K The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY TK

DISCOVERY OF A BLACK FEMALE FORM OF

PTEROURUSAPPALACHIENSIS (PAPILIONIDAE: PAPILIONINAE) AND

ADDITIONAL OBSERVATIONS OF THE SPECIES IN WEST VIRGINIA

HARRY PAVULAAN ^

494 Fillmore Street, Herndon, VA 20170

DAVID M. WRIGHT

124 Heartwood Drive, Lansdale, PA. 19446

ABSTRACT: The univoltine Pterourus appalachiensis was described from the southern Appalachian Mountain region of the

United States as a sympatric sibling of P. glaucus (Pavulaan & Wright 2002). A black form was unrecognized at that time.

Subsequent sampling of populations at Spruce Knob, West Virginia, revealed a unique black female phenotype present among

typically yellow female appalachiensis. We believe that this phenotype represents a black female form of P. appalachiensis,

thus broadening our understanding of this unusual species and firmly establishing its distinction from P. canadensis.

Additional key words: mimicry, female-linked polymorphism, sympatry, introgressive-type hybridization

REVIEW OF THE BLACK FORM OF PTEROURUS GLAUCUS (LINNAEUS)

Papilio glaucus was described by Linnaeus (1758) from a black female, which he believed was a

distinct species unrelated to the large yellow swallowtails of eastern North America. As was customary in

18^*^ century natural history, no type specimen was designated and the name existed for over two centuries

without a name-bearing type. Honey & Scoble (2001) in their exhaustive survey of butterflies in the

Linnaean collections found no surviving P. glaucus specimens. In the absence of syntypes from which to

select a lectotype, we designated a typical black female of the summer brood (Fig. 6) as the neotype of

Papilio glaucus Linnaeus (see Pavulaan & Wright 2002). This specimen was collected on September 10,

2000, in the Sandbridge section of Virginia Beach, VA, a few hundred feet from the Atlantic Ocean.

Scott (1981) introduced the infrasub specific name nigra for the black female to distinguish it from the

“normally yellow” form. However, this name has no official taxonomic standing under the rules of the

ICZN (1999). The black female of P. glaucus is believed to be a mimic of the poisonous model Battus

philenor (Linnaeus), a dark-colored swallowtail that sequesters aristolochic acids from its larval hosts

(Sime et al. 2000). The mimetic black female of P. glaucus occurs in differing frequencies within the

species range, constituting a greater percentage of females in the southern portion and a lesser percentage

in the north. This distribution is believed to be in response to the frequency of B. philenor, which is more

common in southern United States and less common in the north.

DISCOVERY OF AN UNUSUAL BLACK PHENOTYPE OF PTEROURUS APPALACHIENSIS

During our original field study of Pterourus appalachiensis in years 1985-2001, no black females

corresponding to the size and behavior of typical yellow females were found in the mountains of North

Carolina, Virginia, Maryland, and southern Pennsylvania. The few black specimens retrieved for identi¬

fication proved to be rather typical black females of P. glaucus. We inferred that P. appalachiensis lacked

* Research Staff, The International Lepidoptera Survey, 126 Wells Road, Goose Creek, South Carolina 29445

a black female because of its general resemblance to P. canadensis (Rothschild & Jordan), a univoltine

species known to have no black female (Hagen et al. 1991). On June 10, 2001, two unusual black females

were captured among approximately 100 yellow P. appalachiensis females on the summit of Spruce

Knob (4861 ft.) in West Virginia. These black females (similar to Fig. 2) initially confounded us. Their

size matched the distinctive large size of P. appalachiensis (Fig. 5), and their overall appearance was

noticeably different than the local black female form of P. glaucus (Fig. 1). At that point, we were

undecided whether they were aberrations or an undescribed black female form of P. appalachiensis.

In the following years (2002-2004), several more black females of this phenotype were collected

on Spruce Knob, suggesting this form existed in a low but stable frequency in West Virginia. On June 5,

2002, four were collected among approximately 100 yellow P. appalachiensis females at Spruce Knob

and Spruce Knob Lake. On June 24, 2003, nine were collected or observed at these two locations, again

among approximately 100 yellow females. On May 29, 2004, the number increased significantly. Eleven

black females were collected or observed among approximately 50 females in two hours on Spruce Knob

summit. The reason for this sudden increase is unknown; it may reflect natural fluctuations in population

size, local weather conditions, and/or quality of nectar sources. (A sampling bias also cannot be ruled

A repeat search for this black female form in other mountainous portions of Virginia, Maryland,

and southern Pennsylvania in 2002-2004 was unproductive. Serendipitously, we received a report from

Richard Romeyn, who collected an unusual black female specimen on May 10, 2002, at Buck Creek in

western North Carolina (Clay Co.), the type locality of P. appalachiensis. Examination of the specimen

(Eig. 3) revealed a striking similarity between it and the black females from Spruce Knob. We concluded

that they were all black females of P. appalachiensis. The presence of the black form in the extreme

southern Appalachians and West Virginia implies it occurs throughout the species range. We attribute the

apparent scarcity of the black female to its low frequency and the general elusiveness of P. appalachiensis

females.

Spruce Knob is an excellent natural setting to observe large numbers of females. Why is this peak

so attractive to P. cppalachiensisl The answer most likely involves local topography and behavioral

tendencies of the butterfly. P. appalachiensis is primarily a forest canopy species and females are rarely

seen in the forest understory except when nectaring (Pavulaan & Wright, 2002). At 4861 ft. elevation.

Spruce Knob (Eig. 7, Top) is West Virginia’s highest peak and resides in the boulder-strewn Canadian

Zone. It has a relatively open canopy of Betula lenta (Black Birch), Nemopanthus mucronatus (Mountain

Holly), Picea rubens (Red Spruce), Prunus alleghaniensis (Allegheny Plum), Prunus pensylvanica (Pin

Cherry), Quercus rubra (Red Oak), Ribes rotundifolia (Smooth Gooseberry), Sambucus canadensis

(Black Elderberry) and Vaccinium sp. (Blueberries). Populus tremuloides (Quaking Aspen), a host of

Pterourus canadensis (Canadian Tiger Swallowtail) (Hagen et. al. 1991), also occurs in this area. The

western side of Spruce Knob slopes gradually away from the summit and is persistently windswept. The

landscape here has a distinctly subalpine character. P. appalachiensis has been observed flying up this

rocky slope toward the summit pushed on by strong westerly winds. On the summit are numerous

protected crannies characterized by montane heath vegetation (Eig. 7, Bottom), where adults may rest and

nectar. Rhododendron nudiflorum (Pinkster Flower or Pink Azalea) is locally abundant and blooms

precisely during the peak flight of P. appalachiensis (Fig. 8).

To the east of Spruce Knob, the terrain drops steeply from the summit. The forest on the eastern

slope is primarily Transition Zone. It consists of a vast variety of trees, including Acer pensylvanicum

(Striped Maple), Acer rubrum (Red Maple), Acer saccharum (Sugar Maple), Acer spicatum (Mountain

Maple), Betula alleghaniensis (Yellow Birch), Betula lenta (Sweet Birch), Fagus grandifolia (American

Beech), Fraxinus americana (White Ash), Liriodendron tulipifera (Tulip Poplar), Magnolia acuminata

(Cucumber Tree), Magnolia fraseri (Fraser Magnolia), Picea rubens (Red Spruce), Prunus serotina

(Black Cherry), Quercus prinus (Chestnut Oak), Quercus rubra (Red Oak), Sorbus americana (American

Mountain Ash), Tilia americana (Linden or American Basswood), and Ulmus americana (American

Fig. 1. Pterourus glaucus, black ? spring form, May 29, 2004, Spruce Knob, near Judy Gap, Pendleton Co., WV.

Fig. 2. Pterourus appalachiensis, black ? form, same data as Fig. 1. Fig. 3. P. appalachiensis, black ? form. May

11, 2002, Buck Creek, Clay Co., NC. (P. appalachiensis type locality). Fig. 4. P. appalachiensis, intermediate

“dusted” 9 form, same data as Fig. 1. Fig. 5. P. appalachiensis, yellow 9 form, June 10, 2000, Blue Mountain

Summit, near Linden, Warren Co., VA. Fig. 6. Neotype Papilio glaucus, black 9 summer form, September 10,

2000, Sandbridge, Virginia Beach, VA. All figures natural size.

Elm). The dominant trees are well protected from the prevalent winds of the western s]

nearly to the summit, forming a solid canopy. From vantage points on the summi t trail, one

the forest canopy to the east and observe P. appalachiensis flying above the treetops. Br

uphill over the forest canopy suddenly arrive on the summit among abundant bloomin

congregations of swallowtails. The strong westerly winds encountered on the immediate v

summit may have the effect of “herding” large swallowtails back to the summit area.

)s of the wing margin are more ai

le entire wing. In P. appalachiensis, these crescents a

le blue crescents in cells RS and Sc+Rl are often m

(50-65 mm) than P. glaucus females of the spring flight (34-53 mm) and slightly larger than the later summer P. glaucus

females (49-64 mm).

Venter: Typical striped pattern of 1

Hindwing ground color dark blackish brown with broad black margin and fringe edged with yellow. The classic tiger-

stripe pattern is evident. Black marginal area with row of six elongated submarginal lunules as in yellow appalachiensis

females. The first lunule in cell Sc+Rl rounded and deep orange. Lunules in cells RS, Ml and M2 whitish yellow but variably

tinged with orange. Lunule in cell M3 is a yellow curved crescent extending somewhat into the base of the tail at vein M3.

Lunule in cell Cul orange and often reduced to a narrow streak. Within the submarginal area, interior to the submarginal

lunules, is a terminal band of grayish-blue clouding, as in the yellow females, extending from cell Sc+Rl to cell Cul. Narrow

black median stripe crosses wing as in the yellow female. Dark discal veins form an arch. Veins M3 and Cul outlined with

additional black scaling enhancing thickness. Unlike the yellow females, the cells M3 and Cul between these veins do not

contain areas of orange coloration.

The venter of the female hindwing departs from the general appearance of the black form of P. glaucus primarily by

its more elongated submarginal lunules. Unlike yellow appalachiensis females, the lunules of known specimens tend not to be

rectangular, though they are not as arched as in P. glaucus. As in the yellow appalachiensis females, the inner edge contour of

the broad black marginal area generally forms a straight line in cells RS and Ml, angling outward in cell Sc+Rl and angling

inward in cell M2. In P. glaucus the inner edge of the margin is scalloped inwardly in each cell.

Specimen records. Since infrasubspecific form names have no official standing under the ICZN

rules, we refrain from naming this form and prefer to simply designate our specimens as 'Tterourus

appalachiensis black female form”. The specimen in Fig. 2 from Spruce Knob summit, 4861 ft. elev.,

Pendleton Co., WV, May 29, 2004, is retained in the collection of the senior author. It depicts the key

phenotypic features given in the description above. Additional specimens from the same location and

forming part of the descriptive study series are as follows: June 10, 2001 (n=2), June 5, 2002 (n=4), June

24, 2003 (n=9). May 29, 2004 (n=10). It is important to note that none were found at this location in mid

or late summer after the P. appalachiensis flight period. This supports it as a normal P. appalachiensis

form and not a form of P. glaucus or hybrid.

Additional Nectaring Observations in WV. On the summit of Spruce Knob, P. appalachiensis

adults nectar chiefly on Rhododendron nudiflorum (Pinkster Flower or Pink Azalea). This shrub thrives

in the rocky landscape (Fig. 7, Bottom), where stands of Red Spruce trees provide shelter from the wind.

Elaeagnus commutata (Autumn Olive) and Diervilla lonicera (Bush Honeysuckle) are used as nectar

sources just below the exposed summit area. Along the summit approach road in the Transition Zone

forest, adults use flowers of Crataegus crus-galli (Cockspur Thorn) at higher elevations and Erigeron

ramosus (Daisy Fleabane) at lower elevations. At Spruce Knob Lake, 3840 ft. elev., 3 mi. west in nearby

Randolph Co., D. lonicera is the primary nectar source. South of Spruce Knob, at Seneca State Forest in

Pocahontas Co. (Transition Zone), 2600-3600 ft. elev., P. appalachiensis adults including black females

have been observed and photo-documented on Kalmia latifolia (Mountain Laurel).

A REEVALUATION OF TIGER SWALLOWTAILS IN WEST VIRGINIA

In reexamining The Butterflies of West Virginia and Their Caterpillars (Allen 1997), an important

observation surfaced regarding the tiger swallowtails in Plate 3 (p. 253). Three specimens in this color

plate display key features described for Pterourus appalachiensis in Pavulaan & Wright (2002) and for

the black female form described above. In Row 1, the left specimen [Rt. 41, Prince, Fayette Co., WV,

May 22, 1987] fits the criteria for an appalachiensis male, while the right specimen is a rather typical

spring form glaucus male. In Row 2, the left specimen is a rather typical summer form glaucus female,

whereas the right specimen [East Fork of Glady Creek, Randolph Co., WV, June 18, 1984] fits the key

characters of the black female form of appalachiensis. In Row 3, the left specimen [East Eork of Glady

Creek, Randolph Co., WV, June 18, 1984] is a female specimen of intermediate “dusted” phenotype

fitting the general features of appalachiensis and is also similar to a “dusted” specimen we collected at

Spruce Knob (Eig. 4); the right specimen is a rather typical black summer form glaucus female. It is

interesting to note that all six specimens in this plate were photographed together in a single frame. From

this perspective, a general comparison of their relative sizes can be made. The appalachiensis specimens

are noticeably larger than the glaucus specimens. We also regard the male in Plate 20 (p. 286), Row 1,

right specimen [Lanesville, sic. Tucker Co., WV, June 18, 1973], as a typical P. appalachiensis male.

In Pavulaan & Wright (2002), we reported that many of the small individuals from the moun¬

tainous regions of West Virginia were phenotypically canadensis-like and we suspected that the small

phenotype might represent relictual P. canadensis. It appeared that P. canadensis and P. appalachiensis

overlapped in a small area of sympatry in the highlands of West Virginia, as evidenced by the two size

segregates that flew contemporaneously. However, we now believe that true P. canadensis is absent from

West Virginia. A preliminary electrophoresis study of the small canadensis-likt phenotypes from Spruce

Knob found only glaucus allozymes (Scriber & Ording, unpublished results). This new evidence

confirms the presence of glaucus at higher altitudes of the Appalachians, albeit as diminutive forms which

are far outnumbered by P. appalachiensis. These small variants of glaucus are more common at low and

mid elevations in early spring and can be easily confused with the northern species (Scriber 1990b).

To our knowledge only P. appalachiensis and P. glaucus occur in West Virginia. The southern

limit of true P. canadensis populations in eastern United States is poorly defined, but it is at least several

hundred miles north of West Virginia in northern New York (Adirondacks) and northern New England.

Immediately south, the two species P. canadensis and P. glaucus hybridize in a narrow zone across

central New York, northern Pennsylvania, and Massachusetts. Individuals from these hybrid populations

exhibit a composite of canadensis and glaucus traits (Scriber et al. 2003) and fly in polymodal fashion

(Hagen & Lederhouse 1984). It had been predicted that the southern Appalachians might provide a

corridor for the extension of the present hybrid zone or altitudinal refugia for canadensis types (Scriber

1996). Currently, P. appalachiensis satisfies neither hypothesis fully. Although Allen (1997) concluded

they were hybrids, appalachiensis individuals differ in many details from those in the present hybrid zone

(e.g. size, yellow female phenotype, black female, female behavior, neonate larva, flight mode, mtDNA).

Instead of viewing P. appalachiensis as a relictual ''canadensis type” in a southern Appalachian refuge, a

preferable treatment is to view this species as a relictual montane "glaucus type”.

SIGNIFICANCE OF BLACK FEMALE

AND THE TAXONOMIC STATUS OF P. APPALACHIENSIS

The discovery of a black female in P. appalachiensis is consistent with our previous conclusion

that this univoltine species evolved primarily from a glaucus genome. In many yellow Pterourus

swallowtails the black female is genetically suppressed (West & Clarke 1988). It is absent in the northern

species canadensis and the western species rutulus (Lucas), eurymedon (Lucas), and multicaudatus (W.F.

