(DRAFT) - Taxonomy
Species BUTTERFLY, SWALLOWTAIL, SCHAUS
Species Id ESIS504001
Date 13 MAR 96
TAXONOMY
NAME - BUTTERFLY, SWALLOWTAIL, SCHAUS
OTHER COMMON NAMES - BUTTERFLY, SWALLOWTAIL, SCHAUS; SWALLOWTAIL, SCHAUS;SWALLOWTAIL, SCHAUS'; SWALLOWTAIL and KEYS
ELEMENT CODE -
CATEGORY - Terrestrial Insects
PHYLUM AND SUBPHYLUM - ARTHROPODA,
CLASS AND SUBCLASS - INSECTA,
ORDER AND SUBORDER - LEPIDOPTERA,
FAMILY AND SUBFAMILY - PAPILIONIDAE,
GENUS AND SUBGENUS - HERACLIDES,
SPECIES AND SSP - ARISTODEMUS, PONCEANUS
SCIENTIFIC NAME - HERACLIDES ARISTODEMUS PONCEANUS
AUTHORITY -
TAXONOMY REFERENCES -
COMMENTS ON TAXONOMY -
Schaus Swallowtail Butterfly
Heraclides aristodemus ponceanus Small, 1911
KINGDOM: Animal GROUP: Insect
PHYLUM: Arthropoda CLASS: Insecta
ORDER: Lepidoptera FAMILY: Papilionidae
The Schaus swallowtail, Heraclides aristodemus ponceanus Schaus
(or more completely: Heraclides (=Papilio) aristodemus Esper 1794,
ssp. ponceanus Schaus 1911) is a large blackish-brown swallowtail
butterfly with contrasting markings mostly dull yellow (01,02,03).
Antenna black with a yellow knob, except for black tip. Forewing with
dull yellow median band from apex to about midpoint of inner margin,
with short side branch to costa about 1/3 distance from apex.
Subterminal and terminal lines of lunular yellow spots from apex to
anal angle. Hindwing with yellow median band continuing that of
forewing, and submarginal row of large yellow lunular spots; yellow
edging in concavities of deeply scalloped outermargin. Blackish
"tail" is straight-edged (not teardrop-shaped), and is bordered with
yellow. Hollow red spot along anal margin just above anal angle;
bluish scaling basad of it. Small, inconspicuous red dash sometimes
present basad of the second yellow lunule from the anal angle (between
Taxonomy - 1 (DRAFT) - Taxonomy
Species BUTTERFLY, SWALLOWTAIL, SCHAUS
Species Id ESIS504001
Date 13 MAR 96
vein M2 and Cu1).
Underside of wings yellow with black shading mostly in the median
and submarginal areas of forewing and in terminal area and tails of
the hindwing. Dull brownish red median band extends from costa to
inner margin of hindwing, narrowing before touching these margins.
Extensive bluish scaling along outer edge of reddish band. Wingspan:
2 7/8 to 4 inches (8.6 to 9.5 cm) (01,02,29).
This swallowtail is most easily confused with the giant
swallowtail, Heraclides (=Papilio) cresphontes Cramer, which is
widespread in eastern North America and which also occurs in habitat
occupied by the Schaus swallowtail. The two are easily separated in
that the giant swallowtail is larger and more nearly coal black with
lines brighter yellow. Its median forewing band is broader but more
broken into spots, and is less separated from the submarginal band
toward the apex. Its antenna is solid black and its tail is
teardrop-shaped, yellow inside bordered with black edging. Its
reddish underside markings are less brownish and much less extensive
than in the Schaus swallowtail (03).
In its original description (07) the Schaus swallowtail was
considered to be a full species, Papilio ponceana, with the type
locality Miami, FL. Schaus noted that it was "allied to P.
aristodemus from Haiti, and P. temenes Godt. from Cuba." It was first
placed as a subspecies of aristodemus Esper in 1917 (08), and has been
cited as such since that time except for Holland's revised version of
The Butterfly Book (09). Now considered an Antillean species with
high propensity to vary geographically, P. aristodemus consists of 5
subspecies (10): Nominate P. aristodemus (Hispaniola), P. aristodemus
temenes Godart (Cuba), P. a. ponceanus Schaus (Florida), P. a.
driophilus Clench (Bahamas), and P. a. bjorndalae Clench (Bahamas).
Validity of some of these subspecies has been questioned (15).
However, it is believed that the Schaus swallowtail may represent a
distinct species (30).
Orthography of the specific name was "ponceana" in the original
description (07) and in some earlier references (08,11,12).
Widespread use of the "ponceanus" spelling resulted from its first
major appearance in The Butterfly Book (09). Both subspecies status
and the "ponceanus" orthography have been stable since 1951 (01).
Generic usage has fluctuated recently, with North American
species traditionally placed in Papilio Linnaeus grouped into 3
genera, one of which is Heraclides Hubner (13). This usage has been
followed by some recent workers (02,23), but not by others (03,06,14,
15). These conservative workers retain Papilio, considering
Heraclides to be at most a subgenus. Heraclides (Papilio) aristodemus
ponceanus has been used incorrectly since Papilio is not a subgenus.
Heraclides (=Papilio) aristodemus ponceanus is the correct
representation.
Common names for this species include Schaus swallowtail
butterfly, Schaus (or Schaus') swallowtail, and Keys swallowtail.
Holotype male and allotype female of ponceanus Schaus are in the
Dept. of Entomology, U.S. National Museum of Natural History,
Washington, DC 20560 (15). Holotypes female and paratypes of
driophilus Clench and bjorndalae Clench are in the Carnegie Museum of
Natural History, Pittsburgh, PA 15213. The locations of 246 specimens
Taxonomy - 2 (DRAFT) - Taxonomy
Species BUTTERFLY, SWALLOWTAIL, SCHAUS
Species Id ESIS504001
Date 13 MAR 96
of ponceanus are tabulated in (15). It is not known if the types of
aristodemus Esper and temenes Godart still exist.
