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common name: pipevine swallowtail, blue swallowtail
scientific name: Battus philenor (Linnaeus 1771) (Insecta: Lepidoptera: Papilionidae: Papilioninae: Troidini)

Introduction - Nomenclature - Distribution - Description - Host Plants - Life Bicycle - Natural Enemies - Defenses - Mimicry - Acknowledgements - Selected References

Introduction (Back to Top)

The pipevine swallowtail, Battus philenor (Fifty.), is one of our most cute swallowtails. It is too known as the bluish swallowtail (e.g., Howe 1988, Iftner et al. 1992). Its life cycle was beautifully illustrated during the 18th century past John Abbot (Smith 1797) (Figure 1).

Figure 1. Life bike of the pipevine swallowtail, Battus philenor (L.). Drawing by John Abbot (From Smith 1797). (For aesthetic reasons, the color temperature of the epitome was adjusted to compensate for yellowing of the manuscript due to historic period).

Nomenclature (Back to Pinnacle)

The pipevine swallowtail was originally described by Linnaeus (1771) and placed in the genus Papilio with the other swallowtails. Information technology was later moved to the genus Battus (Scopoli 1777). The proper name "Battus" is from Battus I, founder of the ancient Greek colony Cyrenaica and its capital, Cyrene, in Africa. The specific epithet is from the Greek word "philenor" which means addicted of married man or conjugal (Opler & Krizek 1984).

Larvae of the pipevine swallowtail and those of the other swallowtails belonging to the tribe Troidini feed on plants in the genus Aristolochia and are ordinarily referred to as the Aristolochia swallowtails.

For synonymy, see the excerpt from Pelham (2008) at the Butterflies of America web page on Battus philenor (Accessed December 23, 2016).

Distribution (Back to Top)

The U. S. distribution of the pipevine swallowtail extends from southern Connecticut s to cardinal Florida and w to Arizona with an isolated population in northern California (Figure two). The pipevine swallowtail is likewise found north to southeastern Ontario, Canada and s to southern Mexico (Cech & Tudor 2005, Opler & Malikul 1998, Scott 1986).

Figure ii. Pipevine swallowtail, Battus philenor (L.), U.S. distribution map.

Published distribution maps (e.g., Brock & Kaufman 2003, Scott 1986, Moth Photographers Group, Battus philenor species page, Butterflies and Moths of North America) differ due to records of adult "strays" and to planting of larval host plants as ornamentals (Iftner et al. 1992, Layberry et al. 1998) outside their native ranges. For instance, the simply documented records for Battus philenor larvae in Ontario are from Aristolochia macrophylla vines planted as ornamentals (Layberry et al. 1998).

Description (Back to Peak)

Adults: The wingspread is 2 3/4-five 1/8 in. (72-132 mm) (Opler & Malikul 1992). The dorsal surfaces of the wings of males are mostly black with blue or blue-green iridescence on the hind wings (Effigy 3). The dorsal aspect of the hindwings of females (Figure 4) has duller iridescence than that of males. At that place are light marginal and sub-marginal spots on both forewings and hindwings. The ventral hind wings of both sexes accept blue iridescence and a row of vii bright orangish sub-marginal spots (Figures 4 & 5). There is a row of white spots on the lateral aspect of the abdomen (Figure 5).

Effigy 3. Developed male pipevine swallowtail, Battus philenor (L.), dorsal view showing bright iridescence. Photograph by Donald West. Hall, Entomology and Nematology Section, University of Florida.

Figure iv. Adult female pipevine swallowtail, Battus philenor (L.), dorsal (left) and ventral (right) views. Photograph by Donald Westward. Hall, Entomology and Nematology Department, University of Florida.

Figure 5. Newly-emerged adult male person pipevine swallowtail, Battus philenor (L.), with wings folded showing undersides of the wings and white spots on abdomen. Photograph by Donald Westward. Hall, Entomology and Nematology Department, Academy of Florida.

California specimens are smaller, with hairy bodies. Their larvae likewise have shorter thoracic filaments than those of eastern specimens. The California population also differs from eastern populations in some aspects of biology (Sims & Shapiro 1983b) and may represent a different subspecies, Battus philenor hirsuta (Sourakov & Daniels 2002). According to Field (1938, p. 200), individuals of the spring generation in Kansas resemble those from California populations.

Eggs: The eggs are ruby-red-orange (Figure 6) in contrast to those of the related Polydamas swallowtail, Battus polydamas (L.), which are yellow or yellowish-orange. The eggs of all Aristolochia (troidine) swallowtails are partially covered by a hard, nutritious secretion that is usually laid down in vertical bands (Comstock & Grimshawe 1935, Tyler et al. 1994). In that location are numerous relatively big droplets on the bands. The secretion is produced past a big gland that lies above the female's ovipositor duct (Tyler et al. 1994).

Figure half-dozen. Eggs of the pipevine swallowtail, Battus philenor (Fifty.). Photograph by Donald W. Hall, Entomology and Nematology Section, Academy of Florida.

Larvae: Full-grown larvae are approximately 50 mm (nigh 2 in.) in length (Minno et al. 2005, Stehr 1987, Wagner 2005).

First instar larvae have numerous short orange tubercles – each with a unmarried seta (Effigy 7). 2nd instars accept longer tubules - each bearing multiple setae (Figure 8). The tubercles of third and quaternary instars are proportionately longer, and the exoskeleton takes on a slightly glossy appearance (Figure 9).

Figure 7. First instar larva of the pipevine swallowtail, Battus philenor (L.). Photo by Donald West. Hall, Entomology and Nematology Department, Academy of Florida.

Figure viii. Second instar larva of the pipevine swallowtail, Battus philenor (L.). Photograph past Donald W. Hall, Entomology and Nematology Department, University of Florida.

Figure 9. 3rd instar larva of the pipevine swallowtail, Battus philenor (L.). Photograph by Donald West. Hall, Entomology and Nematology Department, Academy of Florida.

Full-grown larvae are typically dark dark-brown to black with sub-dorsal and lateral rows of bright orangish tubercles (Figure 10), but some are red (Figure eleven). In western Texas and southern Arizona, the ruby-red form predominates under conditions of higher temperature (in a higher place about 30°C) (Nice & Fordyce 2006, Nielsen & Papaj 2015, Papaj & Newsome 2005). The factor(s) responsible for ruby-red larvae in the eastern U.S. has not been investigated. The cerise larva in Figure 11 was photographed in Gainesville, Florida on July 29, 2016, during the heat of the summertime.