Kirby). Its presence in both P. appalachiensis and P. glaucus underscores the critical importance of

mimicry to achieve maximal fitness in habitats where a poisonous model exists. Battus philenor is very

common in the southern Appalachians. The abundance of Battus philenor in the mountains can be

attributed in part to the density of its local larval host,

Aristolochia macrophylla (Dutchman’s Pipe). A. macro-

phylla flourishes in the mountains of West Virginia

(Strausbaugh & Core 1978), where it is reported to be the

host for philenor (Allen 1997). (See Fig. 9. A.

macrophylla, dark gray. A. serpentaria, light gray. FNA

2000.) A single plant of this broad-leafed species can

support the growth of several larvae, whereas a single

plant of A. serpentaria (Virginia Snakeroot) is not

enough for one philenor larva to mature to pupation

(Rausher 1980). The role of ecological factors in natural

selection and in speciation cannot be overstated. The

absence of a black mimetic female may have selected

against P. canadensis and other hybrid "canadensis

types” in this region.

The distinctive large size and reduced blue scaling of the P. appalachiensis black female suggest a

possible aberration or developmental abnormality. We doubt this is the case, since this phenotype appears

regularly at Spruce Knob year after year and seems to be unchanged. An extensive search of the literature

relating to aberrations of tiger swallowtails found no natural examples matching the P. appalachiensis

black female (Clark 1932, 1936; Clark & Clark 1951; Clarke & Clarke 1983; Edwards 1884; Gunder

1927; Heithaus 2003; Howard 1899; Scriber 1990a; Scriber & Evans 1988; Scriber et al. 1987; Strecker,

1878; Tyler et al. 1994). To understand the development of the mimetic wing pattern we must turn to

glaucus where it has been studied extensively (Clarke & Sheppard 1959). The pattern is constructed from

several independently inherited characters that evolved in several steps. A single sex-linked gene controls

the black background. Any gene that produces a great deal of black will produce tolerable mimicry. This

female-limited gene appears to be an efficient switch because intermediates are rare. The other elements

in the mimetic pattern (blue scaling and red spots) are under polygenic control. The suite of genes

responsible for these elements are distributed on one or more autosomal chromosomes and inherited

independently of black. The regulation of autosomal blue scale genes is poorly understood. Interspecific

hybridization is a significant causal mechanism of unusual color morphs, and, occasionally, dark females

with reduced blue scaling result from hand-paired laboratory crosses (Scriber & Evans 1988; Scriber et al.

1987; Scriber et al. 1995). However, these hybrid individuals are predisposed to chromosomal alterations

that severely jeopardize their integrity and they (or their progeny) rarely survive. In contrast, the black

female phenotype in P. appalachiensis appears to be unique in nature and fixed as part of a stable female-

linked polymorphism.

We presently consider P. appalachiensis to be an established species and the dominant species at

higher elevations of the southern Appalachians. Its range is fully sympatric with P. glaucus and the two

species are easily distinguishable in the field. In our earlier paper (Pavulaan & Wright 2002) we

speculated that P. appalachiensis evolved initially as a montane race of P. glaucus, either independently

or through the introgression of unique genes. A recent electrophoresis study of specimens from Spruce

Knob discovered that appalachiensis individuals carry a mix of allozyme alleles {canadensis-iypt LDH,

glaucus-iypo^ PGD), suggesting introgressive-type hybridization and a relatively modern origin (Scriber &

Ording, unpublished results). The acquisition of favorable genes, such as an obligate diapause gene and

genes expanding tolerances of larvae and pupae, may have acclimated proto-appalachiensis to the

southern Appalachians. The full extent of introgressive-type hybridization in evolution and animal

diversification is unclear. Unlike plants, hybrid animal taxa appear to be relatively rare. However, this

may be due to negative attitudes toward hybridization and the difficulty in detecting such examples

(Dowling & Secor 1997). Recently, Papilio joanae Heitzman (in the Ozarks) and P. hrevicauda Saunders

(in eastern Canada) of the machaon group of black swallowtails have been put forth as examples of taxa

of hybrid origin (Sperling 2003; Sperling & Harrison 1994). Einally, we caution that further detailed

studies are needed to establish the age and origin of P. appalachiensis. Presently, it cannot be ruled out

that P. appalachiensis is ancestral to P. canadensis. P. appalachiensis may have independently evolved

unique adaptive genes and launched pxoio-canadensis into the vast northern territories.

ACKNOWLEDGMENTS

We gratefully acknowledge the following people and thank them for their invaluable assistance: J. Mark

Scriber and Gabe Ording, Department of Entomology, Michigan State University, for electrophoresis analysis of

West Virginia specimens and engaging discussions; Richard Romeyn for providing specimens from Buck Creek,

North Carolina; Emily Romeyn for providing digital photographs; Amos Showalter for providing photographic

records of specimens from Seneca State Forest, West Virginia; and Eileen Mathias, Librarian, The Academy of

Natural Sciences of Philadelphia, and Charles Greifenstein, Librarian, American Philosophical Society,

Philadelphia, for assistance in pertinent literature research.

LITERATURE CITED

ALLEN, T.J. 1997. The Butterflies of West Virginia and Their Caterpillars. Pittsburgh, PA: University of Pitts¬

burgh Press, 388 pp.

CLARK, A.H. 1932. The butterflies of the District of Columbia and vicinity. U. S. NatT Mus. Bull. 157: 1-337.

_ 1936. The swallowtail butterflies. Smithsonian Institution Annual Report, 1935: 383-408.

CLARK , A.H., & L.F. CLARK. 1951. The butterflies of Virginia. Smithsonian Misc. Coll. 116(7): 1-239.

CLARKE, C. & E.M.M. CLARKE. 1983. Abnormalities of wing pattern in the Eastern Tiger Swallowtail

butterfly, Papilio glaucus. Systematic Ent. 8: 25-28.

CLARKE, C.A., & P.M. SHEPPARD. 1959. The genetics of some mimetic forms of Papilio dardanus, Brown, and

Papilio glaucus, Linn. Journal of Genetics 56: 236-260.

DOWLING, T.E., & C.L. SECOR. 1997. The role of hybridization and introgression in the diversification of

animals. Ann. Rev. Ecol. Syst. 28: 593-619.

EDWARDS, W.H. 1884. The Butterflies of North America. Vol. 2. Boston: Houghton, Mifflin and Company, 357

pp., 51 pi.

ENA (Elora of North America Association). 2000. Elora of North America. ENA Online. [Available at:

http://www.fna.org/ENA/ ]. Last downloaded: 9 November 2004.

GUNDER, J.D. 1927. Transition forms (Lepid., Rhopalocera). Ent. News 38: 263-271.

HAGEN, R.H., & R.C. LEDERHOUSE. 1984. Polymodal emergence of the tiger swallowtail, Papilio glaucus

(Lepidoptera: Papilionidae): source of a false second generation in central New York State. Ecol. Ent. 10(1):

19-28.

HAGEN, R.H., R.C. LEDERHOUSE, J.L. BOSSART, & J.M. SCRIBER. 1991. Papilio canadensis and P. glaucus

(Papilionidae) are distinct species. J. Lepid. Soc. 45 (4): 245-258.

HEITHAUS, P. 2003. Mosaic tiger swallowtail at Kenyon College. Ohio Lepid. 25(4): 41.

HONEY, M.R., & M.J. SCOBLE. 2001. Linnaeus’s butterflies (Lepidoptera: Papilionoidea and Hesperioidea).

Zoological Journal of the Linnaean Society 132: 277-399.

HOWARD, L.O. 1899. An abnormal tiger swallow-tail. Insect Life 7: 44-47, 1899.

ICZN (International Commission on Zoological Nomenclature). 1999. International Code of Zoological Nomen¬

clature. 4^^ ed. London: The International Trust for Zoology Nomenclature, 306 pp.

LINNAEUS, C. 1758. Systema Naturae. 10'^ ed. Vol.l Stockholm, 824 pp.

PAVULAAN, H.P., & D.M. WRIGHT. 2002. Pterourus appalachiensis (Papilionidae: Papilioninae), a new

swallowtail butterfly from the Appalachian region of the United States. The Taxonomic Report 3(7): 1-20.

RAUSHER, M.D. 1980. Host abundance, juvenile survival, and oviposition preference in Battus philenor.

Evolution 34: 342-355.

SCOTT, J.A. 1981. New Papilionoidea from North America. Papilio (New Series), No. 1, 12 p.

SCRIBER, J.M. 1990a. Two aberrant forms of the tiger swallowtail butterfly from the Great Lakes hybrid/transition

zone (Lepidoptera: Papilionidae). The Great Lakes Ent. 23(3): 121-126.

_ 1990b. Interaction of introgression from Papilio glaucus canadensis and diapause in producing

“spring form” Eastern Tiger Swallowtail butterflies, P. glaucus (Lepidoptera: Papilionidae). The Great Lakes

Entomologist 23(3): 127-135.

_ 1996. Tiger tales: Natural history of native North American swallowtails. American Entomologist

42(1): 19-32.

SCRIBER, J.M., M. DEERING, & A. STUMP. 2003. Hybrid zone ecology and tiger swallowtail trait dines in

North America, (pp. 367-391) In Boggs, C. L., W. B. Watt, and P. R. Ehrlich (eds.). Butterflies: Ecology and

Evolution Taking Flight. Chicago: Univ. of Chicago Press.

SCRIBER, J.M., & M.H. EVANS. 1988. Bilateral gynandromorphs, sexual and/or color mosaics in the tiger

swallowtail butterfly, Papilio glaucus (Lepidoptera: Papilionidae). J. Res. Lepid. 26(1-4): 39-57.

SCRIBER, J.M., M.H. EVANS, & D.B. RITLAND. 1987. Hybridization as a causal mechanism of mixed color

broods and unusual color morphs of female offspring in the Eastern Tiger Swallowtail Butterflies, Papilio

glaucus. (pp. 119-134) In M. Huettel (ed.). Evolutionary Genetics of Invertebrate Behavior. Gainesville: U. of

Florida.

SCRIBER, J.M., R.C. LEDERHOUSE, and R.V. DOWELL. 1995. Hybridization studies with North American

swallowtails, (pp. 269-281) In J. M. Scriber, Y. Tsubaki, and R. C. Lederhouse (eds). Swallowtail Butterflies:

Their Ecology and Evolutionary Biology. Gainesville: Scientific Publishers.

SIME, K.R., P.P. FEENY, & M.M. HARIBAL. 2000. Sequestration of aristolochic acids by the pipevine swallow¬

tail, Battus philenor (L.): evidence and ecological implications. Chemoecology 10: 169-178.

SPERLING, F. 2003. Butterfly molecular systematics: from species definitions to higher level phylogenies. (pp.

431-458) In Boggs, C.L., W.B. Watt, and P.R. Ehrlich (eds.) Butterflies: Ecology and Evolution Taking Flight.

Chicago: Univ. of Chicago Press.

SPERLING, F.A.H. & R.G. HARRISON. 1994. Mithochondrial DNA variations within and between species of

Papilio machaon group of swallowtail butterflies. Evolution 48: 408-422.

STRECKER, H. 1878. Butterflies and Moths of North America. Reading, PA: Press of B. F. Owen, 283 pp.

STRAUSBAUGH, P.D., & E.L. CORE. 1978. Flora of West Virginia. ed. Morgantown, WV: Seneca Books,

Inc., 1079 pp.

TYLER, H.A., K.S. BROWN, Jr., and K.H. WILSON. 1994. Swallowtail Butterflies of the Americas: A Study in

Biological Dynamics, Ecological Diversity, Biosystematics, and Conservation. Gainesville: Scientific

Publishers, 376 pp.

WEST, D.A., & CLARKE, C.A. 1988. Suppression of ihe black phenotype in females of the P. glaucus group

(Papilionidae). J. Res. Lepid. 26(1-4): 187-200.

[Editor’s note. A photo of the venter of the black P. appalachiensis female can be found at the TILS web site photos section.]

Visit The International Lepidoptera Survey on the World Wide Web at: http://www.tils-ttr.org

- and -

Join the discussion at our list serve on YahooIGroups at: TILS-Leps-Talk

Volume 6

18 April 2005

Number 2

ic Report L T J

PTERA SURVEY

The Taxonomic

)F THE INTERNATIONAL LEPIDOPTERA ,

IMMATURE STAGES OF COLIAS OCCIDENTALIS SULLIVANI

FROM OREGON (LEPIDOPTERA: PIERIDAE)

JACK HARRY

47 San Rafael Court, West Jordan, UT 84088

ABSTRACT: The imm ature stages of Colias occidentalis sullivani Hammond and McCorkle are described and figured. The

larval host is Lathyrus rigidus White.

Additional keywords: alexandra, Christina, Harney County, head capsule, larva, ova, pupa.

Colias occidentalis sullivani was described in 2003 by Paul Hammond and Dave McCorkle. The type

locality is near Ten Cent Lake in Harney County, Oregon. The immature stages reared in this study were

from the type locality.

MATERIALS AND METHODS

Oviposition: Three females were observed ovipositing on L. rigidus. All three eggs were placed on or

near new growth at the ends of the plant stems. Therefore, the newly hatched larvae would have tender

growth readily available.

Sprigs of the hostplant were put into water in a small container and then placed inside a gallon size

plastic container with a chiffon cover. Females collected at the type locality in May 2002 were put into

the gallon container for ovipositing. At first, indirect sunlight was tried but the females did not respond

by ovipositing. Some Colias species will oviposit under incandescent light but the sullivani females did

not. When the container was placed in direct sunlight, about 100 eggs were readily obtained. Each was

oviposited singly on leaves of the larval hostplant. Care was taken to not let the container overheat. The

sprigs remained in water and the eggs were left in place until shortly before hatching.

Rearing: Cut stems of the hostplant were placed in water in a small container and put inside a plastic

container with a chiffon cover. Just before the eggs were to hatch, the leaves with the eggs were placed

among the stems of the hostplant for rearing. The newly hatched larvae could easily find the fresh leaves.

When the cut stems needed to be replaced, the leaves the larvae were on were placed among fresh stems

and leaves. The majority of the larvae were reared on L. rigidus. A few larvae were reared on Lathyrus

brachycalyx Rydberg which was obtained near Salt Lake City.

The larvae used for measurements were set aside on the leaf they were on after they quit eating and

were preparing to molt. After the larvae molted, they were placed on fresh stems.

Diapausing larvae were put into a small plastic container which had many holes for air circulation.

The container with the larvae was put into a screen cage in a shady spot in the yard until there was snow

in the mountains - which was in November in the Wasatch Mountains near Salt Lake City. Then the

container was put in a cage in the snow for the winter. The post diapause larvae take a few days to start

eating after put into room temperature. Many larvae did not survive the winter.

Observations: Morphological observations and measurements of the eggs, first instar larvae and head

capsules were made with the aid of a stereomicroscope and a 0.1 mm scale. Length measurements of the

second instar and larger were made with a millimeter scale. Measurements of the head capsule width are

of the molted head capsule. Measurements of the length were made at pre-molt time (except the fifth

instar). Length of the fifth instar larvae is given for the mature larvae during the most common resting

position. When feeding, the larvae are often longer. N=10 for the egg, head capsule, and length measure¬

ments.

DESCRIPTION

Eggs: The eggs (Fig. 3) are typical of the genus Colias. The eggs are fusiform in shape, 0.48 mm wide

(range 0.46 to 0.50) and 1.35 mm long (range 1.20 to 1.47). Each egg has longitudinal ribs with small

transverse ridges. The top is rounded and contains the micropyle. Eggs are creamy white when

oviposited and become orange with a creamy tip within 2 days. Prior to hatching the eggs exhibit a black

tip, the head of the larvae can be seen through the shell. Eggs hatch in about 5 days at room temperature.

Larvae: C. o. sullivani has five larval instars. The larvae have one lateral stripe on each side. Second

through fifth instar are similar and differ only in size, there is one exception which is noted in fourth

instar. Also, the fourth instar larvae are dark blue or purplish just after molting but become green when

feeding and growing. The appearance of the first, fourth and fifth instars and pupa is described.

First instar: Neo-natal body: brown then turns yellow green in a few hours. Head: black with tiny white hairs.

Body: yellow green and more yellowish posterior; mid-dorsal stripe: dark brown, faint, and yellowish posterior;

on each side of the body there are three white hairs on each segment except first segment which has five white

hairs; many black dots on yellow body under 40 X magnification; thoracic legs: black, spiracles: black. Length:

3.5 mm (range, 3.4 to 3.7). Head width: 0.36 mm (range, 0.35 to 0.37).

Second instar: Length: 5.46 mm (range, 5.0 to 5.9). Head width: 0.56 (range, 0.53 to 0.58).

Third instar: Diapause length: 7.34 mm (range, 6.8 to 7.8). Post diapause length: 4.81 mm (range, 4.5 to 5.1).

Molt length: 8.03 mm (range, 7.5 to 8.5). Head width: 0.81 mm (range, 0.78 to 0.85).

Fourth instar: Hairs: on second and third instar, all are white; on fourth instar, black on upper part of head and

body and white on lower part. Length: 14.2 mm (range, 13.2 to 15.3). Head: width 1.34 mm (range, 1.28 to 1.45).

