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Gulf Coast Pitcher Plant Bogs
Gulf Coast Pitcher Plant Bogs
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here saturation results from a seasonally high water table. Pitcher plant bogs may also be found
on stream terraces or at the edges of sinkholes.
In all sites movement of water through the substrate is slow. Flooding seldom occurs, but
scattered small pools are often present during the wet season. The soils are typically sands, loamy
sands, or sandy loams. In recent soil surveys they are mapped mainly as members of the Atmore,
Myatt, Pansey, Plummer, and Rains series, although some bogs do occur on soils of other series.
The soils are typically strongly acidic, ranging from pH 3.5 to pH 5.0. The acidity results from
the activity of mineral components of the soil, with organic acids contributing little. Although the
pitcher plant bogs are low-energy wetlands in which flowing water moves very little material, the
amount of organic litter that accumulates is small because of frequent fires. Input of nutrients
from outside the bogs is minimal in most cases. Nutrient cycling is therefore dependent on the
organisms
Except for the insect species associated with pitcher plants of the genus Sarracenia, the fauna of
the bogs is poorly known. Means and Moler (1978) noted that small pools in the bogs are an
important habitat for the larvae of the endangered pine barrens tree frog, Hyla andersonii. The
odd flightless grasshopper, Gymnoscirtetes morsei (Acrididae), appears to be restricted to this
habitat (M. E. Dakin, pers. comm.), as is the spittlebug, Lepyronia angulifera (Cercopidae) (J. B.
Chapin, pers. comm.). Several species of burrowing crayfish of the genera Cambarus and
Procambarus are often abundant and may play a role in bringing leached nutrients to the surface.
Ants and earthworms are sometimes common but seem to be less so in bogs where the normal
cycles of moisture and fire occur. Bamforth (1976) sampled fungi, bacteria, and protozoa in
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southeastern Louisiana and found populations generally lower in soil surrounding the roots of
plants in bog sites than in other habitats. The general absence of surface litter results in a sparse
litter fauna in the bogs as contrasted with areas that do not burn as frequently.
Carnivorous plants
Nowhere on the continent and perhaps nowhere in the world can one find a diversity of
carnivorous plants equaling that in the pitcher plant bogs. Well over half of the approximately
forty-five North American carnivorous species occur along the Gulf Coast, with as many as
thirteen species in four genera found in a single bog. Even extensively disturbed sites often
harbor four or five species.
The works of Darwin (1899) and Lloyd (1942) include information on some of the carnivorous
species inhabiting the bogs, and more recently Schnell (1976) has described the North American
forms in a popular work. However, the details of the evolution and ecology of the carnivorous
species remain unclear. In the bogs, the carnivorous flora include species of sundews (Drosera:
Droseraceae), bladderworts (Litricularia: Lentibulariaceae), butterworts (Pinguicula:
Lentibulariaceae), and pitcher plants (Sarracenia: Sarraceniaceae).
Tolerance of both fire and soil saturation characterizes the carnivorous species, but the exact role
that carnivory plays in their ability to survive in the bogs is not fully understood. It has long been
hypothesized (Darwin 1899; Lloyd 1942) that absorption of prey products supplements nutrients
in the soil, allowing carnivorous species to compete in nutrient-poor habitats. Plummer (1963)
found that soils in carnivorous plant habitats in Georgia were low in calcium, potassium, and
magnesium, but not exceptionally low in phosphorus. He conjectured that the plants benefit more
from
mineral ions than from nitrogenous compounds derived from their prey. Eleutarius and Jones
(1969) examined bog soils in Mississippi and found no deficiencies in nitrogen, phosphorus, or
potassium. When they applied 6-12-12 fertilizer and ammonium nitrate, productivity in
Sarracenia alata actually declined. Christensen (1976) fed insects to S. flava and found no
indication of a resulting increase in concentrations of calcium, magnesium, or potassium in leaf
tissue, although levels of nitrogen and phosphorus were significantly higher than those of
controls. He concluded that carnivory might enhance plant nutrition on soils deficient in nitrogen
and phosphorus.