Lateral tubercles on the thorax and abdominal segments 2, 7, 8, and nine of larvae are modified into elongated filaments. The filaments on the prothorax are particularly long (Scott. 1986, Minno et al. 2005, Stehr 1987). Total-grown larvae are covered with fine hairs that give them a velvety appearance (Howe 1975).

Figure 10. Total-grown night larva of the pipevine swallowtail, Battus philenor (L.). Photograph by Donald Westward. Hall, Entomology and Nematology Department, University of Florida.

Figure 11. Full-grown red larva of the pipevine swallowtail, Battus philenor (L.). Photograph by Donald Due west. Hall, Entomology and Nematology Section, Academy of Florida.

The sex of larvae of all ages tin be determined past the structure of sex-specific pits on the ventral surface of the 8th and 9th abdominal segments (Underwood 1994).

Pupae: Pupae may be either green or brown (Figure 12). Unlike pupae of other swallowtails, the sides of the bodies of Battus pupae are widened into lateral flanges. In the lateral views of pupae shown in Effigy 12, the lateral flanges announced as bluish-purple ridges along the anterior half of the belly.

Effigy 12. Light-green pupa and brown pupa of the pipevine swallowtail, Battus philenor (50.). Photographs past Donald West. Hall, Entomology and Nematology Department, University of Florida.

Host Plants (Back to Height)

Larval host plants: Pipevine swallowtail larvae feed on plants belonging to the genus Aristolochia in the family Aristolochiaceae (Cech & Tudor 2005, Minno et al. 2005, Opler & Krizek 1984, Scott 1986). Reports in the literature (eastward.yard., Arnett 2000, Howe 1975, Robinson et al. undated, Schull 1987, Tyler 1975) of larvae feeding on wild ginger, Asarum canadense L. (Aristolochiaceae), knotweed, Polygonum (Polygonaceae), and morn glories, Ipomoea (Convolvulaceae), are probably erroneous (Heitzman & Heitzman 1987, Opler & Krizek 1984, Scott 1986) and likely due to misidentification of the host found by Blatchley (1896) in the case of Asarum (see Weintraub 1995) and to wandering larvae for the other institute species.

Aristolochia species are commonly known as pipevines or Dutchman's pipes because the flowers of some species are shaped like tobacco pipes (Figure xiii). They are also known as birthworts ("wort" is Sometime English for herbaceous plant) considering of their historical use in child birth. The proper name Aristolochia is derived from the Greek roots aristos (all-time) and lochia (delivery or child nascency) (Crosswhite & Crosswhite 1985, Flora of North America undated). All Aristolochiaceae are believed to contain pharmacologically active aristolochic acids (Chen & Zhu 1987).

Figure xiii. Unopened flower of woolly pipevine, Aristolochia tomentosa Sims, and tobacco pipe graphic showing the similarity of shapes. Photo of blossom bud past Donald W. Hall, Entomology and Nematology Section, University of Florida. Tobacco pipe graphic from PNG Clip Art (graphic modified and pasted into photograph by Donald Westward. Hall, University of Florida).

Although they are at present officially banned in many countries, Aristolochia-derived herbal products or parts of the plants themselves are still used in many areas of the world for various conditions including snake bite, gastrointestinal issues, respiratory problems, wounds, infectious diseases, and fever (Austin 2004, Chen & Zhu 1987, Knuckles 2001, Schaneberg et al. 2002).

Virginia snakeroot, Aristolochia serpentaria Fifty., has been used for many medical applications (Austin 2004, Knuckles 2001, Heinrich et al. 2009, Moerman 1998), and preparations made from it are nonetheless for auction online. An excerpt of the southwestern pipevine, Aristolochia watsonii Wooton & Standl., was the main ingredient in the snakeroot oil sold by traveling "snakeroot doctors" at medicine shows in the Old West during the 19th century (Crosswhite & Crosswhite 1985). Aristolochic acids in the products accept been implicated every bit causative agents of renal toxicity and also may be carcinogenic (Heinrich et al. 2009, Schaneberg et al. 2002).

Aristolochia serpentaria 50., ranges from primal Florida northward in much of the eastern Us. In the Southeast, it has broad-leaved and narrow-leaved forms (Figures fourteen, 15 & 16). The broad-leaved form is more than common in richer mesic forests while the narrow-leaved form is more than common in drier, sandy areas (Allard 2002, D.W. Hall personal ascertainment).

Figure 14. Virginia snakeroot, Aristolochia serpentaria L. (broad-leaved class), a host of the pipevine swallowtail caterpillar, Battus philenor (Fifty.), with bloom. Photographs by Donald W. Hall, Entomology and Nematology Department, University of Florida.

Figure xv. Virginia snakeroot, Aristolochia serpentaria L. (broad-leaved form), a host of the pipevine swallowtail caterpillar, Battus philenor (50.), with seed capsules. Photographs past Donald W. Hall, Entomology and Nematology Department, University of Florida.

Figure 16. Virginia snakeroot, Aristolochia serpentaria 50. (narrow-leaved form), a host of the pipevine swallowtail caterpillar, Battus philenor (L.). Photo by Donald W. Hall, Entomology and Nematology Section, University of Florida.

Aristolochia serpentaria 50. and Aristolochia macrophylla Lam. (Effigy 17) take been assayed for aristolochic acid, and both species were adamant to comprise significant quantities of aristolochic acrid I (Schaneberg et al. 2002). Because of their toxicity and distastefulness, the aristolochic acids play a major role in the biology of pipevine swallowtails.

All of our native species of Aristolochia within the range of the pipevine swallowtail are documented larval hosts (for distribution maps see their corresponding USDA Plants Database species pages – links below):

• Virginia snakeroot, Aristolochia serpentaria L. (Figures 14, fifteen &16) USDA Plants Database species page.
• Pipevine, Aristolochia macrophylla Lam. (synonym, Aristolochia durior Hill) (Figure 17) USDA Plants Database species page.
• Woolly Dutchman's pipe, Aristolochia tomentosa Sims. (Figure 18) USDA Plants Database species page.
• Texas Dutchman's pipe, Aristolochia reticulata Jacq. USDA Plants Database species folio.
• Watson's Dutchman's pipe, Aristolochia watsonii Wooton & Standl. USDA Plants Database species page.
• California Dutchman's pipage, Aristolochia californica Torr. USDA Plants Database species page.

Figure 17. Pipevine, Aristolochia macrophylla Lam. (synonym, Aristolochia durior Hill) a host of the pipevine swallowtail caterpillar, Battus philenor (L.). Photograph by Donald W. Hall, Entomology and Nematology Department, University of Florida.

Figure 18. Woolly Dutchman'south pipe, Aristolochia tomentosa Sims., a host of the pipevine swallowtail caterpillar, Battus philenor (L.), leaf (left), unopened bloom (middle), opened flower (right). Photographs past Donald W. Hall, Entomology and Nematology Department, University of Florida.