Fifth instar: Head: green with many black spots which bear black or white hairs; eyes and mandibles black.

Body: green with many black spots which bear short black or white hairs, hairs are black dorsally and white

ventrally on body and head; thoracic legs: green with a bit of brown on the outside; mid-dorsal stripe: dark green

except near head where it is light green; lateral stripe: white with a bit of pink on some segments; spiracles: white;

black spots on body ringed with light green. Length: about 30 mm. Head width: 2.5 mm (range, 2.45 to 2.6).

Pupa: Head: green with front a darker green, light green horizontal line in middle of head. Body: green, lighter

green posterior: mid-dorsal stripe: dark green; lateral stripe: light green and is expressed on wing case; spiracles:

light green; sublateral stripe: brown on three segments immediately behind wings; small black spot between

lateral stripe and sublateral stripe on two segments immediately behind wings. Length: 19.9 mm (range, 18.0 to

21.5); width: .49 mm (range, 4.5 to 5.5); height: (dorsal to ventral) 6.1 mm (range, 5.5 to 6.5). As the pupae

develop the colors and patterns of the adult become visible. The posterior end of the pupa is attached to a silk pad

on the substrate and the body is held loosely to the substrate with a silk girdle.

Behavior: First through third instar larvae eat the spongy mesophyll from the leaf and leave the veins and

membrane. The first three instar larvae rest on the upperside of a leaf along the mid vein. Fourth and fifth

instar larvae eat the entire leaf. Partially grown third instar larvae diapause. Diapausing larvae become

light brown. Post-diapause larvae slowly turn green as they feed. Mature larvae leave the hostplant and

wander off to pupate. Occasionally one will pupate on a stem of the hostplant in the rearing container.

RESULTS

There was considerable loss of pre-diapause larvae due to unknown reasons. Colias larvae are very sus¬

ceptible to disease so that may have been part of the problem. There were 24 larvae reared on L. rigidus

and three on L. brachycalyx that diapaused. Fifteen of the L. rigidus larvae and one of the L. brachycalyx

larvae survived winter diapause and began eating in the spring. Eleven adults were obtained from the L.

rigidus reared larvae and one from the L. brachycalyx reared larva.

The one post-diapausal larva reared on L. brachycalyx began eating 15 April, pupated 2 May, and the

adult emerged on 11 May. This adult was the largest of all the reared adults. It is a female equal in size

to the smallest wing caught adult female in my collection. Therefore, this adult is 9 percent smaller than

the norm. The average of all reared adults is 10.5 percent smaller than the wing caught adults.

Ken Hansen, Jacque Wolfe, and I attempted to rear three subspecies of C. occidentalis (chrysomelas,

occidentalis & sullivani) on Hedysarum mackenziei. In all attempts, the larvae died before becoming

second instar. This evidences that C. occidentalis cannot use H. mackenziei as a larval host.

DISCUSSION

The cut stems of Lathyrus and Vida do not transport water to the leafs very well. This seems to be the

case with many other legumes as well. The larvae feeding on cut stems of the two Lathyrus species used

would start to leave the plant and wander around the container two or three days after they were given

freshly cut stems. This indicated that they were getting discontented with the condition of the plant. This

is probably the reason for the small size of the adults. After four or five days the leaves of the plant

seemed to be in good condition, but apparently they were not good enough for the larvae. Therefore, it

would be preferable to use growing plants; either planted or potted hostplants would probably suffice.

In 2003, some larvae were placed on L. rigidus at the type locality and a sleeve put over the plant.

Ten diapausing larvae were later recovered. These larvae all entered diapause in the second instar. 2003

had a dry spring and the eggs were oviposited after the middle of the flight. The plant leaves probably

desiccated so early that the larvae could not continue beyond second instar. The rearing indicates that the

larvae will not exceed third instar even if they still have edible plant.

The larvae of Colias occidentalis chrysomelas (pers. comm. Ken Hansen, Jacque Wolfe) and C. o.

pseudochristina (pers. comm. J. Harry, J. Wolfe) turn light brown when they diapause. This evidences

that sullivani is indeed a subspecies of occidentalis.

The larvae of Colias alexandra edwardsi (pers. Obs. J. Harry), Christina krauthi (pers. comm. J.

Wolfe), and Christina kluanensis (pers. Obs. J. Harry) do not turn brown when they diapause. This

indicates that they are not conspecific with occidentalis. Since the larvae of no other North American

Colias change color when they diapause; this is considered to be a significant genetic character.

ACKNOWLEDGEMENT

Thanks are extended to Dave McCorkle for reviewing this manuscript and providing helpful suggestions.

REFERENCE

HAMMOND, P. C. and D. V. McCORKLE, 2003. A New Desert Subspecies of Colias occidentalis (Pieridae) from

Southeastern Oregon. J. Lepid. Soc. 57:274-278.

Figs. 1-14. Developmental stages of Colias occidentalis sullivani. Figs. 1/6. Male: topotype, D/V. Figs.

in. Female: topotype, D/V. Fig. 3. Ova after three days. Fig. 4. Larva in process of hatching. Fig. 5.

First instar larva. Fig. 8. Second instar larva. Fig. 9. Diapausing third instar larva. Fig. 10. Newly emerged

fourth instar larva. Fig. 11. Fifth instar larva. Fig. 12. Prepupa. Fig. 13. Pupa. Fig. 14. Pupa with female

adult ready to emerge. All photographs by Jack Harry. Figs. 1 & 2 actual size. D: dorsal. V: ventral.

Volume 6

18 April 2005

Number 3

IMMATURE STAGES OF COLIAS JOHANSENI

FROM ARCTIC CANADA (LEPIDOPTERA: PIERIDAE)

JACK HARRY

47 San Rafael Court, West Jordan, UT 84088

ABSTRACT: The immature stages of Colias johanseni are described and figured. The probable larval host plant is Hedysamm

mackenziei, a common arctic Legume.

KEYWORDS: Colias hecla. Northwest Territories, Nunavut, Coppermine, drumlin, fusiform.

The purpose of this paper is to describe the immature stages of Colias johanseni Troubridge and Philip,

1990. C. johanseni was first collected by Frits Johansen at Bernard Harbor, Northwest Territories, Canada

in 1916 during the Canadian Arctic Expedition. Bernard Harbor is 107 km north of Coppermine along the

coast of the Arctic Ocean. Coppermine, a native village, now goes by the name Kugluktuk and this area is

now in the Nunavut Territory. One male C. johanseni was taken by Johansen. Colias johanseni was next

collected by Troubridge and Philip at Bernard Harbor in 1988. They described this taxon from these

specimens. Colias johanseni was next collected by this author during the summer of 1998 at Bernard

Harbor and Cape Kendall. One male was taken at Cape Kendall which represents the only known location

other than Bernard Harbor. Cape Kendall is 87 km south of Bernard Harbor and 20 km north of

Kugluktuk.

MATERIALS AND METHODS

Oviposition: One female Colias johanseni was collected, at Bernard Harbor, in copulo with a male C.

johanseni. The pair was safely contained until after copulation was completed. The female was put into a

plastic container with a screen cover. Sprigs of an Astragalus species for an oviposition substrate were

inserted into water inside the container. The female would not oviposit on the Astragalus so the

Astragalus was replaced with Hedysarum mackenziei Richards on which she readily oviposited.

Troubridge and Philip reported H. mackenziei as the probable larval host plant by association with the

females. The container was placed in indirect sunlight for the female to oviposit. The female oviposited

eggs singly on the upper side of a leaf. Ova were removed in two or three day intervals and placed in a

container to await hatching.

Rearing: The larvae were reared on cut stems of H. mackenziei that were placed in water to retain

freshness in a plastic container with a fine mesh cover. Larvae were disturbed only when it was apparent

that they had quit eating and were preparing to molt or pupate.

DISCUSSION

From the time the post diapause larvae started eating to emerging as adults was 16 days. All the larvae

diapaused as fully grown third instar, so it appears that they would not exceed that stage in nature even if

weather permitted. It is quite probable that most larvae do not reach that stage in nature due to the onset of

cold temperatures. Overwintering Colias eurytheme in my Utah yard, began to eat on February 24 even

though it was a rather cold winter and there were pupae by the end of March. Presumably post diapause

C. johanseni larvae begin to eat early and develop fairly rapidly even when the weather is still quite cool.

Only six adults were obtained from this rearing, three males and three females. Most of the larvae died

from overwintering. The three males have the androconial patch on the hind wing. This androconial patch

is the primary distinction between C. johanseni and C. hecla. The three females are all intermediate in

color and pattern with all females of both C. johanseni and C. hecla that were collected.

The natural life history, the mating and taxonomic relationship with C. hecla at Bernard Harbor remains

to be studied. Due to the remote location and arctic conditions where C. johanseni inhabits natural studies

would be rather difficult.

ACKNOWLEDGEMENTS

I thank James Scott for reviewing this manuscript and providing helpful suggestions. I also express gratitude to

Aime Ahegona, a Kugluktuk native, for transporting me to Bernard Harbor with his boat, for the use of his cabin at

Bernard Harbor, and companionship while at Bernard Harbor.

LITERATURE CITED

Troubridge, J. and K. Philip. 1990. A new species of Colias from Arctic Canada. Canadian Entomologist, 122:15-

20 .

Figs. 1-12 Developmental stages of Colias johanseni. Fig. 1. Male: topotype. Fig. 2. Female: topotype.

Fig. 3. Ovum. Fig. 4. First instar larva. Fig. 5. Fourth instar larva. Fig. 6. Diapausing third instar larva.

Fig. 7. Fifth instar larva. Figs. 8 & 9. Freshly emerged reared female. Fig. 10. Pupa with female adult

ready to emerge. Fig. 11. Pupa. Fig. 12. Prepupa. All photographs by Jack Harry. Figs. 1 & 2 actual size.

Volume 6

18 April 2005

Number 4

\rhe Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

IMMATURE STAGES OF ESTIGMENE ACREA

FROM GUATEMALA (LEPIDOPTERA: ARCTIIDAE)

JACK HARRY

47 San Rafael Court, West Jordan, UT 84088

ABSTRACT: The imm ature stages of Estigmene acrea Drury are described and figured. The larval host utilized was Brassica

oleracea L. var. botrytis (cauliflower).

KEYWORDS: Guatemala, Chimaltenango Department, fasciated inflorescence, agricultural pest.

Estigmene acrea Dury, 1773 is a well known Tiger Moth that ranges from southern Canada to Honduras. It’s

common name, Saltmarsh Caterpillar, disguises the fact that it is a well known polyphagus agricultural pest.

Nonetheless, it is found very sporadically over this wide range and is generally not often thought of as a “pest

species” among lepidopterists. http://www.npwrc.usgs.gov/resource/distr/lepid/moths/usa/1824.htm is the link

to its uses range map.

The immature stages in this study are from the southwest side of the town of Teepan, Chimaltenango Depart¬

ment, Guatemala. This location is an agriculture area with many farms at an elevation of 2330 meters. A

farmer gave the author two cluster of ova - one had 54 eggs and the other had 36 eggs. The eggs had been

oviposited on leaves of cauliflower {Brassica oleracea L. var. botrytis) which was then utilized in rearing

the larvae for this study.

MATERIALS AND METHODS

Rearing: Cauliflower is easy to obtain in grocery stores and markets. Most of the leaves of the cauliflower

were cut from the stem. The stem of the cauliflower was placed into a small container with some water and

the gaps between the stem and the container were plugged with wads of plastic. The cauliflower was then

placed into a gallon size plastic container. The container had a lid of chiffon material to allow for some air

circulation and avoid moisture accumulation.

The leaves the ova were on were trimmed to nearly the size of the egg cluster. When the eggs began to hatch

the clusters were placed on the cauliflower. The larvae could easily find the fresh plant. When the larvae were

small, a container of cauliflower was used for each cluster. When the larvae were large, about 15 were put

into a container. They appeared to be somewhat crowded but that didn’t seem to be a problem. From the

second instar on, each time the larvae molted they were transferred to a fresh head of cauliflower. Those larvae

utilized for measurements were measured for length before being disturbed, then they were placed in a

separate container to molt. This made the head capsule easy to find, especially during the small instars.

Observations: Morphological observations and measurements of the ova, first instar larvae, and head capsule

were made with the aid of a stereomicroscope and a 0.1 nun scale. Measurements of second instar larvae and

larger, and the pupae were made with a millimeter scale. Measurements of the head capsule width were made

of the molted head capsule. Measurements of the length of the larvae were made when the larvae had stopped

eating and were preparing to molt. Maximum length of the sixth instar larvae is given for the mature larvae in

the most conunon resting position. N=10 for all of the measurements.

DESCRIPTION

Eggs: Eggs are light blue, smooth, and spherical with a diameter of 0.6 mm.

Larvae: The larvae have six instars.

First instar: Neo-natal larvae: 1.8 mm long. Head: black with black hairs and a light brown triangle just above

mouth. Body: creamy white, each segment with five black spots on each side; a long black hair arises from each

spot, each black spot on first segment has two hairs; a large black spot above anus; light brown areas on each

segment; thoracic legs: black and white. Length: 3.8 mm (range: 3.6 to 4.0); width of head capsule: 0.41 mm

(range: 0.40 to 0.43).

Second instar: Similar to the first instar. Length: 6.6 mm (range: 6.2 to 7.0); width of head capsule: 0.56 mm

(range 0.55 to 0.58).

Third instar: Similar to the first instar except that each spot has several hairs. Length: 10.7 mm (range: 10.0 to

11.5); width of head capsule 0.91 mm (range 0.84 to 0.97).

Fourth instar: Head: black with black hairs, eyes white; frontoclypeus light brown or creamy, somewhat trans¬

parent on the molted head capsule. Body: creamy white with brown areas that form a marbled wide stripe on each

side of mid-dorsum; marbled brown laterally and underneath, some individuals appear more brown than others;

creamy white diagonal stripe on each segment from middle of body down to legs, this diagonal stripe is also on

the fifth and sixth instars and barely apparent on the third instar; the spots give rise to many hairs; two spots

closest to mid-dorsum are black, others are creamy white or light brown; some hairs black, some light brown, and

some white. Length: 17.1 mm (range: 16.0 to 18.0; width of head capsule: 1.56 mm (range: 1.48 to 1.58).

Fifth instar: Head: similar to fourth instar. Body: primarily black marbled with white, each segment with six

bumps on each side, the two bumps nearest mid-dorsum are black, others light brown; posterior two segments

have one less bump; each bump gives rise to many hairs which most are light brown and some are black. Length:

26 mm (range: 25 to 27), width of head capsule: 2.41 mm (range: 2.34 to 2.47).

Sixth instar: Similar to fifth instar except body somewhat darker, still marbled. Some larvae have mostly reddish

brown hairs, others mostly black hairs. Length: approximately 45 mm; head capsule data not obtained.

Pupae: Primarily cylindrical, rounded on both ends with a sharp point on the posterior end, and dark brown.

Length: 19.5 mm (range: 18.5 to 21); diameter 8.5 mm (range: 8 to 9)..

Behavior: Larvae eat the fasciated inflorescence and leaves, but seem to prefer the inflorescence. Third instar

larvae wander around to find a place to molt. Thus, it appears they are gregarious for the first and second instar

and solitary for the third through sixth instars. However, the larger instars did not have a problem being with other

larvae. Hairs of the last instar are loosely formed around the pupa, and may also be loosely attached to a substrate.

RESULTS

The eggs hatched on December 10, 2004. Each instar took 3-4 days to complete except the sixth instar which

took 10 days to pupation. The adults started emerging on January 17, 2005 with sixty-eight adults obtained.

Acknowledgements

Thanks are extended to Cristina Bailey for determination of the species. Thanks also to Robert Mower and Alan Myrup for

reviewing the manuscript and providing suggestions.

Figures 1-10. Developmental stages of Estigmene acrea. Fig. 1. Male: from rearing. Fig. 2. Female: from

rearing. Fig. 3. Egg mass. Fig. 4. Eggs mass with emerging first instar larva. Fig. 5. Second instar larvae.

Fig. 6. Third instar larva. Fig. 7. Fourth instar larva. Figs. 8. Fifth instar larva. Fig. 9. Sixth instar larva.

Fig. 10. Pupa. All photographs by Jack Harry. Figs. 1 & 2 enlarged.

Volume 6

20 July 2005

Number 5

The Taxonomic

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

TAXONOMIC ANALYSIS OF THE GENUS MEGISTO (SATYRIDAE)

IN THE EASTERN UNITED STATES.