Descriptions of the Schaus swallowtail are to be found in 01, 02,
03, 06, 07, 09, 10, and 11. Color illustrations in 01, 03, 04, 05,
06, 09, and 14. A black-and-white plate comparing the Schaus and
giant swallowtails, with diagnostic arrows, appears in reference 16.
Taxonomy - 3 (DRAFT) - Status
Species BUTTERFLY, SWALLOWTAIL, SCHAUS
Species Id ESIS504001
Date 13 MAR 96
STATUS
Coded Status
Florida; Federal Endangered
Florida; Officially Listed
E: Federal Endangered
Game (Consumptive Recreational)
Ornamental
COMMENTS ON STATUS -
U.S. STATUSES AND LAWS:
The Schaus swallowtail butterfly (Heraclides aristodemus
ponceanus) has been designated an Endangered species pursuant to the
Endangered Species Act of 1973 (50 CFR 17.11; P.L. 93-205, 87 Stat.
884; 16 U.S.C. 1531-1540), as amended. The subspecies has this status
wherever found including the State of Florida.
This subspecies is protected by the Lacey Act (P.L. 97-79, as
amended; 16 U.S.C. 3371 et seq.) which makes it unlawful to import,
export, transport, sell, receive, acquire, or purchase any wild
animal (alive or dead including parts, products, eggs, or offspring):
(1) in interstate or foreign commerce if taken, possessed,
transported or sold in violation of any State law or
regulation; or
(2) if taken or possessed in violation of any U.S. law,
treaty, or regulation or in violation of Indian tribal law.
It is also unlawful to possess any wild animal (alive or dead
including parts, products, eggs, and offspring) within the U.S.
territorial or special maritime jurisdiction (as defined in
18 U.S.C. 7) that is taken, possessed, transported, or sold in
violation of any State law or regulation, foreign law, or Indian
tribal law.
RESPONSIBLE FEDERAL AGENCIES:
USFWS -Responsible for the management/recovery, listing, and
law enforcement/protection of this species.
NPS -Responsible for the law enforcement/protection of this
species with applicable State and Federal laws on
public lands under their control. Also responsible
for conservation (Nat. Park System Organic Act - 16
U.S.C. 1, 2-3)/management/recovery on National Park
Service lands. Taking, possessing, or disturbing of
Federally listed species is prohibited on NPS lands
(36 CFR 2.1, 2.2, and 2.3).
All Federal agencies have responsibility to ensure that any
action authorized, funded, or carried out by that agency is not likely
to jeopardize the continued existence of the species or result in the
destruction or adverse modification of Critical Habitat (50 CFR 402),
and to utilize their authorities to carry out programs for the
Status - 1 (DRAFT) - Status
Species BUTTERFLY, SWALLOWTAIL, SCHAUS
Species Id ESIS504001
Date 13 MAR 96
conservation of the species.
STATE STATUSES AND LAWS:
STATE: Florida
DESIGNATED STATUS: Endangered
ADMINISTRATIVE AGENCY: Florida Game and Fresh Water Fish
Commission and Florida Department of
Natural Resources.
STATE STATUTE: Wildlife Code of the State of Florida, Title 39,
Florida Administrative Code.
INTERNATIONAL STATUSES, TREATIES, AND AGREEMENTS:
The Schaus swallowtail butterfly (Papilio aristodemus ponceanus)
is listed as Endangered in the IUCN Red Data Book (1983).
ECONOMIC STATUSES:
The species is a beautiful butterfly that has cultural value to
man. It has also been popular with butterfly collectors and as such
has ornamental value. The species may have had some commercial value
but the extent of this is unknown.
75/04/22:40 FR 17757/17758 - Proposal to list as Threatened
76/04/28:41 FR 17736/17740 - Final rule, listed as Threatened
81/02/27:46 FR 14652/14658 - Five year review
83/08/29:48 FR 39096/39099 - Proposal to list as Endangered
84/08/31:49 FR 34501/34504 - Final rule, listed as Endangered
Status - 2 HABITAT ASSOCIATIONS
HABITAT - TERRESTRIAL
TERRESTRIAL
COASTAL
LAND USE -
Transportation, communications, and Util
Evergreen Forest Land
COMMENTS ON HABITAT ASSOCIATIONS -
The habitat of the Schaus swallowtail butterfly is exclusively
the mature subtropical dry forest (hardwood hammocks), now extensive
only in the upper Florida Keys in Dade and Monroe Counties (15).
Adults may fly in clearings and along roads and trails, or even out
over the ocean for short distances from island to island (19,22); but
they typically remain in the hammocks proper (19). Nectaring activity
usually occurs on blossoms of wild-coffee (Psychotria nervosa), guava
(Psidium guajava), cheese shrub (Morinda royoc), in or at the margins
of these hammocks, rarely in open sunlit places (15,19).
Oviposition occurs on foodplants within the hammock or at its
margin, not in open spaces subjected to full sunlight (19). The
primary food is foliage of torchwood, Amyris elemifera L., and a
secondary one is wild lime, Zanthoxylum fagara L., both of which are
in the rue family (Rutaceae). The Schaus swallowtail is rarely very
far from hammock habitat in which one or both of these plants are
plentiful (15,19).
Other characteristics of the habitat include relatively high
elevation (3 to 4.6 m above sea level), away from tidal waters, and a
mature overstory of characteristic trees such as the foodplants, gumbo
limbo (Bursera simaruba), pigeon plum (Coccoloba diversifolia), black
ironwood (Krugiodendron ferreum), mahogany (Swietenia mahagoni), and
wild tamarind (Lysiloma latisiliquum) (16). These plants grow on a
substrate of Key Largo limestone, which characterizes the upper keys.
