Carnivorous plants –

classic perspectives and new research

Barry Rice
The Nature Conservancy, Davis, USA

The ranks of known carnivorous plants have grown to approximately 600 species. We are
learning that the relationships between these feeders and their prey are more complex, and
perhaps gentler, than previously suspected. Unfortunately, these extraordinary life forms are
becoming extinct before we can even document them!

Carnivorous plants are able to do four things: they attract, false signals, the trigger hairs must be bent, not once, but
trap, digest and absorb animal life forms. While these four two or more times in rapid succession. In effect, the plant
abilities may seem remarkable in combination, they are, can count! When the trap first closes, the lobes fit together
individually, quite common in the plant kingdom. All very loosely, the marginal spines interweaving to form a
plants that produce flowers for the purpose of summoning botanical jail. Prey items that are too small to be worth
pollinators are already skilled at attracting animals. Many digesting can quickly escape, and the trap will reopen the
plants trap animals at least temporarily, usually for the next day. But, large prey remain trapped, and their
purposes of pollination. Digestion may seem odd, but all panicked motions continue to stimulate the trigger hairs.
plants produce enzymes that have digestive capabilities – This encourages the traps to seal completely, suffocating
carnivorous plants have only relocated the site of enzy- the prey, and to release digestive enzymes. (Children who
matic activity to some external pitcher or leaf surface. feed dead flies to their pet Venus flytraps are often disap-
Finally, absorption of nutrients is something that all pointed when, the next day, the uninterested plants open
plants do (or, at least, all that survive past the cotyledon their traps and reject the inanimate morsels – only live
stage). Carnivorous plants have simply combined these prey stimulate the leaves enough to complete the digestion
conventional plant traits into a novel foraging strategy. process.) After about one week, the leaves reopen to reveal
the crispy exoskeleton that survived the digestion process.
While Dionaea muscipula is but one species, we know of
Types of traps
about 600 other carnivorous plants in 17 genera across ten
The Venus flytrap (Dionaea muscipula; title image) is the different plant families. With so many species, it should be
best known carnivorous plant. It has a ground-hugging no surprise that the bear-trap approach is not the only
rosette of leaves that look and function like hinged, trick that carnivorous plants use! In Table 1, carnivorous
foothold bear traps. A detailed study reveals a fascinating genera are listed along with their primary methods of prey
array of refinements in its hunting technique. Rapid leaf capture. The diversity of techniques these plants employ is
closure is triggered when the scrabbling of prey bends impressive.
hairs on the leaf-lobe surface. But, in order to weed out The simplest kind of trap is the pitfall. In this, the plants
create a chamber (in basic form, a funnel or vase) into
Title image: The red maw of the Venus flytrap (Dionaea muscipula) which prey plummet. Digestive enzymes in the bottom of
beckons! the vase perform the expected function, and nutrients are

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Table 1. Types of carnivorous plants Another carnivorous tactic is to develop leaves

