lucia has also been adopted as a form name (f. “lucia”) that appears in many Celastrina
taxa. This form is characterized by coalesced macules in the middle of the VHW resulting in a black
discal patch. The form commonly occurs in early spring taxa like lucia and ladon, but also occurs in
light-ventered allochronic taxa like idella and serotina (Fig. 25). By itself, the presence of a dark VHW
patch is not a diagnostic trait of any specific taxon.
14

ACKNOWLEDGMENTS
We acknowledge our deep gratitude to John Calhoun, Doug Ferguson, David Iftner, Gordon Pratt, Ross Layberry,
Dale Schweitzer, and Reginald Webster who provided many stimulating discussions regarding this new taxon and Celastrina
biology.
We thank two reviewers for suggestions to improve the manuscript.
We sincerely thank the following individuals who provided us with specimens, photographs, notes, observations,
and/or reports: Chris Adams, Diane Adams, David Albaugh, Anurag Agrawal, Tom Allen, Jeffrey Belth, Richard Boscoe,
Walter Bosse, Richard Bray, Lenny Brown, John Calhoun, Brian Cassie, Rich Chyinski, Harry N. Darrow, Bob Dirig, Rick
Enser, Rachel Farrell, Jeff Fengler, Douglas C. Ferguson, Ed Force, John Friedel, Karl Gardner, Robert Gardner, Wm. David
Garrah, Jr., Ron Gatrelle, Hank Golet, Richard E. Gray, Paul Grey, Alex Grkovich, Helen Ghiradella, Louis Handfield, Alan
Hanks, Dave Hoag, Jack Holliday, David Iftner, Bernard Jackson, Marc Johnson, Phil Kean, Don Lafontaine, Ross Layberry,
Chris Leahy, Richard Lutman, Eugenia Marks, Kent McFarland, Andrew McGinnis, Todd McLeish, Mark Mello, Joanne
Michaud, Don Miller, Tony Moore, Mogens Nielsen, David Norris, Jane O’Donnell, David Parshall, John Peacock, Arthur
Plitt, Linda Eklof Read, Robert K. Robbins, Dale Schweitzer, James Scott, John Shuey, Jackie Sones, Daryll Speicher, Jim
Springer, Melanie Statts, Clare Stone, Clay Taylor, Anthony Thomas, David Wagner, Mark Walker, Steve Walter, Reed
Watkins, John Weber, Jr., Reginald Webster, Marty Wencek, Glenn Williams, William D. Winter, Jr., Ryan Woolwine, Bill
Yule, J. Benjamin Ziegler.
We extend our warmest thanks to the following individuals and their respective institutions for providing access to
institutional reference collections: The Academy of Natural Sciences of Philadelphia (Don Azuma, Jon Gelhaus, and Jason
Weintraub), Allyn Museum of Entomology (Jackie and Lee Miller), American Museum of Natural History (Fred Rindge and
Eric Quinter), Carnegie Museum (John Rawlins), Cleveland Museum of Natural History (Sonja Teraguchi), Dayton Museum
of Natural History (Reed Watkins), Delaware County Institute of Science (John Hallahan), Eastern College (David Unander),
Field Museum (Phil Parillo), Florida State Collection of Arthropods (John Heppner), Illinois Natural History Survey (Kathleen
Zieders), Los Angeles County Museum of Natural History (Julian Donahue), Michigan State University (Mo Nielsen),
Museum of Comparative Zoology (David Furth and Stephan Cover), National Museum of Natural History - Smithsonian
Institution (Robert Robbins), Natural History Museum London (Phil Ackery and Kim Goodger), New Jersey State Museum
(David Parris), Newark Museum (John Michalski), The Ohio State University Museum of Biological Diversity (Eric Metzler),
Pennsylvania Department of Agriculture (Karl Valley), Pennsylvania Department of Forestry (John Quimby), Pennsylvania
Natural Diversity Inventory (Barbara Barton), Purdue University (Arvin Provonsha), Reading Museum (Michael Feyers),
Royal Ontario Museum (Chris Darling), Rutgers University (Timothy Casey), Staten Island Institute of Arts & Sciences
(Edward Johnson), University of California Riverside (Greg Ballmer and Gordon Pratt), University of Delaware (Dale Bray
and Tom Wood), University of Guelph (Steve Marshall), University of Louisville (Charles V. Covell), University of Michigan
Museum of Zoology (Mark O’Brien), University of Pennsylvania (Andrew Binns), Wagner Institute (Eugene Bolt and Susan
Glassman), and Yale Peabody Museum (Charles Remington).
We extend our sincere appreciation to participants involved in two butterfly atlas mapping projects in New England
who kindly loaned their vouchers for examination. We thank the members of the Massachusetts Butterfly Atlas Project
(MBAP): Kathleen S. Anderson, Carroll M. Arbogast, James Baird, Jim Berry, William Bither, Jeff Boettner, Jason Brissette,
Tom Carpenter, Brian Cassie, Jonathan Center, J. Choimere, Richard A. Daub, Anne Deitrich, Tom Dodd, Edna L. Dunbar,
Ann S. Earley, Will Earley, E. Einhorn, Mark Fairbrother, Marilyn Flor, R. A. Foster, T. H. Fowler, T. W. French, David
Gregg, Stephan Goldstein, Richard A. Harlow, R. W. Hildreth, Karen Homes, A. P. Jacques, Mark J. Kasprzyk, Jane S.
Kessler, B. King, Ruth Kruger, Jim Lafley, Rene Laubach, Chris Leahy, Ernest A. Leblanc, Paul A. Levine, Lyn Lovell, David
Ludlow, William T. Maple, Mark Mello, Russell B. Miller, Fred R. Morrison, Donna Munafo, Ed Neumuth, Neil Osborne, R.
W. Peace, Wayne R. Peterson, Edward R. Pinkham, R. Prescott, T. Raymond, Don Reid, Steve Roble, Edward C. Rooks,
Shetterly & Hathaway, Nice Simpson, Sharon Stapleton, Tom Tysing, Richard K. Walton, Dennis R. Welin, Ron Wolanin, and
David Young; and the members of the Connecticut Butterfly Atlas Project (CBAP): A. Brand, Richard Chyinski, Victor
Demasi, J. Fengler, Ed Force, Lawrence Gall, B. Godwin, P. Godwin, Greg Hanisek, Ronald Harms, John Himmelman,
Richard Hyde, Chris Maier, Paxton Mallard, T. Marsh, Robbie Marshall, Bill Martha, Pat Martha, Leslie Mehrhoff, Bob
Muller, Jane O’Donnell, Linda Page, Annie Paradis, Noble Proctor, Marion Richardson, Ed Sluzenski, David Stevens, Clay
Taylor, Eric Thomas, Mike Thomas, C. Tondreau, Monty Volovski, and David Wagner. Special thanks are extended to Chris
Leahy, Massachusetts Audubon Society, Lincoln, MA, and Jane O’Donnell, University of Connecticut, Storrs, CT, for
arranging loans of specimens.
We thank Chris Adams and Steve Walter for making available their superb photos. A special thanks is extended to
Eileen C. Mathias (Librarian of the Ewell Sale Stewart Library of The Academy of Natural Sciences of Philadelphia) for
aiding in retrieval of important references. We are grateful for the expertise and efforts of D.L. Vincent, Paul Marsh and
Richard D. Smith, Research Entomologists (Systematic Entomology Laboratory, USDA), who identified the parasitoids and
ant attendants.
15

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_______________________________________
Date of publication: 1 December 2005
18

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