Diverse exotic Aristolochia species are planted every bit ornamentals because of their unusual and sometimes beautiful flowers. Some of these may be as well toxic (or too distasteful) for pipevine swallowtail larvae and may be "expiry traps" for the larvae. Elegant Dutchman'southward pipe or calico blossom, Aristolochia elegans Mast (synonym, Aristolochia littoralis Parodi) (Figure 19), is attractive to female pipevine swallowtails for oviposition, merely larvae usually do not survive on it (Kendall 1964, Scott 1986, Tyler 1975). Kendall (1964) suggested that Aristolochia elegans is probably simply distasteful and not toxic to Battus philenor larvae and that the larvae probably die due to starvation considering of their refusal to eat it. Therefore, planting of Aristolochia elegans and other exotic pipevines is not recommended where Battus philenor (Fifty.) occurs (fundamental Florida and north).

Figure nineteen. Elegant Dutchman's pipe or calico flower, Aristolochia elegans One thousand.T. Mast [synonym: Aristolochia littoralis Parodi], a cultured exotic which is a death trap for the pipevine swallowtail caterpillar, Battus philenor (L.). Photograph by Donald Due west. Hall, Entomology and Nematology Department, Academy of Florida.

Seed Dispersal: Aristolochia tomentosa and Aristolochia macrophylla are tall vines (Pfeifer 1966) with flattened seeds. Their seeds are probably dispersed by wind and possibly likewise by animals and flowing h2o. Two Russian Aristolochia vines with similar seeds that grow in like habitats have been shown to be primarily dispersed past wind and water and to a lesser extent by birds (Nechaev & Nakonechnaya 2009).

Aristolochia serpentaria is a small herb. Its seeds have an attached oil trunk (elaiosome), and the seeds are dispersed by ants (myrmecochory) that collect the seeds for the edible elaiosomes (D.W. Hall, personal ascertainment, Osorio 2010) (Figure twenty).

Figure 20. Myrmecochory (ant dispersal) of Virginia serpent root, Aristolochia serpentaria L., seed by trap-jaw emmet, Odontomachus brunneus (Patton): a) dehisced (opened) seed capsule showing seeds with attached oil bodies (elaiosomes). b) trap-jaw pismire, Odontomachus brunneus (Patton), removing seed from capsule c) trap-jaw pismire, Odontomachus brunneus (Patton), carrying seed by the elaiosome. Photographs by Donald Westward. Hall, Entomology and Nematology Department, Academy of Florida.

Both Aristolochia serpentaria and Aristolochia reticulata are minor herbs, and their flowers (and seeds) are usually formed near the bases of the plants, commonly underground or at to the lowest degree under leafage litter. Rausher and Feeny (1980) suggested that this may be an adaptation to protect the flowers and seeds from predation past Battus philenor caterpillars. However, it may also be an adaptation for dispersal of the seeds by ants. The mechanism of dispersal of seeds for Aristolochia reticulata is unknown simply probable as well involves ants.

Pollination:

Insect pollination: Aristolochia species have fascinating pollination biologies (Proctor et al. 1996). When the flowers first open, the stigmas are receptive, but the anthers are not releasing pollen – a phenomenon known as protogyny (from the Greek roots "proto" [first] and "gyne" [female person]). During the kickoff day of bloom, the flowers are attractive to flies (ordinarily small flies - ofttimes, but non e'er, scuttle flies in the family Phoridae [Hall & Brown 1993]) which enter the flowers conveying pollen from another flower. The flies are so prevented from leaving by the presence of downward projecting guard hairs or tiny spines and slippery surfaces in the tube of the flower (Proctor et al. 1996, Zomlefer 1994) (Figures 21, 22, & 23).

Figure 21. Virginia snakeroot, Aristolochia serpentaria L., bloom: forepart view (left), side view (correct), and inside of tube with slippery surface and small downward pointing hairs (inset). Photographs by Donald W. Hall, Entomology and Nematology Department, University of Florida.

Effigy 22. Woolly pipevine, Aristolochia tomentosa Sims, blossom (left) and longitudinal section (right) showing inside of tube with slippery surface and small downward pointing spines (inset). Photographs by Donald Westward. Hall, Entomology and Nematology Section, University of Florida.

Effigy 23. Elegant pipevine, Aristolochia elegans 1000.T. Mast (synonym: Aristolochia littoralis Parodi). a ) flowers in front and side view orientations and down pointing guard hairs (inset). b) cartoon of longitudinal department of bloom showing inside of tube with downwards pointing guard hairs. Photographs by Donald W. Hall, Entomology and Nematology Department, University of Florida. Drawing by Margo Duncan.

On the 2nd solar day (later on pollination), the stigmas become non-receptive, and the anthers dehisce (open up) dusting the flies with pollen. The baby-sit hairs in the flower tubes wither, and in some species (e.g., Aristolochia gigantea Mart. & Zucc.) the flowers also droop into a more horizontal position (Burgess et al. 2004) releasing the flies that are now carrying the new pollen.

Cleistogamy: In add-on to its open (chasmogamous) insect-pollinated flowers, Aristolochia sepentaria also has flowers that never open (cleistogamous) and are self-pollinated (Ahles 1959, Pfeifer 1966). The terms chasmogamous and cleistogamous are from Greek roots ("chasma" [open up or gaping], "cleist", [closed], and "gam", [marriage]) (Borror 1960). Cleistogamy is really a common phenomenon in angiosperms (flowering plants). It is known from at least 693 species in 228 genera and fifty families (Culley & Klooster 2007).

The cleistogamous flowers of Aristolochia serpentaria (Figures 16 & 24) and their resultant seed capsules (Figure 24) are lilliputian more half the size of the chasmogamous flowers and their seed capsules (Figures 14 & 15). Likewise, the cleistogamous flowers do not have the characteristic "tobacco pipe" shape as they practice not trap insects for pollination. Both cleistogamous and chasmogamous flowers often occur on the same found. In dissimilarity to the chasmogamous flowers, the cleistogamous flowers are often formed later in the flavor in some species (Mondal 2016) and higher on the plant (Figure 24) where those of Aristolochia serpentaria are more susceptible to being eaten by Battus philenor larvae.

Figure 24. Virginia snakeroot, Aristolochia serpentaria L., plant with insets showing sequential stages of evolution of cleistogamous flower to mature seed capsule. Photographs past Donald West. Hall, Entomology and Nematology Department, Academy of Florida.