The lower keys (Big Pine Key to Key West) have Miami Oolite overlying
the Key Largo Limestone (16). The lower keys are characterized by a
more open, slash pine/silver palm vegetation, unsuitable for Schaus '
swallowtail habitat (14).
Climatic factors include mean annual temperatures ranging from 23
degrees C (74 degrees F) in the Miami area to 25 degrees C
(77 degrees F) in the middle and lower keys. Annual rainfall in the
Miami area is 1524-1651 mm, with that in the upper keys 1143-1270 mm,
and that in the southern keys 889-1016 mm. South Florida rainfall
patterns are seasonal, with a November-May dry season and a bimodal
wet season with peaks in June and September-October. Long-term
periodicity is evident, with particularly wet periods occurring in
apparently 5-year cycles (16).
Hurricane factors are also discussed in the Recovery Plan (16):
"Tropical cyclones (winds greater than 61 kph) and hurricanes (winds
greater than 119 kph) occur on an average of once every 5 to 7 years
in southern Florida. The probability of a 'great hurricane'
(sustained winds greater than 210 kph) striking the area in a given
year ranges from 2% to 7%. Hurricane storm surges of from 3.0-5.5 m
above mean sea level have been recorded in southern Florida, heights
capable of submerging any portion of the keys. Such surges and their
associated high winds can severely damage coastal hammocks, but such
effects may be ameliorated somewhat by hammock canopy shape, which
tends to deflect winds, and by large quantities of accompanying rain,
which flush sea water from the hammock soil."
The most detailed ecological summary for Schaus swallowtail
Habitat Associations - 1 butterfly habitat can be found in the 1984 status report (14), the
Recovery Plan (16), and a 1986 status report (30).
Habitat Associations - 2 (DRAFT) - Food Habits
Species BUTTERFLY, SWALLOWTAIL, SCHAUS
Species Id ESIS504001
Date 13 MAR 96
FOOD HABITS
TROPHIC LEVEL -
HERBIVORE
LIFESTAGE FOOD FOOD PART
General Evergreen Shrubs-Leaves/Twigs
General Coniferae
Food Habits - 1 (DRAFT) - Environment Associations
Species BUTTERFLY, SWALLOWTAIL, SCHAUS
Species Id ESIS504001
Date 13 MAR 96
ENVIRONMENTAL ASSOCIATIONS
G = General A = Adult
LIM = Limiting RA = Resting Adult
J = Juvenile FA = Feeding Adult
RJ = Resting Juvenile BA = Breeding Adult
FJ = Feeding Juvenile P = Pupae
L = Larvae E = Egg
RL = Resting Larvae
FL = Feeding Larvae
LIFESTAGE ENVIRONMENTAL ASSOCIATIONS
G
G Coastal Features: Vegetated offshore islands
Environment Associations - 1 (DRAFT) - Life History
Species BUTTERFLY, SWALLOWTAIL, SCHAUS
Species Id ESIS504001
Date 13 MAR 96
LIFE HISTORY
FOOD HABITS:
The Schaus swallowtail butterfly's food habits can be divided
into larval food habits and adult food habits.
Females of the Schaus swallowtail butterfly have been observed
ovipositing on leaves of only two plant species. These are torchwood,
Amyris elemifera L. (17,19,22), and wild lime, Zanthoxylum fagara L.
(19,23). Both belong to the citrus family (Rutaceae). In all cases,
eggs were laid on the upper surface of the tender new leaves of the
foodplant. Usually a single egg is laid per leaf; but occasionally
there may be two (15). This species has been reared (from an egg) in
captivity on another rutaceous plant, prickly-ash (Zanthoxylum
americanum Mill.). The egg was recovered from a leaf of wild lime in
nature (19).
Young caterpillars apparently utilize only tender, young leaves
of the foodplants, avoiding tougher older leaves (19). Fifth (final)
instar larvae have been observed eating tougher older leaves of
torchwood in nature (15), and also prickly-ash in the laboratory (19).
Adults have been observed taking nectar from blossoms of guava
(Psidium guajava L., Myrtaceae) (19), cheese shrub (Morinda royoc L.,
Rubiaceae) (15,19), and wild coffee (Psychotria nervosa Sw.,
Rubiaceae) (15). Guava seemed to be the nectar source preferred by
individuals observed by Rutkowski, and he suggested that the
butterflies will fly some distance from their hammock haunts to find
blooming guava flowers (19). Emmel (30) observed frequent nectaring
at 7 plant species on Elliott Key. These nectar sources include:
cheese shrub (Morinda royoc), blue porterweed (Stachylarpheta
jamaicensis), sea grape (Cocoloba uvifera), Dog's tail (Heliotropium
angiospermum), wild-sage (Lantana involucrata), salt-and-pepper
(Melanthera nivea), and wild-coffee (Psychotria nervosa).
HOME RANGE/TERRITORY:
The Schaus swallowtail butterfly is territorial to the extent
that males have been observed to investigate other males entering
their territories within the hardwood hammock. "They alight
frequently on torchwood leaves about 3-4 feet off the ground and
"bask" with outspread wings in the sunfleck for a few minutes; . . .
they investigate other ponceanus that enter their territory and fly
closely (but slowly) around them for a few moments . . . " (14).
Males have also been reported as they "patrolled the tree tops at a
height of 10 feet or more" during the "hot afternoon" on "bright
days", sometimes "descending into open spaces to investigate any other
ponceanus" (19). Emmel also notes that they are remarkably adapted to
flight within the hammock, and are able to pick their way among
branches and spider webs. However, individuals show wing damage very
soon after emergence. Some of this is caused by striking various
obstacles; other damage such as beak marks must result from bird
attacks (14).