PART I: TYPES, TYPE LOCALITIES,

AND TAXONOMIC RELATIONSHIPS OF THE AVAILABLE NAMES.

RONALD R. GATRELLE ^

126 Wells road, Goose Creek, South Carolina 29445

ABSTRACT. The genus Megisto Hiibner, 1819 in the eastern United States and southeastern Canada is currently comprised

of the original epithets Papilio eurytus Fabricius, 1775, Papilio cymela Cramer, 1777, Papilio eurytris, Fabricius, 1793,

and Neonympha eurytris viola Maynard, 1891. Because the eastern Megisto is considered by some, including the present

author, to contain two or more sibling species, the types and type localities of each of these names were studied. Where no

type was found and no type locality fixed, typification was established by lectotypification or neotypification and a type

locality was fixed. The taxonomic relationship of these taxa were then overviewed. No syntypic specimens were located for

P. cymela and thus a neotype was designated and the type locality fixed as: Colleton County, South Carolina. P. eurytus and

P. eurytris are different spellings of the same Fabrician taxon. Syntypes of P. eurytus/eurytris were found in Glasgow,

Scotland, and considered to represent a different species than P. cymela. A lectotype was designated for P. eurytus/eurytris

and the type locality fixed as Berkeley County, South Carolina. The type specimens of N. eurytris viola were not located but

are possibly at the Museum of Comparative Zoology, Harvard. Its type locality is Enterprise, Volusia County, Florida.

FOUNDATIONAL EASTERN MEGISTO NAMES

The first taxon to be described in the Modem genus Megisto Hiibner, 1819 was Papilio eurytus

Fabricius, 1775. This name is however a junior primary homonym of Papilio eurytus Linnaeus, 1758 (a

central African nymphalid known today as Pseudacraea eurytus), and as such, is an available but perman¬

ently invalid name in zoological nomenclature. The next oldest name is Papilio cymela Cramer, 1777. In

1781 and 1793 Fabricius again published on the taxon he named eurytus. In both instances, he repeated

verbatim his 1775 original description: however, in the 1793 publication the spelling was given as

eurytris. The 1793 text also differed in that therein the origin of the specimens was attributed to Banks

while in the 1775 and 1781 texts they were attributed to Hunter. In both 1781 and 1793, Fabricius synon-

ymized his taxon with Cramer’s cymela, establishing that he then viewed these as the same taxon.

The situation today is that the oldest available name cymela has long and consistently been applied

to a ventrally light tan, late spring to early fall entity, flying widely in the eastern states and southeast

Canada. This name is thus established by prevailing usage to a specific taxon regardless of what the original

types may have been (or are) and what the original description states. In these investigations, and by this

author’s taxonomic concept, the two syntypes of eurytus/eurytris represent a different species than cymela.

^ Staff Researcher, The International Lepidoptera Survey, Goose Creek, South Carolina.

Thus these three names, in this taxonomic view, are not synonyms. There is thus a difficult and unusual no-

menclatural situation with the name eurytris. The author therefore presented this situation to the Inter¬

national Commission of Zoological Nomenclature (ICZN) official World Wide Web list serve for advice.

The name eurytris can not be a replacement name for the invalid homonym eurytus as Fabricius did

not indicate that this was what it was intended to be in any way (Article 72.7). In fact, there is nothing to

lead one to think that Fabricius even thought of his eurytus as a homonym to Papilio eurytus Linnaeus,

1758. After much discussion, it was deduced that the most likely explanation for the different spelling in

1793 was that it was simply a printer’s misspelling - faulty type setting.

The etymology of eurytus is a reference to king Eurytus of Greek mythology. Fabricius was a very

learned person and the informal opinion in the ICZN discussion was that there is no “proper” Latinized

grammatical meaning with the word ""eurytris,"" and, thus, there is no reason to think he would have spelled

this any other way than eurytus. Thus, it is an incorrect subsequent spelling, and, as such, is an unavailable

name (Article 33.3). However, the epithet eurytris has a history of usage prior to about 40 years ago. The

name therefore becomes available as a correct original spelling per Article 33.3.1 with the correct citation

being Papilio eurytris Fabricius 1775 - not 1793. The only alternative is to consider both spellings as

invalid names which would necessitate the erection of a totally new name for this species. This would seem

to only add more confusion to what is already a nomenclatural mess. In addition, it is to be noted that

Maynard’s 1891 description of viola was correctly authored (Gatrelle, in press) as a subspecies of eurytris

(Neonympha eurytris viola). In that work, he also first gave a species description of nominate Neonympha

eurytris. To some, eurytris might be considered as available by adoption from Maynard’s 1891 taxonomic

act. In this view, Neonympha eurytris Maynard, 1891 would still have Fabricius’s 1775 specimens as

syntypes. The least disruptive way to deal with these names is to consider eurytris available per 33.3.1.

If indeed, the 1793 spelling of eurytus as eurytris was a typesetter’s error, then there is an issue of

why the type specimens were attributed in 1793 to Banks and not Hunter as in 1775 and 81. The explanation

here is the same as before, a typesetter’s error. The clue to this is that in the three identical 1775, 1781 and

1793 texts the type locality is stated as Jamaica. That being consistent, the origin would logically have been

the same collector / collection. In this author’s search for syntypic material, no Banks specimens were

found, just the two syntypes in the Hunterian collection in the University of Glasgow, Scotland. (Even if

Banks specimens of this taxon were found they would not be syntypes as only those of 1775 are eligible.)

A third error with these Fabrician names is the type locality. This taxon does not occur in Jamaica.

This is addressed convincingly in the Miller & Brown Lep. Soc. checklist note #627. There was a switch¬

ing of two type localities, and thus the correct type locality of these names is “Carolina”. This is most

certainly today’s South Carolina (Gatrelle, 2000) as several taxa described by various old workers are

from the vicinity of the busy Colonial port and cultural center of Charleston. But the best evidence is the

syntypes themselves as they are identical to specimens of this taxon which are still common today just

inland in the coastal area of South Carolina, west and north of Charleston.

Thus, there are two Megisto species in the east originally described under the names Papilio cymela

Cramer, Mil md Papilio eurytris Fabricius, 1775. Today, these dLVQ Megisto cymela md Megisto eurytris.

ESTABLISHING TYPES AND TYPE LOCALITIES

Because there are two sibling species involved (Gatrelle, in press), it is essential that they be

established objectively in accord with the Code’s Principle of Typification. The original description of

eurytris and its syntypes render it fairly self explanatory. Herein, the better conditioned of the syntypes is

simply designated as the lectotype of both Papilio eurytus Fabricius, 1775 and Papilio eurytris Fabricius,

1793. The type locality is here further restricted to Goose Creek, Berkeley County, South Carolina. A red

label with the words: Lectotype, Papilio eurytus and Papilio eurytris, has been sent to the Hunterian

collection. University of Glasgow to be placed on this specimen (Figs. 1-2).

The author attempted to locate syntypic material of Cramer’s cymela at the Natural History Museum,

London and the Natural History Museum, Leiden, The Netherlands. The staff at the Leiden facility

confirmed that there are none there, and staff at the NHM, London likewise could not locate any. Due to the

prevailing usage of the name Megisto cymela, and the presence of a sibling species in the same region in

which it widely occurs, a neotype needs to be established for all of the reasons stated in the ICZN Code

under Article 75.3 which by this reference are considered here quoted and met. The “exceptional” need for

this as “considered” (75.1) by the author, is that the name cymela can not be objectively determined by the

brief words of the original description nor by the stylized (^fictional) artistic rendering in the OD. The

dorsal painting looks like all eastern Megisto and the ventral painting (both reproduced on header) does not

look like any known Megisto anywhere. No population has the depicted extremely light buff yellow field

on the outer HW. However, in nature in the clarified type locality, cymela does have a noticeably lighter

outer one third on both ventral wings (Fig. 4) (esp. in females), while the sympatric eurytris tends to be

concolorous rich warm brown as can be seen in the lectotype (Fig. 2). In other words, the taxon cymela can

not be objectively determined for either taxonomic or systematic purposes by the OD alone.

The OD type locality of Cape of Good Hope (South Africa), is an obvious error as the Cramer

figures of this taxon are clearly based on eastern North American Megisto. An examination of the type

localities of the taxa Cramer described from the eastern U.S. shows that they ranged from New York

southward along the Colonial seaboard. Thus, the types of cymela could have come from any number of

places in NY, PA, VA or SC. Because these two species are 1) sympatric and 2) easily distinguished just

by phenotype and phenology in coastal South Carolina, including the type localities of both, the cymela

neotype is chosen from the mid coastal region at Jacksonboro, Colleton County, South Carolina (Figs. 3-4).

It bears a red label with the words: Neotype, Papilio cymela Cramer 1777. It is deposited in the collection

of The International Lepidoptera Survey (TILS) in Goose Creek South Carolina.

The taxonomic acts herein lay an essential and objective foundation for the ongoing taxonomic and

systematic investigations into the Megisto of the eastern U.S. and southeastern Canada. All published re¬

search on this genus in the eastern region up to this point, has been speculative simply because without

definitive typification no one could know exactly just what taxon they are commenting upon. The estab¬

lishment of proximal type localities for both taxa where they occur sympatrically and are easily distin¬

guished as species (Gatrelle, in press) greatly eliminates the subjectivity of human opinion from the study of

their systematics and taxonomy.

's H

Figures 1-2. Lectotype: Papilio eurytris [Papilio eurytus], dorsal 1, ventral 2; Megisto eurytris Fabri-

cius, 1775. Colors and markings identical to modem fresh topotypes. Figures 3-4. Neotype: Papilio

cymela, dorsal 3, ventral 4; Megisto cymela Cramer, 1777: 5 June 1992, Jacksonboro, Colleton Co.,

SC., leg. Gatrelle. Illustrations are natural size. Photos: 1-2 by Geoff Hancock, 3-4 by Joseph Mueller.

ACKNOWLEDGEMENTS

My thanks to Geoff Hancock, Curator of Entomology, Hunterian Museum, University of Glasgow, Scotland, for

locating and photographing the two syntypes of Papilio eurytus/eurytris; to Rienk de Jong of the Natural History

Museum, Leiden, The Netherlands for confirmation that no Cramer types of cymela reside there, and for scanning

and digitizing the OD figures of cymela; to Kim Goodger of the Natural History Museum, London for confirming that

Cramer syntypes of cymela were not located there by her; to Jonathan Pelham and the ICZN list serve, esp. Martin

Spies, for nomenclatural and Code relative discussions; to David Wright for peer review and important contributions

and advice.

LITERATURE CITED

CRAMER, P. 1777. De Uitlandsche Kappelen. 2:55.

LABRICIUS, J.C. 1775. Systema Entomologiae. Llensburgi and Lipsiae. pg. 487. 194.

_. 1781. Species Insectorum. II. Hamburg! and Kilonii. pg. 65. 891.

_. 1793. Entomologica Systematica Emendata Et Aucta. III. Hafniae. pg. 157. 485.

GATRELLE, R.R. in press [2005]. Taxonomic Analysis of the Genus Megisto (SATYRIDAE) in the Eastern United

States. Part II: A Case for the Recognition of Three Species of Megisto in Coastal South Carolina. Description of

a New Megisto Species and a New Subspecies. The Taxono mi c Report. The Int. Lepid. Survey. Vol. 6.

_ . 2000. A New North American Swallowtail Butterfly: Description of a Relict Subspecies of

Pterourus troilus (PAPILIONIDAE) from the Southern Tip of Elorida. The Taxonomic Report. The Int. Lepid.

Survey. Vol. 2:4. 1-14.

INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE. 1999. International Code of Zoo¬

logical Nomenclature, Fourth Edition (1 January 2000). London, UK: The Int. Trust for Zoo. Nom. 306 pp.

MAYNARD, C.J. 1891. Manual of North American Butterflies. Pages 109 (text), 113 (Fig. 36) original description of

Neonympha eurytris viola. Boston. De Wolfe, Fisk & Co.

MILLER, L.D., and F.M. BROWN. 1981. A Catalogue/Checklist of the Butterflies of North America North of

Mexico. Lepidopterists’ Society Mem. 2:I-vii, 1-280.

The Taxonomic Report

is a publication of The International Lepidoptera Survey (TILS).

(A Tax Exempt Non-Profit Scientific Organization)

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 PAVULAAN^

P.O. Box 1124, Herndon, VA 20172

AND

DAVID M. WRIGHT

100 Medical Campus Drive, Lansdale, PA 19446

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.)

By early May, lucia individuals showed signs of wear and their numbers began to fall. It was at

this time that a seeond irruption of Azures appeared. This emergenee eonsisted primarily of very light-

ventered individuals, some appearing almost white on the first day of emergenee and eontrasting sharply

with the darker eoarsely-patterned lucia phenotype. These light-ventered Azures were neither as eommon

nor as widespread as the earlier lucia flight, and appeared in only a few loeations sueh 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 pateh, reminiseent of the earlier ''lucia'' forms. The

initial thought was these were f "neglectamajor Tutf’, although the biology of neglectamajor was poorly

understood at the time. Klots (1951) wrote, “In some regions oeeurs a partial seeond brood {neglecta¬

major), large, very bright eolored above, white beneath, with elear-eut markings. This is “spotty” in its

oeeurrenee...” Swayed by Klots’ deseription, the Rhode Island light phenotype was misidentified as C

neglectamajor by Pavulaan (1985). Reginald Webster (pers. eom., 1984) suggested the unique Rhode Is.

butterfly may represent a new speeies, eiting erroneous reports that neglectamajor utilized Viburnum

acerifolium L. (Shapiro, 1966; Opler & Krizek, 1984), a plant that was relatively searee at Great Swamp.

It was reasonably eoneluded that V. acerifolium eould not support the large population of what was

believed to be neglectamajor at this loeation. The first evidenee of possible host assoeiation eame when

several females were flushed from a young P. serotina tree in Warwiek, R.I. on May 22, 1984, though it

was not immediately obvious that the speeies 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 maeulations. Their oeeurrenee was very spotty and eolleetively

their flight was the least eommon of the three flights observed. Popularly thought to be the summer form

of a spring speeies, this notion seemed unfounded beeause C. neglecta individuals were seen in far fewer

numbers and in loeations that often did not eorrespond with the earlier flights. A female was found

ovipositing on White Meadowsweet Spiraea alba var. latifolia (Ait.) along a utility powerline near

Cranston, Providenee Co., on July 17, 1984. Beeause this host and eherry {Prunus sp.) were members of

the same plant family (Rosaeeae), a link seemed possible, but oddly no Azures were found at this loeation

in May. At this point a strong suspieion emerged that the light-ventered May and July flights were

separated and represented different speeies. Shortly thereafter, rearing studies established that the light-

ventered May population was univoltine and reproduetively separated from summer flights of C.

neglecta. Similar studies also established C. lucia was univoltine, although this was not entirely

unexpeeted sinee 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.

C. lucia Auct. C. serotina C, neglecta

Figs. 4-6. Sympatric Celastrina in northeastern United States (spread specimens). SS 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. (DA^). Photos by David

Wright.

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, ‘T 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 19^^ 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, Eintner (1875) in New York, & Saunders (1875) in Ontario challenged

Edwards’ concept and reported lucm 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; Eayberry et al, 1996;

Thomas, 1996; Allen, 1997).

C. neglectamajor

Figs. 7-16. Light-ventered Celastrina in eastern United States. 6'6', $ $ (DA^)- 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. _

C. neglecta

spring phenotype

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 argiolns 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 argiolns (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

argiolns 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].^ 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 argiolns 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 argiolns (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 argiolns (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,

Fautre en fm 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”; PI. 24, row 6 (photo)] [OHIO], Celastrina argiolns 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

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,

PENNSYEVANIA, 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 ISEAND, 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 innommC’ (sp.) in Handfield (1999) [“Selon Wright

(1995) (ensemble nomme ‘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 E. occurs further north (than C. idella} (photo)] [NEW JERSEY, PENNSYE VANIA, 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 ISEAND]. 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].

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 23'’^ mean, n = 40). MASSACHUSETTS: May 3 - June 26 (May 27* mean,

n = 118). RHODE ISLAND: May 2 - June 22 (May 20* mean, n = 698). PENNSYLVANIA: May 16 - June 28 (May 29*

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.

Figs. 18-21. Cherry galls and Celastrina serotina immatures. Fig. 18. Old growth field with Black Cherry {Prunus serotind)

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 4* 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 E. (so. New England & Eong 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 (E.), Colias philodice Godart, C. eurytheme Boisduval, Lycaena phlaeas (E.), Callophrys

(Mitoura) gryneus (Hiibner), C. (Deciduphagus) augustinus (Westwood), C. (Deciduphagus) henrici (Grote & Robinson), C.