Nectar host plants: At that place are many plants that are valuable every bit nectar sources for butterflies. Pink and purple flowers (east.chiliad., phlox [Phlox species], ironweed [Vernonia species], and thistles [Cirsium species]) are particularly bonny to pipevine swallowtails (Scott 1986). Minno and Minno (1999) accept extensive lists of both native and exotic nectar plants for butterflies. When possible, native plants should be planted equally nectar sources rather than exotics that have the potential to be invasive.
About states have native constitute societies that are valuable sources of information on native plants, and many also hold native plant sales. For Florida and the Deep South, the Florida Wildflowers Growers Cooperative is an excellent source of information and also has wildflower seeds for purchase.

To maximize butterfly populations in yards, both caterpillar hosts and nectar plants for adults should be planted.

Life Cycle (Back to Top)

There are three or more generations in the Deep Southward (Gulf of Mexico area and peninsular Florida) and two generations northward (Cech and Tudor 2005, Glassberg et al. 2000, Howe 1975).

Males suck wet from mud to obtain nutrients (particularly sodium) and allure to the mud appears to exist enhanced by the presence of other males (Otis et al. 2006). Males transfer some of the sodium to the female person in the spermatophore as a nuptial gift during mating (Otis et al. 2006).

Mating: Males patrol host plants and flyways to locate females (Lederhouse 1995). Courtship flights are boring with the male hovering above the female (Cech and Tudor 2005). Battus males have androconia (scent scales) hidden in a fold of the inner margin on the upper surface of the hind wing (Field 1938) (Figure 25). The scales are fluted - possibly to increase surface surface area for release of pheromone (Miller 1987, Racheli & Oliverio 1993). When courtship, Battus males are reported to "helicopter" around the females while fanning the androconial chemicals over them (Tyler et al. 1994, p. 53). However, Westward (1983) reported an ascertainment of mating of Battus philenor without fanning of the wings. He also described a method for hand-pairing them in captivity.

Effigy 25. Pipevine swallowtail, Battus philenor (L.), male person showing iridescent scales (left inset) and androconia (right inset). Photographs by Donald W. Hall, Entomology and Nematology Department, Academy of Florida. Lyle J. Buss assisted with the photograph of androconia.

The brilliant iridescent blueish on the dorsal surface of the hind wings of males is believed to serve as a sexual signal for recognition and mayhap for assessment of male quality by females (Rutowski et al. 2010, Rutowski & Rajyaguru 2013). For a general review of the functions of iridescence in animals, come across Doucet and Meadows (2009).

Oviposition: Females search for larval host plants based on foliage shape (Allard & Papaj 1996, Rausher 1978). Plants with new leaf buds are more attractive for oviposition (Papaj 1986b). If females find either eggs or larvae on a plant they avert ovipositing on information technology (Rausher 1979).

Females volition too investigate not-host plants with leaves shaped similar those of the local host plants. In Florida, they frequently visit young Smilax (Smilacaceae) vines that resemble Aristolochia serpentaria (Feeny 1991). Papaj (1986a) observed a female examine a Smilax laurifolia L. found repeatedly before finally depositing an egg on it. Nonetheless, when he transferred larvae to Smilax, they all died. Oviposition mistakes past Lepidoptera are not uncommon. In fact, long-tailed skippers (Urbanus proteus [L.]) will oviposit on Aristolochia tomentosa, but the larvae are only able to survive on information technology during the early instars (Effigy 26) and die before reaching maturity (Marc Minno [personal advice], D.W. Hall [personal observation]).

Effigy 26. Early instar long-tailed skipper (Urbanus proteus [Fifty.]) leaf shelter on woolly pipevine, Aristolochia tomentosa Sims. Inset: leaf shelter opened to show larva. Photographs by Donald W. Hall, Entomology and Nematology Department, University of Florida.

Aristolochic acids I and Ii, the inositols D-(+)-pinitol and sequoyitol, and a monogalactosyl diglyceride (all isolated from Aristolochia macrophylla) were demonstrated to serve equally synergistic contact oviposition stimulants for Battus philenor (Sachdev-Gupta et al. 1993). None of the chemicals by themselves had pregnant action.

Eggs are laid on young foliage or on stems at the bases of leaves - either singly or in small groups on pocket-sized vines or in larger groups on large vines (Allen 1997. Minno et al. 2005). Young larvae are gregarious, but older larvae are solitary (Allen 1997, Scott 1986).

In Lepidoptera eggs, a small quantity of yolk remains trapped between two of the embryonic membranes (amniotic and serosa) (Barbier & Chauvin 1976) that remain inside the egg shells (chorions) after hatching. Soon after hatching, larvae eat the egg shells (Effigy 27), and the balance yolk serves as their first meal (Richards & Davies 1977). Larvae as well swallow their exuviae after molting to conserve nutrients (Effigy 28).

Figure 27. Pipevine swallowtail, Battus philenor (L.), newly-emerged first instar larvae eating egg shells (chorions). Photograph by Donald Westward. Hall, Entomology and Nematology Department, Academy of Florida.

Figure 28. Pipevine swallowtail, Battus philenor (Fifty.), terminal instar larva eating its exuviae. Photograph by Donald W. Hall, Entomology and Nematology Department, University of Florida.

Battus larvae commonly defoliate their host plants (Tyler et al. 1994) and demand to wander in search of new plants. The elongated thoracic filaments of larvae take only tactile sensors and no chemosensory structures. These filaments and the lateral stemmata (eyes) announced to merely aid larvae locate vertical objects, which must then be identified equally host or non-host past the antennae and mouthparts (Kandori et al. 2015). Wandering larvae may find new host plants by detection of plant volatiles. Aristolochia volatiles have been shown to be attractive to larvae of Battus polydamas (Pinto et al. 2009).

When full-grown, larvae (prepupae) wander off the host constitute to find a pupation site. Pupation occurs well off the ground - usually on tree trunks or on cliff faces in the Appalachians (West & Hazel 1979). Larvae rarely pupate on greenish substrates (Hazel 1995). Before pupation they spin a silk girdle for support and a silk pad which they grasp with their terminal prolegs (Figure 29). Afterwards splitting the old larval exoskeleton and wriggling complimentary, the new pupa hooks the cremaster at the tip of its abdomen into the silk pad (Effigy 29).

Figure 29. Pipevine swallowtail, Battus philenor (L.), prepupa beginning pupation. Note the silk girdle. Photograph by Donald W. Hall, Entomology and Nematology Section, University of Florida.

Figure 30. Pipevine swallowtail, Battus philenor (50.), prepupa molting to pupal stage. Annotation the successive positions of the shed larval caput capsule (arrows). Photographs by Donald W. Hall, Entomology and Nematology Department, University of Florida.