PERIODICITY:
Adult Schaus swallowtails fly primarily in May and June, with
most sightings having been recorded between mid-April and mid-July
Life History - 1 (DRAFT) - Life History
Species BUTTERFLY, SWALLOWTAIL, SCHAUS
Species Id ESIS504001
Date 13 MAR 96
(15). A few August and September records suggest either delayed
emergence during a year, or a facultative second brood (15,24). A
unique record of a worn female in March 31, 1961, taken on Key Largo
remains a puzzling anomaly (25).
While this butterfly is apparently univoltine, there is evidence
from rearing that diapause may extend to at least 2 years (17). If
such is the case in nature, the butterfly could survive extreme
drought in the season following that of its larval development by
waiting another year, or perhaps until July-September or later, before
emerging as an adult (19).
As far as is known, the Schaus swallowtail is strictly diurnal.
Rutkowski observed 2 females on different days visiting cheese shrub
blossoms just before 9 am (EDT), his earliest observation; another
female hovering over cheese shrub at 5 pm (EDT) comprised his latest
observation in a day (19). He found both sexes "within the hammocks,
fluttering in diffused light about a foot above the ground at blossoms
of Guava . . ." during the hottest part of the day (from 1 to 2 pm
(EDT)).
MIGRATION PATTERNS:
While no mass migration of the Schaus swallowtail butterfly has
ever been reported, an individual butterfly was followed as it crossed
a half-mile expanse of Biscayne Bay between two islands (22). This
suggests that this species can disperse across open water for a
considerable distance among the upper Florida Keys, and possibly to
and from the nearby mainland. Another such observation was made in
1986, when a Schaus swallowtail crossed 1200 feet from Old Rhodes to
Swan Key (30).
COVER/SHELTER REQUIREMENTS:
All evidence points to the absolute necessity for mature hardwood
hammock habitat with dappled sunlight penetration as requisite for
continued survival of both the Schaus swallowtail butterfly and for
its primary foodplant, torchwood (Amyris elemifera L.) (14,15,16,19,
22,23). The minimum area required is not known.
REPRODUCTIVE SITE REQUIREMENTS:
Actual mating was observed in 1985 by a student of T.C. Emmel,
but not yet published (20). Courtship and oviposition behavior have
been reported for the Schaus swallowtail butterfly. Dense, mature
subtropical hardwood hammock habitat on well-drained substrate is
essential for survival of this species, but the amount needed is not
known. Density of primary and secondary foodplants is also not known.
The butterflies spend much of their time within hammocks, particularly
where sunlight penetrates to give a dappling effect (14). Adult
nectaring may occur there, or out in the open (19). Courtship has
been observed along narrow trails cut through the hammock (19,26).
Open areas such as trails or clearings within or near the dense
hammock are requisite for courtship activity and nectaring. These
open areas may be natural or man-made.
REPRODUCTIVE CHARACTERISTICS:
Courtship has been partially described (19): "At 10:15 AM in a
Life History - 2 (DRAFT) - Life History
Species BUTTERFLY, SWALLOWTAIL, SCHAUS
Species Id ESIS504001
Date 13 MAR 96
dimly-lit trail through brushy hammock I saw a female and two male
ponceanus visiting flowers at opposite ends of a Guava tangle. The
fresher of the two males eventually approached this slightly worn
female while she was still feeding. He hovered over her. She then
settled on the ground with wings flattened and vibrating, raising her
abdomen. The male fluttered on the ground behind and then rose over
her before flying away." Three courting pairs of Schaus swallowtails
were observed in 1982 on Elliott Key (26) and photographed, details
were not recorded. The males were flying behind the females during
part of the flight, which was very low to the ground (1-3 m). In the
pair photographed, the male was worn and heavily damaged, but the
female seemed fresh and whole.
While mating has not yet been reported, oviposition in nature has
been described. Foodplants have been torchwood (Amyris elemifera L.,
Rutaceae) (17,19,22) and wild lime (Zanthoxylum fagara L., Rutaceae)
(19,23). In all cases, females deposited single eggs on the upper
surface of the tips of the leaves. In cases of eggs found in nature,
there is one record of 2 eggs on a leaf (15). Foodplants on which
females oviposit are either at the edge of hammocks along trails,
partially sheltered by the canopy; or they are in the hammocks proper,
perhaps at the edge of a clearing or where a fairly large opening in
the canopy exists. Eggs and larvae found in nature are not on
foodplants in open sunlight; however, in contrast, the giant
swallowtail, Heraclides cresphontes, has been observed ovipositing on
wild lime growing in the open (19).
Development from egg to adult was described by Grimshawe (17) and
Rutkowski (19). Eggs take 3-5 days to hatch. First instar larvae
are "black-brown, except for the thirteenth segment, which is white.
Also, a minute faint light speck is visible on each side of the
seventh ring. This grows more distinct daily, becoming a clear,
round, white spot at the first molt" (17). Grimshawe continues the
description of the remaining 4 instars and pupation:
"The first molt takes place upon the fourth or fifth day.
Immediately after the molting the larva as a rule eats its cast-off
skin, and adorns itself with a white hood.
"With the second molt blue or metallic dots appear, with the
early blackish-brown color transforming into a lighter shade. The
white on the anal extremity extends forward into the twelfth segment.
"The third molt brings other white markings of irregular shape
along each side of the body, while the white spot on the seventh ring
increases in size and projects back into the eighth ring. This
development is fascinating to watch.