(Incisalia) niphon (Hiibner), Chlosyne harrisii (Scudder), Phyciodes tharos (Drury), P. cocyta (Cramer), Nymphalis antiopa

(E.), L. arthemis arthemis (Drury), L. a. astyanax (Fab.), Megisto cymela (Cramer) (type 1), Coenonympha tullia (Muller),

Erynnis icelus (Scudder & Burgess), E. brizo (Boisduval & Ee Conte), E. Juvenalis (Fab.), E. horatius (Scudder & Burgess),

Poanes hobomok (Harris), ?Lndi Amblyscirtes hegon (Scudder).

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-Pele, 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 Riviere; 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; EES COLLINES-DE-

L’OUTAOUAIS (Regional County Municipality): Chelsea; EES 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 COTE-DE-GASPE (Regional County Municipality, formerly Gaspe-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, Colonic; 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.:

Rochester; NASSAU CO.: Bethpage State Park; ONEIDA CO.: Trenton Falls; QUEENS CO.: Corona; ST. EAWRENCE CO.:

Madrid, Morristown; SCHENECTADY CO.: Schenectady, Niskayuna; SCHUYEER CO.: Cayuta Eake; SUFFOEK CO.:

Babylon, Bay Shore, Brentwood, Brookhaven, Commack, Deer Park, East Islip (Heekscher State Park), East Moriehes,

Eastport, Edgewood, Great River, Manorville, Middle Island, North Babylon (Belmont Eake State Park), Oakdale (Connetquot

State Park), Riverhead, Sarnoff Preserve, Sayville, Smithtown, Tiana (Munn’s Pond), West Islip, Yaphank, Wyandanch;

SUEEIVAN CO.: Catskill Township (Catskill Mts.); TOMPKINS CO.: Ithaca, McEean Bogs Preserve, Monkey Run Road;

UESTER CO.: New Paltz, Oliverea, Wittenberg Mountain. OHIO: COEUMBIANA CO.: Beaver Creek State Park.

PENNSYLVANIA: AEEEGHENY CO.: Pittsburgh; ARMSTRONG CO.: Elderton; BEAVER CO.: Eoeality not given;

BRADFORD CO.:Barclay; CARBON CO.: Split Rock; CENTRE CO.: State College; CEARION CO.: Clarion;

CEEARFIEED CO.: Shawville; COEUMBIA CO.: Bloomsburg; CRAWFORD CO.: Eittle Cooley; DAUPHIN CO.: Rt. 325

(3 miles SW of Orwin); EEK CO.: Allegheny National Forest (Buzzard Swamp); FOREST CO.: Allegheny National Forest

(Buzzard Swamp); INDIANA CO.: Clarksburg; EACKAWANNA CO.: Choke Creek (near Thornhurst); EUZERNE CO.:

Mountain Springs Road, Ricketts Glen State Park, Stoddartsville; MONROE CO.: Pocono Eake, Poeono Pines, State Game

Eands #127 ; MONTOUR CO.: Danville; PIKE CO.: Sugar Creek near Carlton Hill; SCHUYEKIEE CO.: near Tamaqua (Owl

Creek); SOMERSET CO.: Seven Springs; SUEEIVAN CO.: Eopez, near Ricketts Glen State Park (Rt. 487);

SUSQUEHANNA CO.: Montrose; WAYNE CO.: Gouldsboro; WESTMOREEAND CO.: Jones Mills, Powdermill Nature

Reserve, Bushy Run Battlefield (5 mi. NE of Irwin), Greensburg; WYOMING CO.: Mountain Springs Road. RHODE

ISLAND: BRISTOE CO.: Barrington, Bristol, Warren ; KENT CO.: Coventry, Greene, Warwick, West Greenwich, West

Warwick; NEWPORT CO.: Jamestown, Eittle 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 ISEE 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.: lee Mountain;

HARDY CO.: River Road east of Moorefield; PENDEETON CO.: 1.5 mile NW of Upper Tract (Briggs Run Forest Road);

POCAHONTAS CO.: Cass; RANDOEPH CO.: Spruce Knob; TUCKER CO.: Blaekwater Falls State Park.

A SHORT DISCUSSION OF CELASTRINA LUCIA

Presently, the name lucia eomprises multiple meanings:

(1) William Kirby (1837) named Polyommatus lucia from material eolleeted at Cumberland House,

Saskatehewan, 54° 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 (Erieaeeae) 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 eoarse brown markings. Following

its diseovery, lucia Kirby had been treated for an extended period as a subaretie subspeeies of a single

eontinent-wide speeies {ladon). Reeent reappraisals by Wright (1998), Nielsen (1999), Oehlensehlager &

Huber (2002), and Opler & Warren (2002) have returned C. lucia (Kirby) to a full speeies. The preeise

relationships between lucia populations at Cumberland House (Saskatehewan) and those in western North

Ameriea, Yukon-Alaska, and eastern North Ameriea have not been worked out.

(2) In eastern North Ameriea, lucia was loosely eonsidered to be the northern population of the first

spring brood of the nominate eastern subspeeies {ladon). It was pointed out by Klots (1952) and Forbes

(I960) this was not lucia Kirby, rather a highly polymorphie larger entity, solid blue above and quite

variable below (Fig 24). Pratt et al. (1994) designated it ssp. lucia Auetorum (“of authors”) and extended

its range into the eentral Appalaehians. Lucia Auet. differs signifieantly from ladon in seale morphology

and eeology (Wright & Pavulaan, 1999). It is a separate speeies, but its relationship with lucia Kirby and

other lucia populations needs further study. The newly-deseribed speeies {serotina) is elosely linked with

eastern lucia Auetorum. We propose there may be a ehain or great are of alloehronie late spring siblings

from southern Appalaehians, north through northeastern U.S. & eastern Canada, west through the Great

Lakes to the Manitoba & Saskatehewan, and south to the Foot Hills of the Roekies in Colorado. In the

eentral eontinent, argentata Fleteher and humulus Seott & Wright should be examined in this light.

(3) Males from southern New England possess the unique seale of ladon males, but in all other respeets

they are lucia Auet. This is believed to be an example of single eharaeter introgression during a warmer

postglaeial period (Hypisthermal) when southern deeiduous forests invaded southern New England

(Wright, 1971). Beeause of the introgressed trait, wings of males from this region (Fig. 4) eontrast with

males elsewhere (Fig. 24). By good fortune, this eireumstanee allowed us to separate lucia Auet. and

serotina in southern New England not only by flight period but also by a preeise morphologieal analysis.

We found no introgression of the unique seale trait into serotina, boosting our eonfidenee in its status.

(4) The name lucia has also been adopted as a form name (f 'ducid') that appears in many Celastrina

taxa. This form is eharaeterized by eoaleseed maeules in the middle of the VHW resulting in a blaek

diseal pateh. The form eommonly oeeurs in early spring taxa like lucia and ladon, but also oeeurs in

light-ventered alloehronie taxa like idella and serotina (Fig. 25). By itself, the presenee of a dark VHW

pateh is not a diagnostie trait of any speeifie taxon.

ACKNOWLEDGMENTS

■ ■ ■ -. - - - - Calhoun, Brian Cassie, K - - - - - - ■ -

)ouglas C. Ferguson, Ed Force, John Fi

:t, Richard E. Gray, Paul Grey, Alex G

Hanks^Dave Hoag, Jack Holliday, David ffiner, Bernar^ Jackson^ Marc Johnson, Phil Kean,^D^n Lafontmne^I^ss Layberry,

Springer, Melanie Statts, Clare Stone, Clay Taylor, Anthony Thomas, David Wagner,’ Mark Walker, St^e Walter, Reed

Field Museum (Phil Parillo), Florida St

n of Natural History-

•k O’Brien), University of Peni

(MBAP): Kathleen S. Ar

Tom Carpenter, Brian Cc

Ann S. Earley, Will Ear

^dlow,WmiamT^Map

David Young; and the members of the C

k the members of the IV

r, R A. Foster, T. H

i, J. Fengler, Ed Force, Eawrence Gall, B. Godwin, P. Godwin, Greg Hanisek, Ronald Harms, John Himmelman,

ge, Annie Paradis, Noble Proctor, Marion Richardson, Ed Sluzenski, David Stevens,’ Clay

s, C. Tondreau, Monty Volovski, and David Wagner. Special thanks are extended to Chris

LITERATURE CITED

Allen, TJ. 1997. The butterflies of West Virginia and their eaterpillars. Univ. of Pittsburgh Press,

Pittsburgh, Pennsylvania, 388 pp.

Allen, T.J., J.P. Broek & J. Glassberg. 2005. Caterpillars in the field and garden. A field guide to the

eaterpillars of North Ameriea. Oxford Univ. Press, New York, NY, 232 pp.

Boisduval, J.B.A. & J.E. Ee Conte. [1833]. Historie generale et ieonographie des lepidopttes et des

ehenilles de f Amerique septentrionale. Roret, Paris, 228 pp., 78 pi.

Ceeh, R. & G. Tudor. 2005. Butterflies of the East Coast: An observer’s guide. Prineeton Univ. Press,

Prineeton, NJ, 345 pp.

Chew, F.S. & R.K. Robbins. 1984. Egg-laying in butterflies (Chapter 6). In The Biology of Butterflies.

Vane-Wright & Aekery (eds.), Aeademie Press, Eondon, 429 pp.

Cleneh, H.K. 1958. The butterflies of Powdermill Nature Reserve. Researeh Report no.l, Powdermill

Nature Reserve. A Researeh Station of Carnegie Museum, Deeember 30, 11 pp.

Comstoek, J.H. and A.B. Comstoek. 1904. How to know the butterflies. A manual of the butterflies of the

eastern United States. D. Appleton & Co., New York, NY, 311 pp.

Comstoek, W.P. 1940. Butterflies of New Jersey. J. New York Ent. Soe. 48:47-84.

Davis, W.T. 1910. Eist of the Maerolepidoptera of Staten Island, New York. Staten Island Assoe. Arts

Sei. Proe. 3:1-30.

Dirig, R. 2002. Definitive Destination MeEean Bogs Preserve: Finger Fakes Region, New York.

Ameriean Butterflies 10(4) : 4-16.

Doubleday, E. 1841. Deseription of a new North Ameriean Polyommatus. The Entomologist 1:209-211.

Edwards, W.H. 1862. Deseriptions of eertain speeies of diurnal Eepidoptera found within the limits of the

United States and British Ameriea. -No.2. Proe. Aead. Nat. Sei., Phila. 14:54-58.

Edwards, W.H. 1875. Some notes on Lycaena Pseudargiolus. Canad. Ent. 7(5): 81-83.

Edwards, W.H. 1883. On the polymorphism of Lycaena Pseudargiolus. Bois. Papilio 3: 85-97.

Edwards, W.H. 1884. Butterflies of North Ameriea. Vol. 2. Houghton, Mifflin & Co., Boston, MA, 357

pp., 51pl.

Engel, H. 1908. A preliminary list of the Eepidoptera of western Pennsylvania eolleeted in the vieinity of

Pittsburgh. Ann. Carnegie Mus. 5(1): 27-136.

Ferguson, D.C. 1954. The Eepidoptera of Nova Seotia. Proe. Nova Seot. Inst. Sei. 23(3): 161-375.

Ferguson, D.C. 1975. Host reeords for Eepidoptera reared in eastern North Ameriea. USDATeeh. Bull.

1521:1-49.

Forbes, W.T.M. 1960. Eepidoptera of New York and neighboring states. Part IV.Agaristidae through

Nymphalidae ineluding butterflies. Memoir 371. Cornell Univ. Agrie. Exp. Sta. NY State College

of Agrie., Ithaea, NY, 188 pp.

Glassberg, J. 1993. Butterflies through binoeulars: A field guide to butterflies in. the Boston-New York-

Washington region. Oxford University Press, New York, NY, 160 pp.

Glassberg, J. 1999. Butterflies through binoeulars: The East. Oxford University Press, New York, NY,

242 pp.

Goehfeld, M. & J. Burger. 1997. Butterflies of New Jersey: A guide to their status, distribution,

eonservation, and appreeiation. Rutgers University Press, New Brunswiek, NJ, 327 pp.

Handfield, E. 1999. Ee guide des Papillion du Quebee. Broquet, Broueherville, Quebee, 960 pp.

Harris, T.W. 1862. A treatise on some of the inseets injurious to vegetation. Revised edition. (C.E. Flint,

ed.) William White, Printer to the State, Boston, MA, 640 pp.

Iftner, D.C., J.A. Shuey, and J.V. Calhoun. 1992. Butterflies and skippers of Ohio. Ohio Biol. Surv. Bull.

New Series, vol. 9, no. 1, xii + 212 pp. (inel. 40 pis.).

Kirby, W. 1837. Part 4. The inseets. pp. xxxix + 325, pi. I-VIII. In J. Riehardson: Fauna-boreali-

Amerieana; or the zoology of the northern parts of British Ameriea: eontaining deseriptions of the

objects of natural history collected on the late Northern Land expeditions, under command of

Captain Sir John Franklin, R.N. Longman, London.

Klots, A.B. 1951. A field guide to the butterflies of North America east of the Great Plains. Houghton

Mifflin Co., Boston, MA, 349 pp.

Layberry, R.A. 1996. The Spring Azure species complex. In Butterflies of Ontario & Summaries of

Lepidoptera Encountered in Ontario in 1995. Toronto Ent. Assoc. Occ. Pub. #28-96, Section 4.5,

p. 12-15.

Eayberry, R.A., P.W. Hall & J.D. Eafontaine. 1998. The butterflies of Canada. Univ. of Toronto Press,

Toronto, Ontario, 280 pp.

Eeblanc, A. 1985. Ees Eycenides (Eepidoptera: Eycaenidae) du Quebec. Fabreries. Supplement 4, 66 pp.

Eintner, J.A. 1875. On Lycaena Neglecta, Edw. Canad. Ent. 7:122-123.

Nielsen, M.C. 1999. Michigan butterflies and skippers. Michigan State University Extension, East

Eansing, 248 pp.

Ochenschlager, R. & R. Huber. 2002. The butterflies of Wadena County, Minnesota. Scientific

Publications of the Science Museum of Minnesota, New Series 8(1): iv, 1-26.

Opler, P.A. & G.O. Krizek. 1984. Butterflies east of the Great Plains. The Johns Hopkins University

Press, Baltimore, MD, 294 pp.

Opler, P.A. & V. Malikul. 1992. A field guide to eastern butterflies. Peterson field guide series, no.4.

Houghton Mifflin Co., Boston, MA, 396 pp.

Opler, P.A. & A.D. Warren. 2002. Butterflies of North America. 2. Scientific Names Eist for Butterfly

Species of North America, north of Mexico. Contributions of the G.P. Gillette Museum of

Arhtropod Diversity, Colorado State University, 79 pp.

Packard, A.S. 1869. Guide to the study of insects and a treatise on those injurious and beneficial to crops

for the use of colleges, farm schools, and agriculturists. Salem, Naturalist’s Book Agency, 702 pp.

Parker, H.W. 1874. Novelties in Amherst, Mass. Psyche 1(7); 26.

Pavulaan, H. 1985. Field survey of the true butterflies (Papilionoidea) of Rhode Island. J. Eepid. Soc.

39(1): 19-25.

Pavulaan, H. 1989. The butterflies of West Warwick, Rhode Island, and vicinity: A study of urban

butterflies. Y.E.S. Quarterly 6(2):4-31 [April/June].

Pavulaan, H. 1993. Butterflies at Rhode Island's Great Swamp. Y.E.S. Quarterly 10(2): 1-32 [April/June]

& 10(3): 1-32 [July/Sept].

Pavulaan, H. 2002. Azure butterflies redefined. The Connecticut Butterfly Association Newsletter. No. 9,

March, p. 7.

Pavulaan, H. & D.M. Wright. 1994. Out of the azure and into the lab: The current state of Celastrina

research. The Mulberry Wing 10(1):406.

Pavulaan, H. & D.M. Wright. 2000. The biology, life history, and taxonomy of Celastrina neglectamajor

(Eycaenidae: Polyommatinae). The Taxonomic Report of The Inti. Eep. Survey 2(5): 1-19.

Pelikan, M.E. 1998. Island blues: 1998 azure flights on Martha’s Vineyard. Massachusetts Butterflies, no.

11, August, p. 4-8.

Pratt, G.F., D.M. Wright, and H. Pavulaan. 1994. The various taxa and hosts of the North American

Celastrina (Eepidoptera: Eycaenidae). Proc. Entomol. Soc. Wash. 96(3):566-578.