Chocolate-brown pupae predominate (Hazel & Westward 1979, Westward & Hazel 1979). Pupal color is partially influenced by the texture of the pupation substrate but not by the photoperiod during the larval phase (Hazel & West 1979). Short larval photoperiod induces pupal diapause (Hazel & West 1983).

Shortly before emergence of adults, the patterns of the wings and body bear witness through the pupal exoskeleton (Figure 31a). The developed emerges through a serial of cleavage lines: a mid-dorsal cleavage line on top of the thorax, lines between the head and thorax, and lines between the thorax and wings (Scott 1986) (Figure 31b-d). Subsequently emergence, the adult rests nearly the erstwhile pupal case (Effigy 31e) while expanding the wings and joining the 2 halves of the proboscis (tongue) together. During this fourth dimension, it also voids droplets of the scarlet, liquid, pupal waste product products (meconium) (Effigy 31e [inset]).

Figure 31. Pipevine swallowtail, Battus philenor Fifty., stages in emergence of adult from pupal stage. Inset: droplet of pupal waste product (meconium). Photographs by Donald W. Hall, Entomology and Nematology Department, University of Florida.

Natural Enemies (Back to Meridian)

Stamp (1986) observed a third instar coccinellid larva (Hippodamia convergens Guerin-Meneville) feeding on a 2nd instar pipevine swallowtail larva.

Records of parasitoids from Battus philenor are rare. However, there are records of at to the lowest degree one tachinid fly (Compsilura concinnata (Meigen)) (Arnaud 1978, p. 601) and two ichneumonid wasps (Theronia atalantae (Poda) and Apechthis annulicornis (Cresson)) (Sime 2000). Sims and Shapiro (1983a) reported the pupal parasitoid Brachymeria ovata (Say) (Hymenoptera: Chalcididae) from California Battus philenor and also noted significant mortality to pupae from a "disease" of unknown causes - possibly mucus.

Howe (1988) reported many species of birds eating Battus philenor larvae with "enjoy". Anolis lizards volition also eat the larvae (Odendaal et al. 1987).

Defenses (Back to Pinnacle)

Sequestered aristolochic acids: Battus philenor larvae sequester aristolochic acids from their host plants and the acids are passed on to the pupae, adults, and ultimately to eggs of the next generation (Dimarco et al. 2012, Dimarco & Fordyce 2013, Sime et al. 2000). The aristolochic acids are reported to render larvae, pupae, and adults unpalatable to some birds. Adults take tough bodies that allow them to survive being tasted (Scott 1986).

Contradictory reports on the palatability of larvae to birds may be due to differences in the power of some populations (families) of Battus philenor to sequester aristolochic acids (Dimarco et al. 2012). Too, lower aristolochic acrid concentrations in some Aristolochia species or variations in concentrations between individuals of the same species may explicate the contradictions.

The chemical defenses announced to be effective confronting some parasitoids. The ichneumonid wasp Trogus pennator (Fabricius), which normally parasitizes papilionid larvae, rejected Battus philenor larvae afterwards examining them with their antennae (Sime 2002).

Osmeteria: All United States swallowtail larvae have forked, eversible, horn-like organs behind the head known as osmeteria (singular, osmeterium). The osmeteria of Battus philenor larvae are brilliant yellow in all larval instars. Fifth instar Papilio species secrete isobutyric and ii-methyl butyric acids from their osmeteria (Ômura et al. 2006). The chemical makeup of the osmeterial secretion of fifth instar Battus philenor is unknown. However, the secretion of quaternary and fifth instar larvae of the closely-related Battus polydamus reared on Aristolochia elegans is composed of the ii sesquiterpenes: β-selinine and selin-eleven-en-4α-ol (Eisner et al. 1971). These compounds were non found in the host institute in detectable amounts and are causeless to be synthesized past the larvae instead of beingness sequestered (Eisner et al. 1971).

The osmeterial repellent of Papilio species is constructive against ants (Eisner and Meinwald 1965), but Berenbaum et al. (1992) reported that soldier bugs (Hemiptera: Pentatomidae) could attack and eat swallowtail larvae (Papilio species) without evoking extrusion of the osmeteria.

Aposematism (warning coloration or appearance): The orange color on the ventral surface of the wings and the blue iridescence on both ventral and dorsal surfaces have been demonstrated to office as aposematic coloration against assail by vertebrate predators (Pegram et al. 2013, Pegram et al. 2015, Pegram & Rutowski 2014, Pegram & Rutowski 2016).

Mimicry (Back to Elevation)

Female spicebush and black swallowtails have blue on the dorsal surface of the hind wings and announced to be more convincing mimics than males when viewed dorsally. Male spicebush swallowtails accept light green and those of black swallowtails have a lot of yellow and very little blue. Nevertheless, males of these species have orange spots on the ventral surface of the hind wings and are probably mimetic when the wings are closed. Nether experimental conditions, male and female person black swallowtails were as protected when their ventral sides were presented to captive blue jays, simply males were eaten more often than females when their dorsal sides were presented (Codella and Lederhouse 1989).

Müllerian mimicry: Müllerian mimicry is a type of mimicry in which two or more species are similar in advent and are mutually distasteful or dangerous. Therefore, each species acts simultaneously as a model and a mimic and gains protection from the resemblance. Scott (1986, p.73) suggested that Battus philenor larvae and certain millipedes may be Müllerian mimics. Some millipedes are nighttime colored like Battus philenor larvae and release hydrogen cyanide when threatened (Eisner 2003, Eisner et al. 2005).

Acknowledgements (Back to Peak)

The author would like to admit Howard Frank and Marc Minno for reviewing this article and offering helpful suggestions. Also, the assistance of Pam Drach and Laura Lamb, members of the southwestern affiliate of the Indiana Native Institute Order, in providing leafage of woolly pipevine (Aristolochia tomentosa) for a Battus philenor caterpillar I was rearing during a trip to Indiana is greatly appreciated.

Selected References (Back to Top)

• Ahles H. 1959. Aristolochia serpentaria var. nashii as a new proper noun for A. serpentaria var. hastata. Journal of the Elisha Mitchell Science Society 75: 130.
  
• Allard RA. 2002. Aristolochia serpentaria L. Virginia Serpent Root. Conservation and Inquiry Plan for New England. New England Wild Flower Gild. Framingham, Massachusetts. 21 pp. (23 March 2020)
• Allard RA, Papaj DR. 1996. Learning of leaf shape by pipevine swallowtail butterflies: A test using artificial leaf models. Journal of Insect Behavior ix: 961-967.
• Allen TJ. 1997. The Butterflies of West Virginia and Their Caterpillars. University of Pittsburgh Press. Pittsburgh, Pennsylvania. 388 pp.
• Arnaud PH. 1978. A host-parasite itemize of N American Tachinidae (Diptera). United States Department of Agriculture Miscellaneous Publication 1319. Washington, D.C.
  