"At the age of twenty days the caterpillars go into their fourth
and last molt, having gone through molts at intervals of approximately
five-day periods. At this mature stage the white markings have
changed to a creamish-yellow, except those on the anal segments, the
hood, and the markings on the third ring. However, the band or hood
of the second segment is capped with the creamish-yellow.
"Spots of robin-egg blue appear like jewels upon a soft maroon
background. The true legs are blackish-brown; the prolegs are shaded
white; the anal prolegs are a uniform white. Teasing the caterpillar
brings a white, bifurcated or pronged organ out of its nape-ring.
This exudes a strong odor.
Life History - 3 (DRAFT) - Life History
Species BUTTERFLY, SWALLOWTAIL, SCHAUS
Species Id ESIS504001
Date 13 MAR 96
"The mature larvae, at normal rest, measure fifty-six
millimeters. These caterpillars are solitary and enjoy no community
life as do the larvae of the P. polydamus.
"When ready to transform, the larva seeks a place of seclusion,
each for itself, and fastens its anal extremity with a button of silk,
and throws a heavy girdle around the thorax, supporting the body in an
upright, or vertical, position for the long sleep.
"The encased segments of the body of some of the chrysalides are
rusty-brown color; others are gray, etched with moss-green and
conforming generally with that of their supporting object. They take
on a rigid cast and show no signs of life throughout the entire period
of their sleep. Unlike the double and triple-brooded related species
of neotropical Florida, our butterflies remained in the chrysalis
stage either one or two years. As an example, half the caterpillars
transforming into the chrysalis stage July 7, 1935, emerged May 8,
1936; the other half remained, hatching May 13, 1937" (17).
Rutkowski also noted the white osmeteria on the larvae, and
drinking of water droplets by fifth-instar caterpillars (19). While
he observed oviposition on torchwood and wild lime in nature, he
successfully reared one caterpillar to maturity on prickly-ash
(Zanthoxylum americanum) after his supply of acceptable torchwood and
wild lime leaves gave out during the second instar (19). The life
history stages have been described, illustrated, and compared with the
giant swallowtail (30).
PARENTAL CARE:
The Schaus swallowtail female oviposits only on the tips of young
leaves of the foodplants, torchwood (Amyris elemifera L.) and wild
lime (Zanthoxylum fagara L.), and usually deposits one egg per leaf
(15,17,19,22,23). Foodplants in dappled sunlight within hardwood
hammocks or at their edges (with some canopy over them) are used
exclusively (19,20). Selection of proper foodplant, oviposition
singly on new leaves, and preference for protected sites constitute
all known behaviors that might come under the category of parental
care.
POPULATION BIOLOGY:
Limiting factors include:
1) Food Supply. Within the major keys of Biscayne National Park
(Elliott, Old Rhodes, Totten, and Adams Keys) and on northern Key
Largo, the two foodplants of the Schaus swallowtail butterfly seem
adequate to support a healthy population. High numbers of individuals
sighted in 1985 (20) indicate that the butterfly's population is still
capable of periodic peaks;
2) Cover. As now managed, Biscayne National Park provides
adequate cover for both Schaus swallowtail adults and also food plants
(14,15). This cover includes mature and well-drained tropical
hardwood hammock with some natural and man-made openings such as
narrow trails and clearings where nectaring and courting behavior can
take place close to the more enclosed jungle-like forest where adults
spend much of their time (15);
3) Predation. Little is known about predation by spiders,
lizards, birds, or other predators. Damage to wings occurs soon after
Life History - 4 (DRAFT) - Life History
Species BUTTERFLY, SWALLOWTAIL, SCHAUS
Species Id ESIS504001
Date 13 MAR 96
adult emergence, and beak marks on some individuals indicate frequent
bird attacks (14). Flight behavior among the many obstacles in
hammock habitat seems unusually deliberate, in that the butterflies
can fly slowly and painstakingly to avoid the many large orb spider
webs and branches to a remarkable degree (14). Emmel also states that
butterflies are able to remember flight paths through hammocks and
follow them repeatedly. A tendency toward Batesian Mimicry, with the
Zebra butterfly (Heliconius charitonius L., Nymphalidae) as model has
also been observed and discussed (14,19,30). Covell notes that larval
predation is surely minimized by oviposition behavior (one egg per
leaf and few per foodplant), bird-dropping appearance of the larvae
(as in other Papilio larvae), secretive behavior of larvae, and
bad-smelling scents from the osmeteria when larvae are disturbed (17,
19). Crypsis in the pupa (17) as in other swallowtails is also a
factor in avoiding predation. Nothing is known about parasites of
this species;
4) Disease. No information is available regarding diseases of
the Schaus swallowtail. However, high egg mortality has been observed
(15,19);
5) Competition. No direct competition with other butterfly or
moth species for foodplant utilization has been observed. The giant
swallowtail, Heracildes cresphontes, does use wild lime, but oviposits
on plants in open sunlight and not on shaded plants as in the case of
the Schaus swallowtail (15,19);
6) Territorial Behavior. (SEE Home Range/Territory above); and
7) Natural Catastrophes. The main natural catastrophes which
might eradicate the Schaus swallowtail include (1) hurricanes; (2)
fires; and (3) freezes; and (4) droughts. These are discussed in
several works (14,15,16,17,18,10,23,27).
Survival/Mortality Rate: Information on survival rate of adults
is mostly anecdotal, although Emmel's 1985 study should provide some
useful data from mark-release-recapture studies when published (20).
Earlier projections are that adults live only about 2 weeks, and
suffer damage more quickly than similar species that live in more open
areas because of hazards of life in the dense brush of the hammock
(14).
Egg survival rate of 29.7% (n=11 of 37) was cited in one case for
a group of eggs collected in the wild. Further mortality of hatching
larvae resulted in a survival rate of only 5% in the group studied
(20).