Little consideration has been given to the possible role that the availability of micronutrients may
have played in the evolution of carnivory. Soil scientists have long known that the availability of
molybdenum, a trace element essential for plant metabolism, is very low at low pH values such
as typically exist in the bogs (Jones 1957). With our present state of knowledge it could he
contended that supplementation of the molybdenum supply is a major benefit resulting from the
phenomenon of carnivory. A host of other possibilities remain. Bell (pers. comm.) and
Christensen (1976) have pointed out that the breakdown of prey detritus from decaying pitchers
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The southeastern pitcher plants produce leaves the pitcherseach year from perennial
underground rhizomes or rootstocks. The pitchers, which function as pitfall traps, are tubular,
possess downward-pointing hairs on the interior surface, and secrete a fluid containing digestive
enzymes. They capture a variety of insects and other small animals, which are attracted to nectar
secreted by glands near the pitcher orifice. Reproduction is typically by seeds produced from
flowers borne in the spring, hut may also occur by fragmentation of the rhizomes.
Resource partitioning allows coexistence among species that occur together. Hypothetically,
carnivorous plants sharing the same habitat should demonstrate prey partitioning similar to that
found among animal carnivores, and there is evidence of this among Sarracenia species. Fish
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(1976) has noted that S. minor in Florida seems to capture mainly ants, T. C. Gibson (pers.
comm.) has found evidence of prey partitioning among several Sarracenia species he is studying
in the Florida panhandle. My own preliminary studies indicate that S. purpurea obtains a
spectrum of prey species quite different from that of other pitcher plants, often capturing
significant numbers of grasshoppers, crickets, and snails. Microhabitat segregation exists among
species in the same bog and may influence the types of prey obtained. For instance, along the
Gulf Coast S. leucophylla occupies wetter sites than S. flazna, and S. purpurea is often restricted
to the edges of bogs (McDaniel 1971).
Further complicating the assessment of ecological and evolutionary interrelationships among the
Sarracenia species is the presence of hybrids at many sites (Figs. 2 and 3). Almost all the
possible natural hybrids have been found, and occasionally large hybrid swarms are encountered
(Bell 1952; Bell and Case 1956; McDaniel 1971). All possible hybrids have been produced in the
greenhouse, where they seem to be as vigorous and fertile as the parental types (Schnell and
Krider 1976).
The possibility of introgressionthe exchange of alleles between two distinct specieshas been
raised, but there is little evidence that significant genetic exchange is taking place among species
sharing the same habitat. The isolating mechanisms which prevent or reduce genetic exchange
among the species have not been intensively studied. It appears that ecological differences in the
sites preferred by various species are not alone sufficient to prevent hybridization. As many as
five species may inhabit the same bog, and although the ranges of flowering times differ, the
species flower simultaneously in some years (McDaniel 1971). Moreover, in the absence of
burningi.e., when competition is more severemost species flower later. This may result in
overlap in flowering times on burned and unburned sites adjacent to each other.
Pollination of Sarracenuia has been described by MacFarlane (1908) and Jones (1908) and
commented on by Schnell (1978). Bumblebees, the main pollinators, are polytropic, visiting
many plant species. However, during the peak of the Sarracenia flowering season the bees are
effectively monotropic, at least at sites where there are large stands of flowers, visiting only
Sarracenia. It may be significant that Sarracenia species which tend to flower simultaneously
differ in the color, height (length of peduncle), or size of their flowers. For instance, both S. flava
and S. purpurea flower early, but S. flava has yellow flowers, whereas those of S. purpurea are
pinkish- purple and have somewhat shorter peduncles. S. rubra and S. leucophylla flower
simultaneously, and their flowers are essentially the same color. However, S. rubra has small
flowers on short peduncles, whereas S. leucophylla possesses larger flowers on much longer
pedun cles. S. flava and S. alata have flowers of similar color, height, and size, but S. flava
flowers earlier and there is only a small area in which the two species occur together.
purpurea are short, jug-shaped, partially reclining, possess an erect hood, and fill readily with
rainwater. S. alata pitchers, on the other hand, are tall, spindly, erect, taper gradually to a narrow
base, and possess an ascending hood that restricts the entry of water. The hybrid has an open
orifice and is erect, but is characterized by a weak base. When excessive water enters during
heavy rains the pitcher topples, draining its contents and often remaining prostrate. I have found
that hybrids often possess abnormally large amounts of the plant pigment anthocyanin distributed
in unusual patterns. One could speculate that this is indicative of metabolic anomalies that render
hybrids less physiologically fit than parental types.