• Arnett RH Jr. 2000. American Insects: A Handbook of the Insects of America North of Mexico. CRC Press. 2nd Ed. Boca Raton, Florida. 1003 pp.
  
• Austin DF. 2004. Florida Ethnobotany. CRC Press. Boca Raton, Florida. 909 pp.
• Barbier R, Chauvin Chiliad. 1976. 50'enveloppe vitelline chez les Lépidoptères: Transformation lors de l'activation de 50'ovocyte et liaison avec la cuticule sérosale. Bulletin de la Société Zoologique de France 101: 1084-1085.
• Berenbaum MR, Moreno B, Dark-green E. 1992. Soldier bug predation on swallowtail caterpillars (Lepidoptera: Papilionidae): Circumvention of defensive chemistry. Journal of Insect Beliefs 5: 547-553.
• Blatchley WS. 1896. Miscellaneous notes. Canadian Entomologist 28: 265-266.
• Borror DJ. 1960. Lexicon of Word Roots and Combining Forms. Mayfield Publishing Visitor. Palo Alto, California. 134 pp.
• Brock JP. Kaufman G. 2003. Butterflies of North America. Houghton Mifflin. New York, New York. 383 pp.
• Brower JV. 1958. Experimental studies of mimicry in some Northward American collywobbles. ii. Battus philenor and Papilio troilus, P. polyxenes and P. glaucus. Development 12: 123-136.
• Brower LP, Brower JVZ. 1962. The relative abundance of model and mimic butterflies in natural populations of the Battus philenor mimicry complex. Ecology 43: 154-158.
• Burgess KS, Singfield J, Melendez V, Kevan PG. 2004. Pollination biology of Aristolochia grandiflora (Aristolochiaceae) in Veracruz, Mexico. Register of the Missouri Botanical Garden 91: 346-356.
• Cech R, Tudor G. 2005. Butterflies of the East Coast. Princeton University Printing. Princeton, New Jersey. 345 pp.
• Chen Z, Zhu D. 1987. Aristolochia alkaloids. pp. 29-65. In: Brossi A. (ed.). The Alkaloids: Chemistry and Pharmacology. Vol. 31. Academic Press. New York.
• Codella SG Jr., Lederhouse RC. 1989. Intersexual comparison of mimetic protection in the black swallowtail butterfly, Papilio polyxenes: Experiments with captive blue jay predators. Evolution 43: 410-420.
• Comstock JA, Grimshawe FM. 1935. Early on stages of Papilio polydamas lucayus R. & J. Message of the Southern California Academy of Sciences 34: 76-80.
• Crosswhite FS, Crosswhite CD. 1985. The southwestern pipevine (Aristolochia watsonii) in relation to snakeroot oil, swallowtail butterflies, and ceratopogonid flies. Desert Plants 6: 203-207.
• Culley TM, Klooster MR. 2007. The cleistogamous convenance system: A review of its frequency, evolution, and ecology in angiosperms. The Botanical Review, 73:1-30.
• Dimarco RD, Fordyce JA. 2013. Larger clutches of chemically defended butterflies reduce egg mortality: Testify from Battus philenor. Ecological Entomology 38: 535-538.
• Dimarco RD, Nice CC, Fordyce JA. 2012. Family matters: Effect of host plant variation in chemical and mechanical defenses on a sequestering specialist herbivore. Oecologia 170: 687-693.
• Doucet SM, Meadows MG. 2009. Iridescence: A functional perspective. Journal of the Royal Society Interface (Interface Focus Supplement) half-dozen: S115-S132.
• Knuckles JA. 2001. CRC Handbook of Medicinal Herbs. CRC Press. Boca Raton, Florida. 677 pp.
• Eisner T. 2003. Affiliate 2. Vinegaroons and other wizards. pp. 44-73. In: For Honey of Insects. The Belknap Press of Harvard Academy Press. Cambridge, Massachusetts. 448 pp.
• Eisner T, Eisner M, Siegler Thou. 2005a. Chapter nine. Class Diplopoda, Order Polydesmida, Family Polydesmidae, Apheloria kleinpeteri, a polydesmid millipede. pp. 43-47. In: Secret Weapons: Defenses of Insects, Spiders, Scorpions, and Other Many-legged Creatures. Harvard Academy Press. Cambridge, Massachusetts. 372 pp.
• Eisner T, Eisner M, Siegler M. 2005b. Chapter 64. Class Insecta, Order Lepidoptera, Family Papilionidae, Eurytides marcellus, the zebra swallowtail butterfly. pp. 297-303. In: Secret Weapons: Defenses of Insects, Spiders, Scorpions, and Other Many-legged Creatures. Harvard University Press. Cambridge, Massachusetts. 372 pp.
• Eisner T, Kluge AF, Ikeda MI, Meinwald YC, Meinwald J. 1971. Sesquiterpenes in the osmeterial secretion of a papilionid butterfly, Battus polydamas. Journal of Insect Physiology 17: 245-250.
• Eisner T, Meinwald YC. 1965. Defensive secretion of a caterpillar (Papilio). Science 150: 1733-1735.
• Feeny P. 1991. Chemical constraints on the evolution of swallowtail butterflies. pp. 315-340. In: Price Pw, Lewinsohn TM, Fernandez GW, Benson WW. (eds.). Plant-Animal Interactions: Evolutionary Ecology in Tropical and Temperate Regions. John Wiley & Sons. New York. 639 pp.
• Field WD. 1938. A manual of the collywobbles and skippers of Kansas (Lepidoptera: Rhopalocera). Bulletin of the University of Kansas: Biological Serial 39: 199.
• Flora of Northward America. Undated. Vol. 3. [http://world wide web.efloras.org/volume_page.aspx?volume_id=1003]) (23 December 2016)
• Glassberg J, Minno MC, Calhoun JV. 2000. Collywobbles Through Binoculars: A Field, Finding, and Gardening Guide to Butterflies in Florida. Oxford University Press. New York. 242 pp.
• Hall DW, Brown BV. 1993. Pollination of Aristolochia littoralis (Aristolochiales: Aristolochiaceae) by males of Megaselia spp. (Diptera: Phoridae). Annals of the Entomological Society of America 86: 609-613.
• Hazel WN. 1995. The causes and evolution of phenotypic plasticity in pupal color in swallowtail butterflies. pp. 205-210. In: Scriber JM, Tsubake Y, Lederhouse RC. (eds). Swallowtail Butterflies: Their Ecology and Evolutionary Biology. Scientific Publishers, Inc. Gainesville, Florida. 459 pp.
• Hazel WN, West DA. 1979. Environmental control of pupal color in swallowtail butterflies (Lepidoptera: Papilionidae) Battus philenor (Fifty.) and Papilio polyxenes Fabr. Ecological Entomology 4: 393-400.
