Monarch Butterfly Ecology
Karen S. Oberhauser
Department of Fisheries, Wildlife and Conservation Biology
University of Minnesota,
St. Paul, Minnesota, USA
Michelle J. Solensky
Department of Biology,
The
College of Wooster
Wooster, Ohio, USA
Note: This online review is updated and revised continuously, as soon as
results of new scientific research become available. It therefore
presents state-of-the-art information on the topic it covers.
Monarch butterflies (Danaus plexippus)
are native to North and South America. In the 1800's, however, they
spread to other parts of the world. Monarchs were first seen on Hawaii in
the 1840's, and then throughout many South Pacific islands in the 1850's and
1860's (Ackery and Vane-Wright 1984). In the early 1870's, the first
monarchs were reported in Australia and New Zealand (Gibbs 1994). It is not
clear exactly how and why this emigration occurred. One possibility is that
monarchs were transported by ships, either as larvae that found their way
onboard from shipyard milkweed plants, or as adult monarchs that happened to
land on ocean-going vessels. It is most likely that humans were involved in
the process, but it is not known to what extent. Because North American
monarchs often fly over 2,200 km during their migration, it is always
possible that some made the journey on their own (Vane-Wright 1993).
In the western hemisphere, there are two subspecies
of the monarch butterfly: (1)
Danaus plexippus plexippus of southern Canada, the USA, Mexico, most
Caribbean Islands, Central America and northern South America; and (2)
Danaus plexippus erippus of Peru, Bolivia, Chile, Argentina,
Uruguay, Paraguay and eastern Brazil. Neither subspecies has been recorded
in northwestern Brazil.
The foci of this review are the North
American migratory populations, i.e. those monarchs that migrate to overwintering
grounds in central Mexico and the west coast of the USA. The review is
6 pages long and is divided into five parts:
Monarch Breeding Ecology
Fall Migration Ecology
Monarch Overwintering Ecology
Spring Migration Ecology
References and Contact
Information
The first section, Monarch Breeding
Ecology, is two pages long and begins below. The other sections are
each one page long.
Monarch Breeding Ecology
Overview: Monarch larvae are
specialist herbivores, consuming only host plants in the milkweed family (Asclepiadacea).
They utilize most of the over 100 North American species (Woodson 1954) in
this family, breeding over a broad geographical and temporal range that
covers much of the United States and southern Canada. In a typical year, one
generation is produced in the southern part of this range by returning
migrants, and two to three generations are produced in the northern part.
Monarchs and Milkweed. Milkweed
provides monarchs with an effective chemical defense against many predators.
Monarchs sequester cardenolides (also called cardiac glycosides) present in
milkweed (Brower and Moffit 1974), rendering them poisonous to most
vertebrates. However, many invertebrate predators, as well as some bacteria
and viruses, may be unharmed by the toxins or able to overcome them. The
extent to which milkweed protects monarchs from non-vertebrate predators is
not completely understood, but a recent finding that wasps are less likely
to prey on monarchs consuming milkweed with high levels of cardenolides
suggests that this defense is at least somewhat effective against
invertebrate predators (Rayor 2004).
Benefits gained by monarchs from cardenolides are not without cost. Milkweed plants between and even within
species vary a great deal in cardenolide concentration, and both the toxin
and the sticky latex produced by the plants provide defenses against
herbivores. Monarchs appear to be negatively affected by consuming plants
with high cardenolide levels, and may actually starve to death when their
mandibles are glued together by the latex or if their bodies become mired in
a drop of latex formed when the plant is injured (Zalucki and Brower 1992;
Malcolm and Zalucki 1996; Zalucki and Malcolm 1999; Zalucki et al. 2001).
Larger larvae reduce this risk by chewing a notch at the base of the
milkweed leaf midvein, cutting off the flow of sticky latex to the rest of
the leaf and allowing more efficient eating (see
Figure 1).
Like other plants, milkweed quality as a
host for insects varies. Many insects are nitrogen limited (McNeil and
Southwood 1978, Mattson 1980, Scriber 1984, Slansky and Scriber 1985, White
1993). They must consume large quantities of their host plants to accumulate
enough nitrogen for growth and development, since animal tissue generally
consists of 7-14% nitrogen by dry weight (dw) and plants consist of
0.03-7.0% nitrogen dw (Mattson 1980). Leaf nitrogen levels vary within a
season, as plant tissue ages and as plants allocate more resources to
reproductive tissue. In addition, plants grow in habitats with different
levels of available soil nitrogen. Lavoie and Oberhauser (2004) studied the
response of monarch larvae to plants manipulated through fertilizer
treatments to contain varying leaf nitrogen levels, and found that they
compensated for low nitrogen leaves by consuming more plant tissue per day.
If increased consumption makes them more vulnerable to predation or plant
defenses, this could result in decreased fitness levels.
The most important northern host plant is
Asclepias syriaca (common milkweed - Figure 2),
but a number of other species are used as well, including A. incarnata
(swamp milkweed - Figure 3) and A. tuberosa
(butterfly weed - Figure 4). Central Plains host
plants include the vine Cynanchum laeve (sand or honey vine). A
northeastern invasive plant in the same genus (C. nigrum) has spread
west as far as Wisconsin. This species is attractive to ovipositing females,
but monarch larvae do not survive on it (Haribal 1998). In the south, the
most important host plants are probably Asclepias oenotheroides (zizotes
milkweed),
A. viridis (spider milkweed) and A. asperula (antelope horn
milkweed), all fairly common throughout Texas and other southern US states.
Egg production. It is difficult to tell just how many eggs female
butterflies lay during their lives, but the average in the wild is probably
300 to 400. Captive monarch butterflies average about 700 eggs per female
over 2 to 5 weeks of egg laying, with a record of 1179 eggs (Oberhauser
1997). Monarch eggs hatch about 4 days after they are laid, but the rate of
development in this stage, like all other stages, is temperature dependent,
with individuals in warmer environments developing more rapidly (Zalucki
1982). The proteins that are an important constituent of eggs must either be
derived from nutrients ingested during the larval stage or obtained from
males during mating (Boggs and Gilbert 1979, Oberhauser 1997). While an
individual monarch egg (Figure 5) weighs only
about 0.460 mg, about 1/1000 the adult mass, females often lay more than
their own mass in eggs throughout their lives.
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