Rate of Increase: Population fluctuations from year to year have
been discussed at length, with the butterfly abundant some years such
as 1935 (17), 1969-1972 (22,24), and 1985 (20); moderately common in
some others such as 1982 (26) and 1984 (14); and uncommon to rare in
yet others such as 1936 (17) 1973-1978 (14,15,16,27). The populations
general inclination, however, has been "in the direction of decline"
(14). In 1986, it was estimated that the adult Elliott Key population
was 750-1000 individuals, and that populations of adults and immatures
were 50-80 individuals on each of Old Rhodes, Totten, and Adams Keys
(30).
Sex Ratio: No studies on sex ratio have been published, but
Covell has found that males are seen in more abundance than females
(29). In tabulating 245 adult ponceanus in collections, 136 were
Life History - 5 (DRAFT) - Life History
Species BUTTERFLY, SWALLOWTAIL, SCHAUS
Species Id ESIS504001
Date 13 MAR 96
males, 41 females, and 68 had no sex indicated. If these figures were
indicative of natural sex ratios, males would outnumber females by
more than 4:1. However, Covell suspects something closer to 2:1,
males to females. Female butterflies are typically more secretive
than males, and in the case of Schaus swallowtails, a skewed
distribution might be explained in that conscientious lepidopterists
may tend to avoid killing females, heeding the plea to spare females
published by Klots in his popular Field Guide to the Butterflies (01):
"None but males should be collected, and then, at most , only one per
collector." A similar appeal is found in Kimball's book (04).
Recovery Potential: Positive opinions and suggestions regarding
recovery potential are found in the 1984 Status Report (14), the
Recovery Plan (15), and other works (16). Emmel (14) notes a good
nucleus for population recovery in Biscayne National Park.
Recommended recovery approaches include continued survey and
population monitoring, habitat preservation, rearing a captive
population for restocking and as a hedge against natural catastrophes,
range expansion through reintroduction, and further research including
evaluation of pesticide effects on the Hispanolian subspecies and more
detailed life history studies.
Turnover Rate and Longevity: The Schaus swallowtail is
considered to be univoltine (one generation per year), with short
adult longevity (14). There is some evidence of the ability to hold
over at least one year (17), and some that adults are active during
July-September as well as during the normal flight period of late
April through early July (24). Whether the latter condition indicates
partial, facultative bivoltinism or merely late annual emergence is
not known. One worn adult female has been collected in late March
(25), so flight period information is incomplete, but suggests some
flexibility that should assist in its survival.
Population Density Objective Under Management: An increase of
annual numbers and extension of its range to lessen the impact of
natural and man-made catastrophes which could result in extinction are
outlined in the 1984 Status Survey (14) and in the Recovery Plan (15).
SPECIES INTERRELATIONSHIPS:
The survival of the Schaus swallowtail buttefly depends on
survival of sufficient stands of its primary foodplant, torchwood
(Amyris elemifera L.) and, possibly, that of its secondary foodplant,
wild lime (Zanthoxylum fagara L.) in appropriate mature tropical
hardwood hammocks in the upper Florida Keys. These two rutaceous
species are the only known foods for this species in nature (14,15).
OTHER LIFE HISTORY DESCRIPTORS:
None.
Life History - 6 (DRAFT) - Management Practices
Species BUTTERFLY, SWALLOWTAIL, SCHAUS
Species Id ESIS504001
Date 13 MAR 96
MANAGEMENT PRACTICES
RESULT MANAGEMENT PRACTICE
Beneficial Maintaining undisturbed/undeveloped areas
Beneficial Land Acquisition
Beneficial Controlling/Restricting Pesticide Use
Beneficial Stocking captive-reared wild-strain animals
Beneficial Transplanting wild animals
Adverse Incidental Capturing/Killing
Existing Incidental Capturing/Killing
Adverse Poaching
Existing Poaching
Adverse Collecting
Existing Collecting
Adverse Commercial Exploitation
Existing Commercial Exploitation
Adverse
Existing
Adverse Food Supply Reduction
Existing Food Supply Reduction
Adverse Rural Residential/Industrial Areas
Existing Rural Residential/Industrial Areas
Adverse Highway/Railroads
Existing Highway/Railroads
Adverse Soil compaction by heavy equipment in mine areas
Existing Soil compaction by heavy equipment in mine areas
Adverse Shoreline modification/development
Existing Shoreline modification/development
Adverse Applying pesticides
Existing Applying pesticides
Adverse
Existing
Adverse Vegetation Composition Changes
Existing Vegetation Composition Changes
Adverse Fire
Existing Fire
Adverse Harvesting
Existing Harvesting
COMMENTS ON MANAGEMENT PRACTICES -
The Schaus swallowtail is restricted to a habitat where its
primary foodplant, torchwood (Amyris elemifera) grows abundantly (15).
This habitat is limited to coastal southeast Florida and the upper
Florida Keys, in mature tropical hardwood hammocks. Prior to human
influences, populations of this butterfly were probably subject to
naturally occurring population depressions caused by hurricane damage,
drought, and rare freezes (16). The influence of the Labor Day
hurricane of 1935 on the Lower Matecumbe Key population was documented
by Grimshawe (17), though she was incorrect in claiming that the
species became extinct (it was found there and on Key Largo in
succeeding years) (18). However, Grimshawe's careful searching was
Management Practices - 1 (DRAFT) - Management Practices
Species BUTTERFLY, SWALLOWTAIL, SCHAUS
Species Id ESIS504001
Date 13 MAR 96
negative, and her "before and after" experience demonstrates that this
hurricane had a detrimental effect on the biota of the keys southwest
of Key Largo.