Remington, C.E. 1956[58]. Genetics of populations of Eepidoptera. Proc. Tenth Infl Congr. Ent. 2:787-

805.

Saunders, W. 1875. [Note. Ed. C.E. {Canadian Entomologist}]. Canad. Ent. 7(5): 82-83.

Scott, J.A. & D.M. Wright. 1998. A new Celastrina from the eastern slope of Colorado. Papilio n.s., no.

9, 15 pp.

Scudder, S.H. 1869. Entomological correspondence of Thaddeus William Harris, M.D. Occ. Papers

Boston Soc. Nat. History, no.l, 375 pp.

Scudder, S.H. 1876. The relationship of the early spring blues. Canad. Ent. 8(4): 61-66.

Scudder, S.H. 1881. Butterflies: their strueture, ehanges, and life-histories. Henry Holt and Co., New

York, NY, 322 pp.

Seudder, S.H. 1889. Butterflies of eastern United States and Canada. 3 vols.. Published by author,

Cambridge, Massaehusetts.

Seudder, S.H. 1893. Brief guide to the eommoner butterflies of the northern United States and Canada;

being an introduetion to a knowledge of their life-histories. Henry Holt and Co., New York, NY,

206 pp.

Shapiro, A.M. 1966. Butterflies of the Delaware Valley. Speeial Publieation of the Ameriean

Entomologieal Soeiety, Philadelphia, PA, 79 pp.

Shapiro, A.M. 1974. Butterflies and skippers of New York State. Seareh 4(3): 1-59.

Smith, J.B. 1910. The inseets of New Jersey. Annual Report of the New Jersey State Museum 1909.

Trenton, NJ, 888 pp.

Thomas, A.W. 1996. A preliminary atlas of the butterflies of New Brunswiek. New Brunswiek Museum

Publieations in Natural Seienee No. 11, 41 pp.

Wagner, W.H., Jr. & T.L. Melliehamp. 1978. Foodplant, habitat, and range of Celastrina ehenina

(Lyeaenidae). J. Tepid. Soe. 32(l):20-36.

Wagner, D.L. 2005. Caterpillars of eastern North Ameriea: A guide to identifieation and natural history.

Prineeton Univ. Press, Prineeton, NJ, 512 pp.

Walter, S. 1993. Out of the Blue (?) and into the Azure. The Mulberry Wing 9(l):5-6, April-May.

Wright, D.M. 1995. The Ameriean Azures: Our blue heaven. Ameriean Butterflies 3(l):20-28 & 30.

Wright, D.M. 1998. Update on Celastrina ineluding notes on Ohio speeies. The Ohio Lepidopterist 20(2):

18-21.

Wright, D.M. & H. Pavulaan. 1999. Celastrina idella (Lyeaenidae: Polyommatinae): a new butterfly

speeies from the Atlantie eoastal plain. The Taxonomie Report of The Inti. Lep. Survey 1(9): 1-

11.

Wright, H.E, 1971. Late quaternary vegetational history of North Ameriea. (pp. 425-464) In K.K.

Turekian (Ed.) The Late Cenozoie Glaeial Ages. Yale University Press, New Haven.

Date of publication: 1 December 2005

The Taxonomic Report

is a publication of

The International Lepidoptera Survey (TILS).

(A Tax Exempt Non-Profit Scientific Organization)

TTR is published as a public and scientific work for purposes of taxonomic study and nomenclatural acts.

It appears on printed paper in sequential volumes and issues, is regularly disseminated to institutional and individual

subscribers, and is also available as separate issues free of charge upon request at the discretion of authors and/or editor.

TILS Purpose. TILS is devoted to the worldwide eolleetion of Lepidoptera for the purpose of seientifie

diseovery, determination, and doeumentation, without whieh there ean be no preservation of Lepidoptera.

TILS Motto. As a world eommunity, we eannot proteet that whieh we do not know.

Articles for publication are sought. They may deal with any area of taxonomie researeh on Lepidoptera.

Before sending a manuseript, simply write to TILS, P.O. Box 1124, Herndon, VA 20172 to set up

diseussion on how to best handle your researeh for publieation.

Donations are needed

to support and further our efforts

to discover and protect butterflies worldwide.

All donations are US tax deductible. Please help generously.

Donations should be mailed to: TILS, c/o Harry Pavulaan, P.O. Box 1124, Herndon, VA 20172.

Cheeks should be made payable to: TILS. Please indieate if you need an individual reeeipt.

Visit The International Lepidoptera Survey on the World Wide Weh at:

http://www.tils-ttr.org

Join the discussion at our list serve on Yahoo!Groups at:

TILS-Leps-Talk

Volume 6 Number 7

15 March 2007

The Taxonomic Report

OF THE INTERNATIONAL LEPIDOPTERA SURVEY

A new subspecies of Colias gigantea

from arctic Alaska (Pieridae)

Jack L. Harry

47 San Rafael Court, West Jordan, Utah, United States of America 84088

Abstract: A new subspecies of Colias gigantea Strecker from the ‘north slope’ of Alaska is described.

Additional key words: inupiat, philodice

INTRODUCTION

Field work in northern Alaska, United States of Ameriea has revealed that northern populations of Colias gigantea

along the Dalton Highway are suffieiently distinet from the more southerly populations to merit reeognition as a

named subspeeies.

Colias gigantea inupiat Harry, new subspeeies

Description

Males: Forewing length is 20 to 25 mm. Dorsal surfaees: Blaek border is medium to wide, blaek spot at end of

forewing eell is redueed to absent. The hind wing diseal eell spot is white to light orange. Basal dark oversealing is

more extensive than C. g. gigantea. Ventral surfaees: The inner marginal area of forewing is yellow, oeeasionally

beeoming slightly lighter yellow. Hind wing with more greenish over-sealing and not as yellow as interior Alaska C.

gigantea. Diseal spot is red or white with red ring. Satellite spot is present or absent. No submarginal brown spots

on hind wing.

Females: Forewing length 23 to 26.5 mm. Dorsal surfaees: Ground eolor is ereamy yellow to yellow. There is no

border to a slight border, rarely a somewhat extensive border. Hind wing diseal spot is orange. Basal area

oversealing is more extensive than interior Alaska C. gigantea. Ventral surfaees are as in males.

Type Specimens

Holotype male: Alaska, Mile 323 Dalton Hwy, 68°59.0rN 148°49.94'W, 365 meters elevation, 28 June 2003 (Plate

1). Paratype (allotype) female: same loeation as holotype, 12 July 2006 (Plate 1). Paratypes: Mile 323 Dalton Hwy,

(1M2F) 28 June and 2 July 1999; (2M) 2-3 July 2001; (IM) 28 June 2002; (18M6F) 28 June-3 July 2003; (5M3F) 28

June-1 August 2005; (IM) 3 July 2006. Mile 353 and Mile 356 Dalton Hwy, Sagwon Hills, 305 meters elevation,

(IM) 4 July 1999; (7M1F) 3-9 July 2006. Mile 381 Dalton Hwy, 34 miles south of Deadhorse, 90 meters elevation,

(IMIF) 10 July 1999; (3M) 24 June-15 July 2005; (2M) 9 July 2006.

The holotype, allotype, and 5 males will be deposited in the Florida State Colleetion of Arthropods, Gainesville,

Florida, USA. Ten males and 5 females will be retained in the eolleetion of the author. Ten males and 4 females are

in the eolleetion of Robert Worthy, Caterham, Sussex, England. Two males and 1 female are in the Sagamihara City

Brigham Young University, Provo, Utah, USA. Four males and 2 females will be deposited in the Smithsonian

Institute, Washington, D.C. USA. Four males and 2 females will be deposited in the California Aeademy of

Plate 1. Holotype male and allotype female ofColias gigantea inupiat Harry, 2007

with eomparison images of C. philodice. Seale is life size. J. Harry photographs

This subspeeies is given the name inupiat in honor of the native

(Langdon, 2002). The suggested eommon name is Inupiat Sulphur.

climate here is more like the interior than the north flope.^being warmer in the summer and with a

II fiitfi

llJlii 11

spots, some of th

forewmg area at the inner m

Langdon, S. J. 2002. The Native People

The Taxonomic Report

is a publication of

The International Lepidoptera Survey (TILS)

(a tax exempt non-profit scientific organization)

The Taxonomic Report is published for the purpose of providing a public and permanent scientific record.

It appears on printed paper in sequential issues, is regularly disseminated to institutional and individual

subscribers, and is also available as separate issues free of charge upon request at the discretion of authors

and/or the editor. Contents are peer-reviewed but not necessarily through the anonymous review and

comment process preferred by some publishers of serial literature.

TILS Purpose

TILS is devoted to the worldwide collection of Lepidoptera for the purpose of scientific discovery,

determination, and documentation, without which there can be no preservation.

TILS Motto

“As a world community, we cannot protect that which we do not know”

Articles for publication are sought

They may deal with any area of research on Lepidoptera, including faunal surveys, conservation topics,

methods, etc. Taxonomic papers are especially welcome. Before sending a manuscript, simply write to

TILS editor, Harry Pavulaan, P.O. Box 1124, Herndon, VA 20172 to set up discussion on how to best

handle your material for publication; or email harrypav@hotmail.com

Donations are needed

to support and further our efforts

to discover and protect butterflies worldwide.

All donations are US tax deductible. Please help generously.

Donations should be mailed to: TILS, c/o Harry Pavulaan, P.O. Box 1124, Herndon, VA 20172.

Checks should be made payable to: TILS. Please indicate if you need an individual receipt.

Visit The International Lepidoptera Survey on the World Wide Weh at:

httD://www.tils-ttr.org

Join the discussion at our list serve on YahooIGroups at:

TILS-Leps-Talk

http: //groups. v ahoo, com/group/TIL S -leps -talk/

Volume 6 / Number 8

10 April 2007

The Taxonomic Report

OF THEINTEBNATIONAL LEPIDOPTERA SURVEY

BIONOMIC STUDIES OF PAPILIO DEMOTE US LINNAEUS, THE CITRUS BUTTERFLY,

(LEPIDOPTERA: PAPILIONIDAE), FROM LOWER SINDH, PAKISTAN.

ARSHAD MUNIR\ NIKHAT YASMIN SIDDIQUI^ MUHAMMAD A. RAFI^ HARRY PAVULAAN^ & DAVID WRIGHT^

different varieties of Citrus sp. (Citrus) plants. The length of the adult mating period ranged 5 to 8 hours. Following mating,

females laid 27.7 ± 3.3 eggs. Eggs hatched in 1.5 to 7 days. Mean durations of larval instars were 1.8 ± 0.1 (minimum) and 2.9

significant role in mortality ^in all stages of the butterfly.^Survival in most life stages had a negative correlation^ith temper^ature.

le feeding effect of Curry Leaf and te

Wgzf (Cur^ Thee

he Growth Index value was highest on Citrus aurantiifolia (Kaghzi Lime) and lowest on

INTRODUCTION

Papilio demoleus Linnaeus, the Citrus Butterfly (also eommonly known as the Lime Swallowtail), is a

major pest of Citrus sp. (Citrus) and Murray a koenigii (Curry Leaf) plantations (Duport, 1913; Malik,

1970, Badawi, 1981). The butterfly has a wide eeologieal toleranee, enabling it to thrive in a broad variety

of elimatie eonditions (Raft et al, 1999d). It is found in mueh of southern Asia from Saudi Arabia east to

Australia, New Guinea and the Philippines. More reeently, it has been introdueed into the Caribbean

(Eastwood et al, 2006; Guerrero et aL, 2004; Homziat and Homziak, 2006; Wehling et al, 2007). Where

it oeeurs, it is eosmopolitan in distribution (Srivastava and Ahlawat, 1999). In Pakistan, P. demoleus flies

in all four provinees virtually year-round with greatest numbers observed from August through Oetober

(Roberts, 2001).

Citrus is a highly profitable fruit erop that grows between

125 eountries with an estimated annual produetion of 93.7

Pakistan, it is grown over an area of 197,021 heetares with

a major export, it holds a pivol

Pakistan, 1995). Curry Leaf is

in eooking (Wasif, 1991). Be

losses to these plant erops ever

important agrieultural enterpris

i 40° S latitude worldwide in more

3ns (Srivastava and Ahlawat, 1999

l1 produetion of 1.9 million tons. I

’s agrieultural industry (Governme

leaves are traditionally used as a

y deeline or even eollapse in the m

udy presents essential aspeets of P. demoleus bionomies to better understand its funetion as an

;ural pest. The study ineludes biologieal parameters sueh as mating behavior, oviposition, feeundity

igth of life stages, number and duration of generations, and the effeet of biotie and abiotie faetors.

itus is ealibrated by a Growth Index on Curry Leaf and ten different varieties of Citrus plants.

,i.^Natl.Ag.Res.C

d. “tils St

Natural enemies and survival rates at different life stages are determined. The effeets of artifieial diets on

adults are noted.

MATERIALS AND METHODS

Bionomie studies of Papilio demoleus Linnaeus were eondueted at Tando Muhammad Khan, lower Sindh

(Pakistan). Eggs and larvae were first eolleeted for this study in Citrus and Curry Leaf plantations in the

vieinity of Karaehi in 1998. Larvae were reared on Citrus sp. (Citrus) and Murraya koenigii (Curry Leaf)

in plastie jars provided with fresh leaves of hostplants twiee a day until pupation. Ten pairs of the resultant

adult butterflies were released in January 1999 into a wood-framed eage measuring 10x15x10 feet in

size. The eage was eovered on four sides with #15 nylon mesh size, whieh allowed unimpeded air

eireulation. A single small entranee door was plaeed on one side. During the first (three-year) phase of the

study (1999-2001), Citrus aurantiifolia var. ‘Kaghzi Lime’ and Murraya koenigii were planted inside the

eage to serve as ovipositional substrates and larval hostplants. Plant height was eonstantly maintained at

three feet. Seasonal flowering plants were provided as neetar sourees for adult butterflies. In addition, a

supplemental 15% honey solution was sprayed at regular intervals on flowering plants inside the eage.

All eages were thoroughly examined eaeh day to reeord observations and to eheek for immatures. Data

were eolleeted regarding survival rate, pre-mating behavior, mating period, pre-oviposition, oviposition

period, feeundity rate and ineubation period. All the observations were taken eontinuously during 13

daytime hours daily from 6:00 A.M. to 7:00 P.M. Ambient temperature was reeorded frequently. Parasites

and predators of all infested stages were preserved and identified at the Entomology Seetion of the

Department of Zoology, University of Karaehi.

The next generation was maintained in similar eonditions. This study group was initiated using ten random

pairs of adult butterflies bred from eggs of the first generation and released into a seeond similar eage. This

proeess was repeated for thirteen generations over the eourse of eaeh year, alternating the use of two eages.

Observations were eontinued through the duration of the three-year period 1999-2001. Survival data of 39

generations of P. demoleus were analyzed for eaeh life history stage (egg, larva, pupa and adult), ineluding

ealeulation of mean and standard deviation (S.D.).

The seeond phase of this study was eondueted in 2002. Ten pairs of adult butterflies (originating from the

last generation of the previous year) were released in January into the wood-framed eage, with subsequent

generations alternating between two eages. During this phase of the study, Murraya koenigii and ten Citrus

varieties were planted inside the eages to serve as ovipositional substrates and larval hostplants. The Citrus

varieties used were: Citrus aurantiifolia (Christmann) Swingle var. ‘Kaghzi Lime’ (Kaghzi Lime), Citrus

aurantiifolia var. ‘Sweet Lime’ (Sweet Lime), Citrus aurantium Linnaeus var. ‘Khatta’ (Khatta Orange,

Sour Orange), Citrus limon Linnaeus var. ‘Eureka’ (Eureka Lemon), Citrus limonia Osbeek (Common

Jambhiri), Citrus medica Linnaeus (Citron), Citrus reticulata Blaneo var. ‘Coorg’ (Sangtra Coorg

Mandarin), Citrus reticulata Blaneo var. ‘Kinnow’ (Kinnow Sangtra), Citrus reticulata Blaneo var.

salicifolia (Citrus Willow, Willow-leaved Mandarin), and Citrus sinensis Linnaeus var. ‘Washington

Navel’ (Washington Navel Orange). Potted plants, five of eaeh variety, were plaeed LA ft. apart within the

eage. Plant height was eonstantly maintained at three ft. Seasonal flowering plants were provided as neetar

sourees for adult butterflies. In addition, a supplemental 15% honey solution was again sprayed at regular

intervals on flowering plants inside the eage.