Hybrids seem to require more water than parental types and are more susceptible to water stress.
During eight years of observations in Santa Rosa County, Florida, I have found that rhizomes of
hybrids of S. flava and S. leucopinylla produce fewer leaves than parental types, and in
exceptionally dry years may not produce leaves at all. McDaniel (1971) hypothesized that hybrids
cannot compete with parental types and eventually disappear. My observations support this
contention and indicate that hybrids are also less successful in competing with other plants that
invade the bog habitat, especially in the absence of fire.
Grant (1971) showed that genetically viable hybrids may be at a disadvantage because of poorer
pollination. This factor may reduce backcrossing among the Sarracenia speciesthat is, crossing
of a first generation hybrid with a parent. For instance, the hybrid of S. flava and S. minor flowers
at a time when neither parental species is in prime flower and when pollinators are probably
concentrating on other plants. Flower color may also be a factor. Hybrids in which the parents
have yellow and purple flowers respectively produce flowers of an intermediate color that seem
to be less attractive to bumblebees than those of the parental types.
Pitcher plant insects
The interior of the pitcher leaves of Sarracenia is a unique microhabitat unparalleled elsewhere in
nature. Compared to the surrounding environment, the cavity within a pitcher is typically higher
in relative humidity, lower in light intensity, and somewhat less variable in temperature. In
addition, pitchers contain a decomposing mass of entrapped prey which is a potential food source
for other organisms. Although the adaptations that allow pitcher plants to entrap, detain, and
digest prey are highly successful, their presence has presented a counterevolutionary challenge to
organisms that might benefit from an ability to turn the tables on the pitcher plant and colonize
the pitcher environment.
A number of species have evolved the ability to inhabit the pitchers without being entrapped or
digested. Among these, the most fully studied is a mosquito, Wyeomia smithii, whose immature
stages occur in the fluid held on the leaves of S. purpurea. Unlike most insects, these larvae are
neither killed nor digested in the pitcher fluid (Goins 1977 diss.; Bradshaw 1980; Moeur and
Istock 1980). The larvae of four species of sarcophagid flies of the genus Blaesoxiplnia also
inhabit Sarnce,uia pitchers, where they feed on entrapped insects (Forsyth and Robertson 1975).
The larvae of three other fliesMetriocnemus knahi (Chironomidae) (Paterson 1971; Cameron et
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al. 1977); Dolurnipluora cornuta (Phoridae) (Jones 1918); and Bradysia macfarlanei (Sciaridae)
(Jones 1920)are still other occupants of the pitchers.
All three species of the noctuid moth genus Exyra occur in Sarracema pitchers (Fig. 4), where
their larvae feed on the leaf tissue (Jones 1921; Judd 1957; Brower and Brower 1970; Rymal
1980 diss.). Two other moths, Papaipenna appasionata (Noctuidae) and Endothenia daeckeana
(Tortricidae), feed exclusively on pitcher plant tissues but do not enter the pitchers themselves.
By contrast, the aphid, Macrosiphum jeanae, completes its entire life cycle within the pitchers
(Robinson 1972).
Two anoetid mites, Anoetus gibsom and A. hughesi, are known only from Sarracena pitchers
(Nesbitt 1954; Hunter and Hunter 1964), as is the predaceous phyto seiid mite, Macroseius
biscutatas (Chant et al. 1959). At least fourteen of these arthropods are obligate associates of
various Sarracenia species. The mechanisms by which these species escape entrapment and
digestion merit more intensive study.
Natural habitat maintenance
The heliophytes of the bog habitat are dependent on natural phenomena that continually retard the
processes of succession which would eventually eliminate them. First, soil acidity coupled with
low levels of nutrients, at least at some sites, inhibits the invasion of competing species. Second,
anaerobic soil conditions resulting from frequent saturation create an environment inhospitable to
moisture-intolerant types. Third, periodic fires repeatedly eliminate fire-intolerant types. Most of
the bog species have underground rhizomes or rootstocks and hence are not harmed by fire.