• Hazel WN, Due west DA. 1983. The effect of larval photoperiod on pupal color and diapause in swallowtail butterflies. Ecological Entomology eight: 37-42.
• Heinrich Thousand, Chan J, Wanke Due south, Neinhuis C, Simmonds JSJ. 2009. Local uses of Aristolochia species and content of nephrotoxic aristolochic acid 1 and 2 - a global assessment based on bibliographic sources. Periodical of Ethnopharmacology 125: 108-144.
• Heitzman JR, Heitzman JE. 1987. Butterflies and Moths of Missouri. Missouri Department of Conservation. Jefferson, Missouri. 385 pp.
• Howe WH. 1975. The Butterflies of N America. Doubleday. Garden City, NJ. 633 pp.
• Howe WH. 1988. Larval predation of the blue swallowtail, Battus philenor, in Kansas. News of the Lepidopterists' Lodge No. 4, p.61.
• Iftner D, Shuey JA, Calhoun JV. 1992. Butterflies and Skippers of Ohio. Higher of Biological Sciences. The Ohio State University. Columbus, Ohio. 212 pp.
• Kandori I, Tsuchihara Thou, Suzuki TA, Yokoi T, Papaj DR. July 29, 2015. Long frontal projections help Battus philenor (Lepidoptera: Papilionidae) larvae observe host plants. PLOS ONE 10: e0131596.
• Kendall RO. 1964. Larval foodplants for 20-six species of Rhopalocera (Papilionoidea) from Texas. Journal of the Lepidopterists' Society xviii: 129-157.
• Layberry RA, Hall Pw, Lafontaine JD. 1998. The Butterflies of Canada. University of Toronto Printing. Toronto, Canada. 280 pp.
• Lederhouse RC. 1995. Comparative mating behavior and sexual selection in N American swallowtail butterflies. pp. 117-131. In: Scriber JM, Tsubaki Y, Lederhouse RC. eds. Swallowtail Butterflies: Their Environmental and Evolutionary Biology. Scientific Publishers, Inc. Gainesville, Florida. 459 pp.
• Linnaeus C. 1771. Regni animalis, Appendix. Insecta, pp. 529-543. In: Mantissa plantarum altera generum editionis Vi & specierum editionis Ii. Laurentius Salvius. Stockholm, Sweden. pp. 143-588. [description of Papilio philenor, p. 535] (3 February 2017)
• Miller JS. 1987. Phylogenetic studies in the Papilioninae (Lepidoptera: Papilionidae). Message of the American Museum of Natural History 186: 365-512.
• Minno MC, Butler JF, Hall DW. 2005. Florida Butterfly Caterpillars and their Host Plants. University Press of Florida. Gainesville, Florida. 341 pp.
• Minno MC, Minno M. 1999. Florida Butterfly Gardening. Academy Press of Florida. Gainesville, Florida. 210 pp.
• Moerman DE. 1998. Native American Ethnobotany. Timber Press, Portland, Oregon. 927 pp.
• Mondal, P. 2016. Pollination in plants: Types, advantages and disadvantages. (23 March 2020)
• Nechaev VA, Nakonechnaya OV. 2009. Construction of fruits and seeds and ways of dissemination of two species of the genus Aristolochia L. in Primorsky Krai. Biology Message 36: 393-396.
• Nice RW, Fordyce JA. 2006. How caterpillars avoid overheating: Behavioral and phenotypic plasticity of pipevine swallowtail larvae. Oecologia 146: 541-548.
• Nielsen ME, Papaj DR. 2015. Effects of developmental change in body size on ectotherm body temperature and behavioral thermoregulation: Caterpillars in a heat-stressed surround. Oecologia 177: 171-179.
• Odendaal FJ, Rausher Medico, Benrey B, Nunez-Farfan J. 1987. Predation by Anolis lizards on Battus philenor raises questions nearly butterfly mimicry systems. Journal of the Lepidopterists' Society 41: 141-143.
• Ômura H, Honda K, Feeny P. 2006. From terpenoids to aliphatic acids: Further show for late-instar switch in osmeterial defense force as a characteristic trait of swallowtail butterflies in the tribe Papilonini. Journal of Chemic Ecology 32: 1999-2012.
• Opler PA, Krizek Go. 1984. Butterflies Due east of the Peachy Plains. The Johns Hopkins University Press. Baltimore, MD. 294 pp.
• Opler PA, Malikul V. 1998. A Field Guide to Eastern Butterflies. Peterson Field Guides. Houghton Mifflin Company. New York. 486 pp.
• Osorio R. 2010. Ant dispersal in Aristolochia serpentaria (Virginia snakeroot). (23 March 2020)
• Otis GW, Locke B, McKenzie NG, Cheung D, MacLeod Due east, Devil-may-care P, Kwoon A. 2006. Local enhancement in mud-puddling swallowtail butterflies (Battus philenor and Papilio glaucus). Journal of Insect Behavior 19: 685-696.
• Papaj DR. 1986a. An oviposition mistake by Battus philenor L. (Papilionidae). Journal of the Lepidopterists' Lodge 40: 348-349.
• Papaj DR. 1986b. Leaf buds, a cistron in host selection by Battus philenor butterflies. Ecological Entomology 11: 301-307.
• Papaj DR, Newsom GM. 2005. A within-species alarm function for an aposematic signal. Proceedings of the Purple Guild B – Biological Sciences 272: 2519-2523.
• Pegram KV, Han HA, Rutowski RL. 2015. Warning signal efficacy: Assessing the furnishings of colour, iridescence, and time of day in the field. Ethology 121: 861-873.
• Pegram KV, Lillo MJ, Rutowski RL. 2013. Iridescent blue and orange components contribute to the recognition of a multicomponent warning point. Behaviour 150: 321-336.
• Pegram KV, Rutowski RL. 2014. Relative effectiveness of bluish and orange warning colours in the contexts of innate avoidance, learning and generalization. Animal Behaviour 92: 1-eight.
• Pegram KV, Rutowski RL. 2016. Effects of directionality, bespeak intensity, and short-wavelength components on iridescent alert indicate efficacy. Behavioral Ecology and Sociobiology 70: 1331-1343.
• Pelham JP. 2008. A catalogue of the butterflies of the United States and Canada. Lepidoptera Research Foundation. Beverly Hills, California. 658 pp.
• Pfeifer HW. 1966. Revision of the North and Key American hexandrous species of Aristolochia (Aristolochiaceae). Register of the Missouri Botanical Garden 53: 115-196.