After release of ten pairs of adult butterflies, eaeh plant was elosely observed to reeord ovipositional

preferenees. The number of eggs deposited on eaeh variety of plant was reeorded. Ten first instars of eaeh

generation were transferred to separate plastie jars and provided with respeetive host plant leaves twiee a

day. Larval growth rates and survivorship were noted. This protoeol was repeated for eaeh of thirteen

generations over the eourse of the year. The Growth Index (N x 1/T) was ealeulated for developmental

stages of P. demoleus on the seleeted hostplants. [N = % larvae reaehing maturity; T = mean larval period]

The third phase of this study was eondueted in 2003. Ten pairs of adult butterflies (originating from the

last generation of the previous year) were released in January into eaeh of three separate wood-framed

eages, with subsequent generations alternating between two sets of eages. During this phase of the study,

Murraya koenigii and ten varieties of Citrus plants were similarly utilized inside the eages to serve as

ovipositional substrates and larval hostplants. Three adult diets eonsisting of 15% solutions of honey,

suerose and glueose respeetively were tested in eaeh eage. These solutions were sprayed at regular

intervals on flowering plants in the eage. The effeet of eaeh diet on pre-oviposition period, feeundity rate

and longevity was noted. Larval sueeess of this generation was noted on Murraya koenigii and the ten

Citrus varieties.

RESULTS

Immature Stages

Egg (Figs. 1 & 2). Egg round with a smooth and glistening surfaee. Color yellowish-green. Part of the egg

glued to the plant is flat. Slowly turns brown before hatehing (Fig. 2). Fertile eggs with prominent

reddish-brown mieropyle. Females lay eggs on young leaves, petioles, shoots, thorns and flowers of Citrus;

most females prefer to deposit eggs on both sides of the leaves. Mean egg diameter 1.1 mm. Larva. First

instar (Figs. 3 & 4). Neonate larva emerges from egg by small lateral hole. Fully emerged larva usually

eats the empty eggshell. Body eolor light brown; head brown to yellowish-white. Prothorax light

yellowish-brown. Yellowish to milky white pateh eovers 3’^^-5^^ abdominal segments. Abdominal 8^^-9^^

segments yellowish-brown. Remaining segments with darker rings speekled with blaek patehes. Head

narrower than the thorax; remainder of body tapers. Larval eutiele with dark brown spines, whieh arise

direetly from the body on either side of a mid dorsal line. Length 3.5 mm. Width 0.7 mm (mesothorax) and

0.4 mm (9^^-10^^ abdominal segments). A faint bird-dropping pattern appears in this instar; the eamouflage

is mueh more developed in the seeond through fourth instar. Second instar (Fig. 5). Body eolor dark

brown. Head brown. Prothorax yellowish-brown. Prominent dorsal yellowish to milky white pateh

(“saddle”) on the 3"^-4^^ abdominal segment; extends anterolaterally to 2”^ abdominal segment. Abdominal

8*-9^^ segments brownish yellowish. Spines as in the first instar. Length 7.5 mm. Width 1.8 mm

(mesothorax) and 0.8 mm (9^^-10* abdominal segments). Third instar (Fig. 6). Body eolor dark brown.

Head brown. Prothorax yellowish brown with some milky white markings on the lateral side. Prominent

dorsal white marking on 3’^^-5* abdominal segment; runs obliquely and laterally on either side of the 2”^

abdominal segment and extends to 6* abdominal segment. Pair of small lateral white patehes on eaeh side

of 7*-8* abdominal segments. Prolegs yellowish brown. Nine pairs of spiraeles on lateral side of abdominal

segments; one on lateral prothorax. Number, strueture, eolor and position of spines same as seeond instar.

Length 13.5 mm. Width 2.4 mm (mesothorax) and 1.5 mm (9*-10* abdominal segments). Fourth instar

(Fig. 7). Similar to previous instar exeept size. Body eolor dark brown. Head brown. Prothorax yellowish

brown with lateral milky white markings extending to abdominal segment. Two broad blaek patehes

laterally on either side of 2”^ abdominal segment and small white markings laterally at the base of the

prolegs on 3*-6* abdominal segments. White lateral patehes on the 7*-8* abdominal segments. Number,

strueture, eolor and position of spines same as third instar. Length 19.0 mm. Width 4.1 mm (mesothorax)

and 2.2 mm (9*-10* abdominal segments). Fifth instar (Fig. 8). Body eolor green to yellowish green.

Milky white venter and lateral line. Head light brown. Prothorax green with small brown tubereles. Two

dorsal blaek bands between meso-metathorax and abdominal segments. Small eyespot on

dorsolateral metathorax. Oblique lateral brown patehes on 4* and 5* segments. Minute blaek spots on

anterior parts of the abdominal segments, sometimes absent. Last abdominal segment with two

terminal brown spines. Body smooth without any branehing spines. Length 39.5 mm. Width 9.4 mm

(mesothorax) and 5.50 mm (9^^-10^^ abdominal segments). Note: After eaeh molt, larvae eommonly eat

exuviae exeept for hard selerotized parts. Larvae rest approximately 30-45 minutes as new eutiele hardens.

Pupa (Figs. 9-11). Pupation oeeurs 2-5 meters from the host plant. Sites vary and inelude netting, pipes,

wood struetures, potted plants and pots. In prepupal phase, larva forms silk pad on substrate to whieh anal

prolegs are anehored. A silk girdle is formed, passing over the metathoraeie region and holding the

prepupa in position (Fig. 9). The prepupal period lasts 1-2 days. At the end of this period larval skin is

shed. Pupal skin takes approximately 40-70 minutes to harden. Pupal eolor yellowish green (Fig. 10) or

light brown (Fig. 11). Head region with two frontal projeetions. Small dorsal tubereles on eaeh abdominal

segment. Pupae beeome transparent as eelosion nears. Length 30.2 mm. Width 9.8 mm.

Adult

Body eolor blaek dorsally, white ventrally with blaek stripes. Wings blaek, heavily patterned with irregular

white marks. Hindwings laek tails. Red spot at anal angle, eapped with a blue-blaek spot and light blue

irideseent ereseent. Blue-blaek spot and ereseent often absent in males (Fig. 12), but prominent in females

(Fig. 13). Wing eolor beeomes a dull brown with age. Mean wing expanse (spread forewings apex to apex)

of males 80.1 mm (n=10) and females 93.8 mm (n=10).

Duration of Life Stages

Thirteen generations were reeorded as overlapping broods during the eourse of a year. The duration of the

different generations was sensitive to ambient air temperature and varied from 22.5 to 46.5 days depending

upon season. Cool-season generations were the longest-lived. Mean temperatures ranged from 17.5 to

38.0°C during different seasons with eoolest temperatures in Deeember-January and highest in Mareh-July.

Duration of immature life stages varied as follows: egg (1.5-7.0 days), larva (8.0-14.5 days), pupa (5.5-

20.0 days). Mean durations of larval instars were 1.8 ± 0.1 (minimum) and 2.9 ±0.1 (maximum) days

respeetively. Adult longevity averaged 4.9 days for the males and 6.7 days for the females over thirteen

generations.

Mating and Oviposition

The pre-mating period began immediately after emergenee, lasting 4-10 hours at an air temperature range

of 15.9-37.0°C. Mating period lasted 5-8 hours. Females were observed to mate only onee during their

adult lifespan. After mating, females typieally delayed oviposition for 23-40 hours. Deposition of all eggs

was eompleted within 33-54 hours. Oviposition oeeurred between 8:00 A.M. to 6:00 P.M. The number of

eggs deposited per female ranged from 13-40 with a mean of 27.7 ±3.3.

Survival and Mortality of Immature Stages

Mortality rates for immature stages of all 13 generations were as follows: eggs (22.7 ± 4.4%), instar

(22.9 ± 8.3%), 2"“ instar (15.7 ± 5.9%), 3''“ instar (14.0 ± 5.5%), 4* instar (20.4 ± 5.4 %), 5* instar (16.7 ±

5.1%), and pupa (27.0 ± 8.8%). The high rate of egg mortality was due to infertility, falling off leaves,

predators (ants, praying mantis) and the parasitoid Trichogramma chilonis (Isehii) [Hymenoptera:

Triehogrammatidae]. The larval mortality rate was primarily due to falling off leaves and predation by ants

and the Common Garden Lizard (Calotes versicolor (Daudin) [Reptilia: Squamata: Agamidae]). Pupal

mortality was primarily due to predation by the Common Garden Lizard and the hymenopterous parasitoid

Peteromalus puparum (L.) [Hymenoptera: Pteromalidae].

Figs. 1-11. Immature stages of Papilio

demoleus from lower Sindh, Pakistan.

Fig. 1. Fresh egg.

Fig. 2. Mature egg.

Fig. 3 & 4. First instar larva.

Fig. 5. Seeond instar larva.

Fig. 6. Third instar larva.

Fig. 7. Fourth instar larva.

Fig. 8. Fifth instar larva.

Fig. 9. Pre-pupa.

Fig. 10. Pupa (green).

Fig. 11. Pupa (brown).

Host Plant Preference

The host preference of P. demoleus was documented by counting the number of eggs laid on Murraya

koenigii and on each variety of Citrus. Plants with a large number of new leaves usually had a higher mean

number of eggs laid by females (Table 1). The data presented in table demonstrates that Citrus

aurantiifolia var. 'Kaghzi Lime' recorded the highest Growth Index (G.L) value of 13.8, followed by 12.3

for both Citrus reticulata var. salicifolia and Citrus limonia. Citrus sinensis and Citrus medica recorded

the lowest G.L value of 4.00 among Citrus, while Murraya koenigii recorded the lowest G.L overall of 1.8.

On the basis of G.L value, the host preference of Papilio demoleus is as follows, in decreasing order: Citrus

aurantiifolia (Kaghzi Lime) > Citrus limonia (Common Jambhiri) > Citrus reticulata var. salicifolia

(Citrus Willow) > Citrus aurantiifolia (Sweet Lime) > Citrus aurantium (Khatta Orange) > Citrus

reticulata (Kinnow Sangtra) > Citrus reticulata (Sangtra Coorg Mandarin) > Citrus limon (Eureka Lemon)

> Citrus medica (Citron) > Citrus sinensis (Washington Navel Orange) > Murraya koenigii (Curry Leaf).

Scientific Name

Mean Number Eggs

Per Female

Total Number of |

Mean Larval Period I

Pupated 1

Number of Adults

Eclosing 1

Growth Index (G.l.) |

Citrus aurantifolia var. 'Kaghzi Lime'

Kaghzi Lime

20.6

6.5

13.84

90%

Citrus reticulata var. saiicifoiia

Citrus Willow

19.8

6.5

12.30

80%

Citrus iimonia

Common Jambhiri

17.6

6.5

12.30

80%

Citrus aurantium var. 'Khatta'

Khatta Orange

14.4

7.0

10.00

70%

Citrus aurantifolia \/ar. 'Sweet Lime'

Sweet Lime

13.4

7.0

10.00

70%

Citrus reticulata var. 'Kinnow'

Kinnow Sangtra

7.5

8.00

50%

Citrus reticuiata var. 'Coorg'

Sangtra Coorg

11.6

7.0

7.14

40%

Citrus iimon var. 'Eureka'

Eureka Lemon

11.4

7.0

7.14

40%

Citrus sinensis var. 'Washington Navel'

10.0

4.00

30%

Citrus medica

Citron

10.8

10.0

4.00

30%

Murraya koenigii

Curry Leaf

10.4

11.0

1.82

10%

Table 1. Female Ovipositional Preference and Growth Index (G.l.) value of developmental stages of P. demoleus on selected hostplants during 2002.

Natural Enemies

A field survey of natural enemies of P. demoleus eondueted at various Citrus orehards found that in general

birds were the most important predators during larval stages, espeeially 2nd instar to 5th instars. Birds also

attaeked adults. Wild eggs in the field infested by the parasitoid Trichogramma chilonis were noted.

Predator ants (blaek, brown), praying mantises, and Common Garden Lizards were eommon in the field.

Influence of Artificial Diets

Adult P. demoleus females while being fed a 15% honey solution laid a larger proportion of their eggs

(27.7%) in eontrast to females being fed 15% suerose solution (12.4%) and 15% glueose solution (13.3%).

Adult longevity was 4.5-5.0 days (males) and 5.0-7.0 days (females) while being fed a 15% honey solution.

This was empirieally better than adult longevity rates of 3.0-4.5 days (males) and 3.0-4.0 days (females) on

15% suerose solution, and of 3.0-4.0 days (males) and 3.0-4.5 days (females) on 15% glueose solution.

DISCUSSION

We found the egg ineubation period of P. demoleus was 1.5-7.0 days over the range of seasonal ambient

temperatures in Pakistan. This range spans the variable reported ranges in the literature, speeifieally 2-9

days in Mushtaque (1964), 3-6 days in Mishra and Pandey (1965), 4-5 days in Badawi (1968), 3-6 days

(Abu-Yaman, 1973), 3.8-8.9 days (Farahbakhsh and Khashkooli, 1978), 3.1-6.1 days (Badawi, 1981), 2-3

days in Winotai and Napompeth (1981), 3-6 days in Radke and Kandalkar (1989), 4-5 days in Rafi et al,

(1989), 2.4-3.5 days in Rafi et al (1999e), 1.5-4.5 days in Rafi et al (1999d), and 3.2-4 days in Atwal

(1976).

We have determined that there are five larval instars, similar to that reported by Ghosh (1914) and

Mushtaque (1964), but differing slightly from Badawi (1981) who reported 5-6 larval instars. In our study

the duration of the larval stage (8.0-14.5 days) was found to be shorter than ranges reported in the literature,

speeifieally 13-26 days by Mishra and Pandey (1965), 12-34 days by Badawi (1968), 18-25 days by Sharifi

and Zarea (1973), 11-31 days by Badawi (1981), 16 days (Rafi et al., 1989) and 11-30 days by Rafi et al.

(1999d). The longer periods were refleetive of eool-weather larval development.

We found the length of the pupal stage (5-20 days) to be similar to that reported in the literature,

speeifieally 18 days in Rafi et al. (1989), 7-21 days in Rafi et al. (1999d), 11.7 days in Badawi (1968,

1981), and 9.4-12.2 days in Sharifi and Zarea (1970). Sharifi and Zarea (1970) reported that the pupal

stage of P. demoleus is very sensitive to temperature. Even though they hibernated in winter, any rise in

temperature above 25°C eaused adults to emerge. The same results were observed during present study.

Talbot (1939) reported that greenish pupal eolor was due to pupal development in proximity to leaves or

any other green objeet, and brown pupal eolor was due to proximity to brown objeets. Our observations

support these findings. Tauber et al. (1986) reported that pupal polymorphism does not affeet rate of

development of the adult.

Ants are major predators of P. demoleus eggs and young larvae. Owen (1971) deseribed that 90% eggs are

eaten by ants. Jalali & Singh (1990) and Weeravit-Vittayaluk (1991) reeorded Trichogramma sp. and

Telenomus sp. as egg parasitoids. The Common Garden Lizard (Calotes versicolor) voraeiously attaeks

young and older larvae. Pupae are often infested by the hymenopterous parasitoid (Pteromalus puparum).

Abu-Yaman (1973) from Saudi Arabia, Pipatwatankul (1979) and Napompeth (1981) from Thailand, and

Rafi et al. (1999a) from Pakistan report that a majority of P. demoleus pupae are attaeked by Pteromalus

puparum parasite. Rafi et al. (1989) suggest that P. puparum may be the biggest limiting faetor on P.

demoleus population buildup in Pakistan.

P. demoleus adults are very strong and survive in all seasons in lower Sindh. Males are swift fliers and

disperse over a wide area; females tend to fly nearer to the host plants. We found that adults live 4-7 days,

with males averaging 4.9 days and females 6.7 days. Reported ranges in the literature inelude 1-2 days in

(Ghosh (1914), 5-12 days in Abu-Yaman (1973), 6.21 [mean] days in Farahbakhsh and Khashkooli (1978),

5.12 [mean] days in Badawi (1981), approximately 7 days in Rafi et al (1989), 3.5-7.5 days in Rafi et al.

(1999d), and 4-10 days (mean 6 days) in Winotai and Napompeth (1981). Rafi et al. (1999e) reported the

premating period lasting 6-11 hours, whieh elosely approximates the 4-10 hours observed in the present

study. An earlier study eondueted by Sharifi and Zarea (1970) refleeted these results but some pairs of

adults reportedly delayed mating for 1-2 days after emergenee. In this study, the mating period was

observed to last 5-8 hours, whieh is eonsiderably longer than that reported in literature. Sharifi and Zarea

(1970) reported mating to last 107 + 35 minutes, while Rafi et al (1999e) reported mating to last from 20

minutes to 1 hour and 5 minutes. Atwal (1976) reported that female P. demoleus normally mate only onee

in their lifetime. The same observation was made in this study.