There is no real evidence of effects of competition, disease, and
predation/parasitism, but they could have played a role in decreasing
either butterflies or foodplants or both. There is evidence of
indiscriminate spraying of pesticides for mosquito control (28) which
may be affecting the butterfly. The Schaus swallowtail may be very
vulnerable to these types of pesticides which are currently in use
(31).
Clearing of habitat for urban and agricultural purposes in and
around Miami, Homestead, and Lower Matecumbe Key certainly were
instrumental in eliminating the Schaus swallowtail from its type
locality in the extremes of its historical range. Foodplants were
probably either eliminated or reduced to small stands incapable of
sustaining Schaus swallowtail populations (15). Similar clearing has
occurred within its known north Key Largo habitat, but litigation has
slowed development of the area (16). Slight alteration of habitat,
such as dirt roads and trails through the hammocks, seem to be harmful
only in that they permit easy access to collectors, who can catch
butterflies when they fly low along these trails. However, small
clearings and trail edges seem to promote proliferation of torchwood
plants, which spring up in clearings and edges of clearings. Natural
succession in such places, particularly following hurricanes and
fires, could account for population increases in foodplants and the
butterfly (21), however, large fires and extensive forest clearing are
detrimental to the species. Female Schaus swallowtails tend to stay
in the partially shaded areas and oviposit on torchwood and wild lime
growing in these places, ignoring wild lime growing in the open (19).
Paved roads through Schaus swallowtail habitat, particularly
Rt. 905 on northern Key Largo, permit road kill of adults, one case of
which is documented (16). Aerial application of insecticides to
control biting Diptera may affect Schaus swallowtail populations on
Key Largo, but these effects are not known (15,16). The pesticides
Dibrom, Baytex, and Teknar, used in the Keys for mosquito control, are
toxic to the related giant swallowtail (Heraclides cresphontes) in the
laboratory (31).
Collecting of immature stages as well as adults may have reduced
numbers on Key Largo in the period 1969-1974; but again the lasting
effects cannot be gauged (16). Commercial exploitation has existed,
but its extent cannot be assessed due to secrecy on the part of
dealers. There was a case in which a naive individual offered adults
for sale by advertisment. The ad was withdrawn when the individual
was advised that the act was illegal. Also, one case exists of an
apparent intent to poach specimens in Biscayne National Monument in
1985 (29). The culprit was encountered with collecting equipment but
no permit on Elliott Key; however, no specimens were found (20).
The principal future threats to Schaus swallowtail survival are
the following, in descending order:
1. Extreme climatic conditions, especially hurricanes, freezes,
and droughts.
2. Habitat modification, especially developmental clearing and
Management Practices - 2 (DRAFT) - Management Practices
Species BUTTERFLY, SWALLOWTAIL, SCHAUS
Species Id ESIS504001
Date 13 MAR 96
fires.
3. Introduction of pesticides and other hazardous chemicals.
4. Road kills and death by predators, parasites and collectors.
APPROVED PLAN:
U.S. Fish and Wildlife Service. 1982. Recovery Plan for Schaus'
Swallowtail Butterfly (Heraclides (Papilio) aristodemus ponceanus
Schaus) Including Recommendations for Bahamian Swallowtail Butterfly
(Heraclides (Papilio) andraemon bonhotei Sharpe). U.S. Fish and
Wildlife Service, Atlanta, Georgia. 57 pp.
Actions recommended for the recovery of the Schaus swallowtail
butterfly include the following:
1) Maintain existing colonies by surveying for colonies and
acquiring land to protect hammocks from development.
2) Determine limiting factors such as habitat requirements, factors
affecting population dynamics and the effects of currently used
insecticides and their application.
3) Reestablish colonies in appropriate areas within its historic
range.
a. Establish and maintain a captive population.
b. Restock areas with wild or captive bred individuals.
4) Monitor reestablished colonies and habitat.
Management Practices - 3 (DRAFT) - References
Species BUTTERFLY, SWALLOWTAIL, SCHAUS
Species Id ESIS504001
Date 13 MAR 96
References
***** REFERENCES FOR ALL NARRATIVES EXCEPT N-OCCURRENCE *****
01 Klots, A.B. 1951. A field guide to the butterflies of North
America east of the Great Plains. Houghton Mifflin, Boston.
349 pp., 40 pls.
02 Pyle, R.M. 1981. The Audubon Society field guide to North
American butterflies. Chanticleer, New York. 916 pp., 759 pls.
03 Opler, P.A. and G.O. Krizek. 1984. Butterflies east of the Great
Plains. Johns Hopkins University Press, Baltimore. 294 pp.,
54 pls.
04 Kimball, C.P. 1965. Arthropods of Florida and neighboring land
areas. Vol. 1, Lepidoptera of Florida. Florida Dept. of
Agriculture, Div. of Plant Industry, Gainesville. 363 pp., 26 pls.
05 Miller L.D. 1975. Threatened status for two butterflies? Field
Museum Nat. Hist. Bull. 1975:2,15-18,22.
06 Howe, W.H. 1975. The butterflies of North America. Doubleday,
New York. 633 pp., 97 pls.
07 Schaus, W. 1911. A new Papilio from Florida, and one from Mexico
(Lepid.). Ent. News 22:438-439.
08 Barnes, W. and J.H. McDunnough. 1917. Check list of the
Lepidoptera of boreal America. Herald Press, Decatur, IL. 392 pp.
09 Holland, W.J. 1931. The butterfly book, rev. ed. Doubleday,
New York. 424 pp., 77 pls.
10 Clench, H.K. 1978. Papilio aristodemus (Papilionidae) in the
Bahamas. J. Lepid. Soc. 32(4):273-276.
11 Bates, M. 1934. Notes on some tropical Florida butterflies
(Lepid.:Rhopalocera). Ent. News 45:166-169.