Fire is undoubtedly the most important of these
factors. This is demonstrated by the fact that regardless of the moisture regime and soil
conditions, the absence of fire inevitably results in eventual elimination of the bog species. The
paramount role of fire in maintaining these habitats has been documented by a number of workers
(Wells and Shunk 1928; Eleutarius and Jones 1969). The bogs therefore represent a fire
subclimax or disclimax, and they depend on fire not only to eliminate competitors but to release
many nutrients bound up in organic matter as a result of previous growth (Eleutarius and Jones
1969). It should be pointed out that nitrogen is largely volatilized by burning and does not
typically increase in soils after fire.
The season during which fire occurs may influence the floristic composition of the bogs. In the
past, fires apparently occurred most frequently in the summer as a result of lightning (Komarek
1965). At present, most fires are caused by man and occur during the winter. Data on the effects
of this shift are lacking; however, winter fires would seem less effective in opening space for the
germination of seeds of the bog species.
Penfound (1952) and others have conjectured that pitcher plant habitats are at times created by
the cutting of longleaf pine stands. In some cases, reduced evapotranspiration following cutting
may increase soil moisture to the extent that the site becomes suitable for bog species. However,
the creation of drainage ditches and other preparations for replanting produce conditions which
prevent bog communities from developing in most logged sites. The idea that bogs are formed
mainly through cutting of the low pine forests (Penfound 1952, p. 431) is untenable, considering
the vast expanses they occupied before significant disturbance by man. It is possible that
destruction of forests by hurricanes may have played a part in maintaining coastal bogs.
Before mans activities began to affect the pitcher plant bogs, these communities were abundant
and conspicuous on the Lower Gulf Coastal Plain. Based on extensive on-site surveys to
determine habitat suitability and detailed examination of county soil maps, I have calculated that
in pre-Columbian times these communities occupied approximately 2,935 km2, or about 6% of
the colored area in Figure 5. Nearer the coast the bogs were probably larger and more
conspicuous. From the writings of Bartram (1791) and Harper (1918) it seems probable that areas
near the coast from Pensacola, Florida, west to Pascagoula, Mississippi, were nearly continuous
bogs until the late 1800s, with some bogs covering thousands of acres.
The area occupied at present is significantly smaller. Bogs in a natural or nearly natural condition
are now very rare, with perhaps less than 12 km2 remaining. Altered sites which retain the aspect
of bogs and contain many of the component species occur on an additional 60 km2. Even these
liberal estimates mean that at least
97% of the former bogs have been destroyed or seriously altered. The reasons for the
disappearance of these communities are myriad, and I have summarized many of them previously
(1977). The most damaging factors have been draining of land and restriction of fire.
The frequency of fires in pitcher plant bogs is much lower now than in pre-Columbian times and
has greatly decreased in the past few decades. Not only are there intensive efforts to prevent and
control fires, but roads, railroads, tilled cropland, pastures, and fire lanes serve as barriers to
restrict tile spread of fire. A decline in the frequency of fire can be damaging not only because
nutrients remain inaccessible but because of tile extreme heat generated by the eventual
combustion of large amounts of accumulated vegetation, since such heat can affect perennial
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regime if changes in nearby areas threaten to disturb the hydrologic characteristics of the bog.
These requirements could perhaps be met in a national park or preserve consisting of some
10,000 acres distributed in four or five blocks across the Southeast. Similar national parks have
long ago been established to preserve other unique ecosystems such as the Everglades or the cove
forests of the Great Smoky Mountains, or to protect other precious species such as the sequoias
and the saguaros.
Many important questions about the evolution and ecology of pitcher plant communities and
about the biology of carnivorous species remain to be answered. Yet we cannot wait for more
detailed scientific information to justify preservation, for if we do, the communities will be gone
before they can be preserved. The disappearance of the pitcher plant bogs would be a biological
catastrophe of great magnitude, closing the door forever on further study and enjoyment of an
irreplaceable member of earths varied ecosystems.
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