Our study also eonfirmed the reports of Sharifi and Zarea (1970) and Rafi et al (1999e) that mated females

usually start to deposit eggs within two days, while Rafi et al (1989) reported 2-3 days. Sharifi and Zarea

(1970) reported the oviposition period to last 1-4 days, while Winotai and Napompeth (1981) report that

oviposition took plaee over a period of 2-6 days. In our study, females eompleted oviposition within 33-54

hours (1.4-2.2 days), whieh elosely approximates the 35-60 hours (1.5-2.5 days) reported by Rafi et al

(1999e). Sharifi and Zarea (1970) reported that P. demoleus females laid 7-75 eggs with a mean of 31 egg

per females; Farahbakhsh and Khashkooli (1978) likewise reported females deposited 12-77 eggs with an

average of 38.2. Pipatwatankul (1979) reported oviposition with a mean of 96.7 + 31.1 eggs. Rafi et al

(1999e) reported 10-45 eggs with a mean of 26.8 + 13.4. In our study the females deposited 13-40 eggs

with a mean of 27.7 ± 3.3. Interestingly, Winotai and Napompeth (1981) reported a larger range of eggs

(85-240) per female, but this was observed under eontrolled lab eonditions. Sharifi and Zarea (1970)

reported that oviposition took plaee between 9 A.M. and 4 P.M., while Rafi et al. (1999e) reported that

oviposition took plaee between 9 A.M. and 5:30 P.M. In the present study P. demoleus oviposition took

plaee in daylight between 8 A.M. and 6 P.M.

Maxwell-Lefroy (1906) reported that P. demoleus was aetive throughout the year and there were several

generations. We found 13 generations per year in lower Sindh, eaeh lasting 22.5-46.5 days. In the northern

rain fed area of Pakistan, Rafi et al. (1999d) reported 8-9 generations (eaeh lasting 26-167 days depending

on season) with hibernating pupae lasting from November through Mareh. In Iran, Farahbakhsh and

Khashkooli (1978) reported 4-5 generations per year. Wynter-Blyth (1957) reported P. demoleus is found

in large numbers during and after the monsoon in India and flies in all the seasons exeept when the winter

is very eold. Our study found similar patterns and abundanees. Pupae will hibernate during the eoldest

days of Deeember and January.

Papilio demoleus is a major pest of the plant family Rutaeeae (Fleteher, 1917; Pipatwatankul, 1979; Khan,

1940; Atwal, 1976) and ean breed on all varieties of wild and eultivated Citrus. In the present study 10

different varieties of Citrus and Curry Leaf were evaluated as hostplants. The growth index value (G.I.) was

highest on Citrus aurantiifolia (Kaghzi Lime) followed elosely by Citrus reticulata (Citrus Willow) and

Citrus limonia (Common Jamberi). Citrus sinensis (Washington Navel Orange) and Citrus medica (Citron)

had the lowest levels among the Citrus. The lowest G.I. value overall was reeorded on Murraya koenigii

(Curry Leaf). Singh and Gangwar (1989) reported different Citrus affeet the growth rate for P. demoleus in

India. They found the G.I. value was highest in Citrus reticulata, followed by Citrus madurensis, and was

least in Citrus medica. Adult longevity was highest on Citrus reticulata. Rafi et al. (1999b) reported in

their field trial of 11 different Citrus speeies and varieties that ovipositing wild females showed a deeided

preferenee for Citrus aurantium and Citrus reticulata. Females in this study showed similar preferenee for

these two Citrus hosts. Furthermore, in our experienee, plants with greater density of tender leaves had a

higher number of eggs.

Adult swallowtails feed on a variety of food sourees sueh as neetar and pollen (Boggs and Gilbert, 1979;

Nishida et al., 1987). Under experimental eonditions, butterflies fed sugar solutions inerease longevity

and/or egg produetion Stern and Smith (1960), Arms et al. (1974), Morton (1991), Watanabe (1992). In the

present study, our results show empirieal trends indieating adults live longer and lay more eggs while being

fed 15% honey solution. This “naturaf’ solution of sugars and amino aeids produeed better results than

15% suerose solution and 15% glueose.

ACKNOWLEDGEMENTS

Professor Dr. S. Nazneen Rizvi, Chairperson, and Dr. Pervaiz A. Siddiqui, former Chairman, Department

of Zoology, University of Karaehi for providing neeessary faeilities during the progress of this work.

Professor Dr. S. N. H. Naqvi, Department of Zoology (Entomology), University of Karaehi for his eonstant

kind help and eneouragement during this work. Professor Dr. Imtiaz Ahmad and Assistant Professor Dr. S.

Anser Rizvi, Department of Zoology (Entomology), University of Karaehi for their valuable adviee

throughout the progress of this work. Dr. M.A. Matin (N.A.R.C.) for his eontinued interest and valuable

adviee and literature he provided during the duration of this researeh. Mr. Banpot Napon Peth, former

Exeeutive Direetor, National Biologieal Control Researeh Centre, Bangkok, and Mr. Ataul-Mohsin, Ankara

University, Turkey, Dr. Ford William, Entomologieal Researeh Eaboratory, England, and Ms. Madiha

Mustafa, Researeh Seholar, King’s College, Eondon, for providing literature related to this study.

Professor Dr. M. K. Eohar, Department of Entomology, University of Agrieulture, Tando Jam, for his

valuable diseussions. Dr. Khushanood, former Direetor, Pakistan Atomie Energy Agrieulture Researeh

Centre, Tando Jam, for his valuable adviee. Mr. M. Aslam, Assistant Professor, Department of Statisties,

University of Karaehi and Ms. Humaira Ali, Ambreen Ali and Dr. M. Farhanullah Khan (P.A.R.C.) for

their help in reviewing the data. Mr. Azher A. Khan, Department of Zoology, University of Karaehi for

eomposing the original manuseript at the KSK Computation Serviee with his keen sense of interest and

effieieney. Mr. Clive S. Pratt of The Inseet Company, Wiltshire, United Kingdom, for permission to use

photos in the masthead.

LITERATURE CITED

Abu-Yaman, I. K. 1973. Biologieal studies on the eitrus leaf eaterpillar, Papilio demodocus Esp.

(Eepidoptera: Papilionidae) in Saudi Arabia. Zool. Ang. Entomol. 72: 376-383.

Arms, K., P. Feeny and R. C. Eederhouse. 1974. Sodium: Stimulus for puddling behavior by Tiger

Swallowtail butterflies, Papilio glaucus. Seienee 185: 372-374.

Atwal, A. S. 1976. Agrieultural pests of India and South East Asia. Kalyani Publishers, New Delhi, India.

479 pp.

Badawi, A. 1968. Biologieal studies on Papilio demoleus E. a pest of eitrus trees in the Sudan

(Eepidoptera: Papilionidae). Bulletin de a Soeiete Entomologique d’Egypte 52: 397-402.

Badawi, A. 1981. Studies on some aspeets of the biology and eeology of the eitrus butterfly Papilio

demoleus E. in Saudi Arabia (Papilionidae, Eepidoptera). Zeit. Angew. Entomol. 91(3): 286-292.

Boggs, C. E. and E. E. Gilbert. 1979. Male eontribution to egg produetion in butterflies: evidenee for

transfer of nutrients at mating. Seienee 206: 83-84.

Duport, L. 1913. Notes on eertain diseases and enemies of eultivated plants in the Far East. Rev. App.

Entom. Series A, 2: 490.

Eastwood, R., S. E. Boyee and B. D. Farrell. 2006. The Provenanee of Old World Swallowtail Butterflies,

Papilio demoleus (Eepidoptera: Papilionidae), Reeently Diseovered in the New World. Ann. Entomol.

Soe. Am. 99(1): 164-168.

Farahbakhsh, A. and A. Kashkooli. 1978. Biology of the eitrus butterfly, Papilio demoleus demoleus E.

(Eepidoptera, Papilionidae) in Djahrom. Entom. et Phytopathologie Appliquees 46(1-2): 102-109.

Fleteher, T. B. 1917. Report to the Imperial Pathologieal Entomologist. Seientifie Reports, Agrieultural

Researeh Institute Pusa (Caleutta) (9): 64.

Ghosh, C. C. 1914. Eife-histories of Indian inseets. Eepidoptera: The lemon eaterpillar. Mere Dept. Agrie.

India, entom. ser. 5: 33-58.

Government of Pakistan. 1995. Agrieultural Statisties of Pakistan, 1993-94. Ministry of Food, Agrieulture

and Eivestoek. Islamabad, Pakistan. 308 pp.

Guerrero, K. A., D. Veloz, S. E. Boyee, and B. D. Farrell. 2004. First New World doeumentation of an Old

World eitrus pest, the Eime Swallowtail Papilio demoleus (Eepidoptera: Papilionidae), in the

Dominiean Republie (Hispaniola). Ameriean Entomol. 50(4): 227-229.

Homziat, N. T. and J. Homziak. 2006. Papilio demoleus (Eepidoptera: Papilionidae): A new reeord for the

United States, Commonwealth of Puerto Rieo. Florida Entomologist 89(4): 485-487.

Jalali, S. K. and S. P. Singh. 1990. A new reeord of Ooencyrtus papilionis (Hymenoptera: Eneyrtidae) on

the egg of Papilio demoleus (Einn.) from India. J. Biologieal Control 4(1): 59-60.

Khan, A. R. 1940. Inseet Pest number, Punjab Agrieultural College, Eyallpur. Punjab Fruit Journal 13: 40-

41.

Malik, J. M. 1970. Notes on the butterflies of Pakistan in the eolleetion of Zoologieal Survey Department

Karaehi, Part 1. Ree. Zool. Surv. Pakistan 2(2): 25-54.

Maxwell-Eefroy, H. 1906. Indian Inseet Pests. Gov’t. Printing Press, Caleutta, 174 pp.

Mishra, S. C. and Pandey, N. D. 1965. Some observations on the biology of Papilio demoleus Einn.

(Papilionidae, Eepidoptera). Eabdev. J. Sei. Teehnol. 3: 142-143.

Morton, A. C. 1991. Rearing butterflies on artifieial diets. J. Res. Eepid. 18(4): 221-227.

Mushtaque, M. 1964. Studies on the biology and history of lemon butterfly, Papilio demoleus Einn. M.S.

Thesis, Sindh Agrieulture University, Tandojam, Sindh, Pakistan, 66 pp.

Napompeth, B. 1981. Progress of biologieal eontrol in Thailand. National Biologieal Control Researeh

Center, Kasetsart Univ., App. Ent. Ser. 70: 3.

Nishida, R., T. Ohsugi, S. Kokubo and H. Fukami. 1987. Oviposition stimulants of a eitrus-feeding

swallowtail butterfly, Papilio xuthus E. Cellular and Moleeular Eife Seienees 43(3): 342-344.

Owen, D. F. 1971. Tropieal butterflies: The eeology and behavior of butterflies in the tropies with speeial

referenee to Afriean speeies. Clarendon Press, Oxford, 214 pp.

Pipatwatankul, A. 1979. Eeologieal investigation on the Eemon Butterfly Papilio demoleus E.

(Eepidoptera: Papilionidae) and its natural enemies in Thailand. M.S. Thesis. Kasetsart University,

Bangkok, Thailand, 50 pp.

Radke, S. G. and Kandalkar, H. G. 1989. Observations on the biology of Eemon Butterfly, Papilio

demoleus Einnaeus (Eepidoptera: Papilionidae). PKV Researeh J. 13(2): 176-177.

Rafi, M. A., M. A. Matin and S. A. Saghir. 1989. Studies on some bio-eeologieal aspeets of the eitrus

butterfly Papilio demoleus E. (Eepidoptera: Papilionidae) in the Barani eeology of Pakistan. Pakistan.

J. Sei. and Ind. Res. 32(1): 36-38.

Rafi, M. A., M. Irshad and M. A. Matin. 1999a. Bio-eeology of Papilio demoleus E. (Eepidoptera:

Papilionidae) and effieaey of Baetospeine against its larvae. Pakistan J. Entom. Karaehi 6(1-2): 53-61

Rafi, M. A., M. R. Khan and M. Ilyas. 1999b. Host preferenee of lemon butterfly Papilio demoleus E. in

the Northern Barani areas of Pakistan. Pakistan J. Seienee 51(3-4): 93-94.

Rafi, M. A., M. A. Matin and M. R. Khan. 1999e. Biology of eggs of eitrus butterfly Papilio demoleus L.

(Papilionidae: Lepidoptera). Pakistan J. Seienee 51(3-4): 95-99.

Rafi, M. A., M. A. Matin and M. R. Khan. 1999d. Number of generations and their duration of Lemon

Butterfly {Papilio demoleus L.) in rainfed eeology of Pakistan. Pakistan. J. Sei. Res. 51(3-4): 131-136.

Roberts, T. J. 2001. The butterflies of Pakistan. Oxford University Press, Karaehi. 200 pp.

Sharifi, S. and N. Zarea. 1970. Biology of the eitrus butterfly, Papilio demoleus demoleus (Lepidoptera:

Papilionidae). Ann. Ent. Soe. Ameriea. 63(5): 1211-1213.

Singh, Y. P. and S. K. Gangwar. 1989. Biology of the lemon butterfly {Papilio demoleus L.) on Khasi

mandarin and its development on eitrus eultivars. J. Andaman Seienee Assoe. 5(2): 151-153.

Srivastava, K. P. and Y. S. Ahlawat. 1999. Pest Management in Citrus. Researeh Periodieals and Book

Publishing House. New Delhi, India. 320 pp.

Stern, V. M. and R. F. Smith. 1960. Faetors affeeting egg produetion and oviposition in populations of

Coliasphilodice eurytheme Boisduval (Fepidoptera: Pieridae). Hilgardia 29: 411-454.

Talbot, G. 1939. The Fauna of British India, Butterflies, Vol. 1. (Covers Papilionidae and Pieridae).

Today and Tomorrow Printers and Publishers, New Delhi, India. 600 pp.

Wasif, T. A. A. 1991. Common Plants of Pakistan (Trees and Bushes). Nat. Fang. Islamabad, Pakistan, pp.

146-151.

Watanabe, M. 1992. Egg maturation in laboratory-reared females of the swallowtail butterfly, Papilio

xuthus F. (Fepidoptera: Papilionidae), feeding on different eoneentration solutions of sugar. Zool. Sei.

9(1): 133-141.

Weeravit-Vittayaluk. 1991. Eeologieal investigation on the lemon butterfly Papilio demoleus F.

(Fepidoptera: Papilionidae) and its behaviour on Pummelo kasatsart. Univ. Bangkok (Thailand),

Graduate Sehool, pp. 75.

Winotai, A. and B. Napompeth. 1981. Femon butterfly {Papilio demoleus F.) (Fepidoptera: Papilionidae)

and its natural enemies in Thailand. Proe. Nat. Conferenee on the Progress of Biologieal Control in

Thailand, Bangkok, 1981. 7 pp.

Wynter-Blyth, M. A. 1957. Butterflies of Indian region. Bombay Nat. Hist. Soe. 253 pp.

The Taxonomic Report

is a publication of

The International Lepidoptera Survey (TILS)

(a tax exempt non-profit scientific organization)

The Taxonomic Report is published for the purpose of providing a public and permanent scientific record.

It appears on printed paper in sequential issues, is regularly disseminated to institutional and individual

subscribers, and is also available as separate issues free of charge upon request at the discretion of authors

and/or the editor. Contents are peer-reviewed but not necessarily through the anonymous review and

comment process preferred by some publishers of serial literature.

TILS Purpose

TILS is devoted to the worldwide collection of Lepidoptera for the purpose of scientific discovery,

determination, and documentation, without which there can be no preservation.

TILS Motto

“As a world community, we cannot protect that which we do not know”

Articles for publication are sought

They may deal with any area of research on Lepidoptera, including faunal surveys, conservation topics,

methods, etc. Taxonomic papers are especially welcome. Before sending a manuscript, simply write to

TILS editor, Harry Pavulaan, P.O. Box 1124, Herndon, VA 20172 to set up discussion on how to best

handle your material for publication; or email harrypav@hotmail.com

Donations are needed

to support and further our efforts

to discover and protect butterflies worldwide.

All donations are US tax deductible. Please help generously.

Donations should be mailed to: TILS, c/o Harry Pavulaan, P.O. Box 1124, Herndon, VA 20172.

Checks should be made payable to: TILS. Please indicate if you need an individual receipt.

Visit The International Lepidoptera Survey on the World Wide Weh at: http://www.tils-ttr.org

Join the discussion at our list serve on Yahoo! Groups at: http://groups.yahoo.com/group/TILS-leps-talk/