• Pinto CF, Troncoso AJ, Urzúa A, Niemeyer Chiliad. 2009b. Employ of volatiles of Aristolochia chilensis (Aristolochiaceae) in host searching by 4th-instar larvae and adults of Battus polydamas archidamas (Lepidoptera: Papilionidae: Troidini). European Journal of Entomology 106: 63-68.
• Poulton EB. 1909. Mimicry in collywobbles of North America. Annals of the Entomological Society of America 11: 203-242.
• Proctor 1000, Yeo P, Lack A. 1996. The Natural History of Pollination. Timber Press. Portland, Oregon. 479 pp.
• Racheli T, Oliverio M. 1993. Biogeographical patterns of the neotropical genus Battus Scopoli 1777 (Lepidoptera: Papilionidae). Tropical Zoology half-dozen: 55-65.
• Rausher Doctor. 1978. Search epitome for leafage shape in a butterfly. Science 200: 1071-1073.
• Rausher MD. 1979. Egg recognition: Its advantage to a butterfly. Animal Behaviour 27: 1034-1040.
• Rausher MD, Feeny P. 1980. Herbivory, plant density, and plant reproductive success: The event of Battus philenor on Aristolochia reticulata. Ecology 61: 905-917.
• Richards OW, Davies RG. 1977. Chap. 18 Embryology. In: Vol. 1. Imms' Textbook of Entomology. 10th edition. John Wiley & Sons. New York. 418 pp.
• Robinson GS, Ackery PR, Kitching IJ, Beccaloni GW, Hernández LM. Undated. HOSTS - a Database of the Earth'due south Lepidopteran Hostplants. (23 March 2020)
• Rutowski RL, Rajyaguru PK. 2013. Male-specific iridescent coloration in the pipevine swallowtail butterfly (Battus philenor) is used in mate selection by females merely non sexual discrimination by males. Journal of Insect Beliefs 26: 200-211.
• Rutowski RL, Nahm AC, Republic of macedonia JM. 2010. Iridescent hindwing patches in the pipevine swallowtail: Differences in dorsal and ventral surfaces relate to betoken function and context. Functional Ecology 24: 767-775.
• Sachdev-Gupta K, Feeny PP, Carter Grand. 1993. Oviposition stimulants for the pipevine swallowtail butterfly, Battus philenor (Papilionidae), from an Aristolochia host constitute: synergism between inositols, aristolochic acids and a monogalactosyl diglyceride. Chemoecology 4: xix-28.
• Schaneberg, BT, Applequist WL, Khan IA. 2002. Determination of aristolochic acid I and II in North American species of Asarum and Aristolochia. Pharmazie 57: 686-689.
• Schull EM. 1987. The Collywobbles of Indiana. Indiana Academy of Science (Distributed past Indiana University Press). Bloomington, Indiana. 262 pp.
• Scopoli, 1000.A. 1777. Introductio advertisement historiam naturalem sistens genera lapidum, plantarum, et animalium: hactenus detecta, caracteribus essentialibus donata, in tribus divisa, subinde ad leges naturae. Apud Wolfgangum Gerle. Prague. p. 433. (23 March 2020)
• Scott JA. 1986. The Butterflies of North America. Stanford University Printing. Stanford, California. 583 pp.
• Sime Thou. 2000. Two new records of pimpline ichneumonids attacking Battus philenor (Linnaeus) (Lepidoptera: Papilionidae). Journal of Hymenoptera Research nine: 210-212.
• Sime K. 2002. Chemical defense force of Battus philenor larvae against set on past the parasitoid Trogus pennator. Ecological Entomology 27: 337-345.
• Sime Chiliad, Feeny PP, Haribal MM. 2000. Sequestration of aristolochic acids by the pipevine swallowtail, Battus philenor (50.): Show and ecological implications. Chemoecology ten: 169-178.
• Sims SR, Shapiro AM. 1983a. Pupal colour dimorphism in California Battus philenor (L.) (Papilionidae): Mortality factors and selective advantage. Periodical of the Lepidopterists' Society 37: 236-243.
• Sims SR, Shapiro AM. 1983b. Pupal color dimorphism in California Battus philenor: Pupation sites ecology command, and diapause linkage. Ecological Entomology 8: 95-104.
• Smith JE. 1797. Natural History of the Rarer Lepidopterous Insects of Georgia. Vol. one. Bensley, London. 100 pp. (From the observations of John Abbot)
• Sourakov A, Daniels JC. 2002. Is Battus philenor hirsuta a subspecies? News of the Lepidopterists' Society 44: 64-65.
• Stamp NE. 1986. Concrete constraints of defence force and response to invertebrate predators by pipevine caterpillars (Battus philenor: Papilionidae). Periodical of the Lepidopterists' Society 40: 191-205.
• Stehr FW. 1987. Immature Insects. Kendall/Chase. Dubuque, Iowa. 754 pp.
• Tyler HA. 1975. The Swallowtail Butterflies of North America. Naturegraph Publishers. Healdsburg, CA. 192 pp.
• Tyler HA, Brownish KS Jr., Wilson KH. 1994. Swallowtail Butterflies of the Americas. Scientific Publishers. Gainesville, Florida. 376 pp.
• Underwood DLA. 1994. Methods for sexing Lepidoptera larvae using external morphology. Journal of the Lepidopterists' Society 48: 258-263. (includes Battus philenor)
• Wagner DL. 2005. Caterpillars of Eastern Northward America. Princeton University Printing. Princeton, New Jersey. 512 pp.
• Westward DA. 1983. Hand pairing of Battus philenor (Papilionidae). Journal of the Lepidopterists' Lodge 37: xc.
• West DA, Hazel WN. 1979. Natural pupation sites of swallowtail butterflies (Lepidoptera: Papilioninae): Papilio polyxenes Fabr., P. glaucus L. and Battus philenor (L.). Ecological Entomology iv: 387-392.
• Weintraub JD. 1995. Host plant association patterns and phylogeny in the tribe Troidini (Lepidoptera: Papilionidae). pp. 307-316. In: Scriber JM, Tsubake Y, Lederhouse RC. eds. Swallowtail Butterflies: Their Ecology and Evolutionary Biology. Scientific Publishers, Inc. Gainesville, Florida. 459 pp.
• Zomlefer WB. 1994. Plant Families. The University of N Carolina Printing. Chapel Colina, North Carolina. 430 pp.

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