12 McDunnough, J.H. 1938. Check list of the Lepidoptera of Canada
and the United States of America. Part 1, Macrolepidoptera. Mem.
S. Calif. Acad. Sci. 1. 272 pp.
13 Miller, L.D. and F.M. Brown 1981. A catalogue/checklist of the
butterflies of America north of Mexico. Mem. Lepid. Soc. 2.
280 pp.
14 Emmel, T.C. 1985. Status survey of the Schaus swallowtail in
Florida in 1984. Technical Report No. 145, Florida Cooperative
Fish and Wildlife Research Unit, University of Florida,
Gainesville. 39 pp.
15 U.S. Fish and Wildlife Service. 1982. Schaus Swallowtail
Butterfly Recovery Plan. U.S. Fish and Wildlife Service, Atlanta,
GA. 57 pp.
16 Covell, C.V., Jr. 1976. The Schaus swallowtail: a threatened
subspecies? Insect World Digest 3(5):21-26.
17 Grimshawe, F.M. 1940. Place of sorrow: The world's rarest
butterfly and Matecumbe Key. Nature Mag. 33:565-567, 611.
18 Henderson, W.F. 1945. Papilio aristodemus ponceana Schaus
(Lepidoptera:Papilionidae). Ent. News 56(2):126-136.
19 Rutkowski, F. 1971. Observations on Papilio aristodemus ponceanus
(Papilionidae). J. Lepid. Soc. 25(2):126-136.
20 Emmel, T.C. 1985. Personal communication. Department of Zoology,
University of Florida, Gainesville, FL 32611.
21 Baggett, H.D. 1985. Personal communication. Department of
References - 1 (DRAFT) - References
Species BUTTERFLY, SWALLOWTAIL, SCHAUS
Species Id ESIS504001
Date 13 MAR 96
Geology, University of South Florida, Tampa, FL 33620.
22 Brown, L.N. 1973. Populations of Papilio andraemon bonhotei
Sharpe and Papilio aristodemus ponceanus Schaus in Biscayne
National Monument, Florida. J. Lepid. Soc. 27(2):136-140.
23 Loftus, W.F. and J.A. Kushlan. 1984. Population fluctuations of
the Schaus swallowtail (Lepidoptera: Papilionidae) on the islands
of Biscayne Bay, Florida, with comments on the Bahaman swallowtail.
Florida Ent. 67(2):277-287.
24 Brown, C.H. 1976. A colony of Papilio aristodemus ponceanus
(Lepidoptera:Papilionidae) in the upper Florida Keys. J. Ga. Ent.
Soc. 11(2):117-118.
25 Covell, C.V., Jr. and G.W. Rawson. 1973. Project ponceanus: a
report of first efforts to survey and preserve the Schaus
swallowtail (Papilionidae) in southern Florida. J. Lepid. Soc.
27(3):206-210.
26 Covell, C.V., Jr. Unpublished. Butterfly count on Elliott Key,
Biscayne National Park, Florida, 18 May 1982.
27 Covell, C.V., Jr. 1977. Project ponceanus and the status of the
Schaus swallowtail (Papilio aristodemus ponceanus) in the Florida
Keys. Atala 5(1):4-6.
28 Tuskes, Paul M. 1985. Personal communication. 7900 Cambridge,
Houston, TX 77054.
29 Covell, C.V., Jr. 1985. Personal communication. Dept. of
Biology, University of Louisville, Louisville, KY 40292
30 Emmel, T.C. October 1986. Status survey and habitat requirements
of Florida's endemic Schaus swallowtail butterfly. Florida Game
and Fresh Water Fish Commission. Final Report GFC-84-028. 122 pp.
31 Emmel, T.C. October 1986. Pesticide effects on the survival of
the Schaus swallowtail butterfly. Final report to Elizabeth Ordway
Dunn Foundation, Inc. 57 pp.
***** REFERENCES FOR N-OCCURRENCE NARRATIVE ONLY *****
01 U.S. Fish and Wildlife Service. 1982. Schaus Swallowtail
Butterfly Recovery Plan. U.S. Fish and Wildlife Service, Atlanta,
GA. 57 pp.
02 Schaus, W. 1911. A new Papilio from Florida, and one from Mexico
(Lepid.). Ent. News 11:438-439.
03 Grimshawe, F.M. 1940. Place of sorrow: The world's rarest
butterfly and Matecumbe Key. Nature Mag. 33:565-567, 611.
04 Emmel, T.C. 1985. Status survey of the Schaus swallowtail in
Florida in 1984. Technical Report No. 145, Florida Cooperative
Fish and Wildlife Research Unit, University of Florida,
Gainesville. 39 pp.
05 Emmel, T.C. 1985. Personal communication. Dept. of Zoology,
University of Florida, Gainesville, FL 32611.
06 Covell, C.V., Jr. 1976. The Schaus swallowtail: A threatened
subspecies? Insect World Digest 3(5):21-26.
07 Brown, L.N. 1973. Populations of Papilio andraemon bonhotei
Sharpe and Papilio aristodemus ponceanus Schaus in Biscayne
National Monument, Florida. J. Lepid. Soc. 27(2):136-140.
08 Covell, C.V., Jr. and G.W. Rawson. 1973. Project ponceanus: A
References - 2 (DRAFT) - References
Species BUTTERFLY, SWALLOWTAIL, SCHAUS
Species Id ESIS504001
Date 13 MAR 96
report on first efforts to survey and preserve the Schaus
swallowtail (Papilionidae) in southern Florida. J. Lepid. Soc.
27(3):206-210.
09 Emmel, T.C. 1986. Status survey and habitat requirements of
Florida's endemic Schaus swallowtail butterfly. Florida Game and
Fresh Water Fish Commission. Nongame Report GFC-84-028. 122 pp.
References - 3