with glandular, sticky surfaces. These leaves may
Genus Common Name Trap Type emit either a sugary or fungal smell, with different
smells attracting different prey. Mucous-exuding
Aldrovanda waterwheel plant bear trap glands give the plant the appearance of being
Byblis rainbow plant sticky flypaper coated with delicious nectar. On some genera,
Cephalotus Albany pitcher plant pitfall these glands are perched high upon tentacles; on
Darlingtonia cobra lily, California pitcher plant sticky flypaper others (i.e., Pinguicula; Figure 2a) the glands are
Dionaea Venus flytrap bear trap sessile at the leaf surface. Once prey lands upon
Drosera sundew sticky flypaper the leaves, their legs and wings become mired, and
Drosophyllum dewy pine sticky flypaper escape is impossible. Struggle only ensures their
Genlisea corkscrew plant lobster pot death, as mucus coats and suffocates them.
Heliamphora sun pitcher pitfall Digestion occurs on the spot, and the leaf may even
Nepenthes tropical pitcher plant pitfall, sticky flypaper curl over the prey to increase the number of diges-
Pinguicula butterwort sticky flypaper tive glands contacting the invertebrate morsel. The
Roridula (none) sticky flypaper genera of these sticky-leafed carnivores include
Sarracenia trumpet pitcher pitfall, lobster pot Byblis, Drosera (Figure 2b), Drosophyllum,
Triphyophyllum (none) sticky flypaper Pinguicula, Roridula and Triphyophyllum
Utricularia bladderwort suction trap Kingdom Plantae has yet other devious machin-
ery to use against Animalia. Aldrovanda is an
aquatic sister genus to the Venus flytrap. Genlisea
absorbed into the plant. Yet even this simple carnivorous is a lobster-pot aquatic (having entered, prey cannot find
leaf is often modified with innovations to improve the the escape hatch), while Utricularia (Figure 3) has evolved
trap’s efficacy. The trap walls may be covered with down- more than 220 species of carnivores that use bizarre
ward pointing hairs or waxy, slimy surfaces. Enticing suction traps to draw tiny aquatic organisms into their
odours or beguiling pseudo-floral pigmentation patterns bug-thirsty, digestive bladders (Figure 3b).
may attract prey. Strategically placed nectaries may be Some species, dissatisfied with convention, combine
situated directly over the pitfall opening to encourage techniques. Sarracenia psittacina is in a genus of pitfall
browsing insects to wander to the locations of maximal carnivores, but its traps are tilted sideways. It appears to
peril (Nepenthes bicalcarata; Figure 1a). The structure of be mostly harmless except during periodic floodings, when
the pitcher may be modified with overhanging ledges to it captures aquatic animals using a lobster pot technique.
frustrate insects trying to climb to safety. Some pitfall Meanwhile, Nepenthes inermis and a few other species of
traps are equipped with transparent windows that illumi- pitcher plants have slimy inner trap walls, and function as
nate the pitcher interior – this fools insects into believing sticky trap plants.
the shining digestive fluid is a portal to freedom. Once the In a surprising step towards botanical disarmament, one
prey have fallen into the pit, they find themselves in more bladderwort species (Utricularia purpurea) may have
than just a bath of weak enzymes – pitcher fluid has been abandoned its carnivorous congeners, as it seems its blad-
found to contain wetting agents, toxins, and even mild ders may primarily be used to house algae and zooplankton
anaesthetising agents! Genera of pitcher plants occur in a mutualist relationship. Evolution leads to constant
throughout the world: Sarracenia (Figure 1b) and innovation!
Darlingtonia in North America, Heliamphora in South
America, Cephalotus in Australia, and Nepenthes (mostly)
Prey spectrum
in southeast Asia, but with a few species found elsewhere.
Two New World bromeliad genera Catopsis and Brocchinia Various organisms are captured and eaten by carnivorous
are also suspected of being carnivorous. plants. The list includes arthropods, such as insects, arach-
nids, millipedes, centi-
pedes, annelids and
a b crustaceans, as well as
slugs and snails, and
even small vertebrates
such as amphibians
and reptiles. A few
dead rodents have
even been found in
Nepenthes pitchers,
but these captures are
certainly incidental
and rare. This diverse
array of prey presents
a quandary for lexicog-
raphers, who cannot
decide what to call
these plants. None of
the proposed words
work, neither ‘carnivo-
rous’ (insects do not
Figure 1. (a) The fierce pair of spines on Nepenthes bicalcarata are actually only nectaries to attract prey. contain meat) or
(b) Sarracenia leucophylla – a North American pitcher plant – growing in Alabama. ‘insectivorous’

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a b

Figure 2. (a) A gnat is absorbed into the slimy leaf of the butterwort Pinguicula macroceras subsp. nortensis. (b) The small leaves of the
Australian sundew Drosera auriculata can trap comparatively large prey.

(centipedes are not insects). Even ‘animal-eating’ is too habitats. (Specialist horticulturists know that one of the
restrictive a term, since some carnivorous plants (e.g., fastest ways to kill their prized carnivorous gems is to
Genlisea) delight and perhaps even specialise in consum- fertilise them!)
ing protozoa. Examples of nutrient-poor ecosystems that are friendly
to carnivorous plants include epiphytic habitats (using
other plants for structural support) in tropical moist
Why are they carnivorous?
forests, seasonally wet acidic or neutral pH deserts, and a
With such a large number of carnivorous plant species, it wide variety of acidic wetlands such as peat bogs, marshes,
should be no surprise that they live in a wide variety of swamps, wet savannahs and fens. A frequent indicator for
habitats. But carnivorous plant habitats share an impor- habitats suitable for carnivorous plants is the presence of
tant characteristic – they all hunt in habitats that are Sphagnum moss. This moss tends to buffer the pH to
deficient in some essential nutrients, nutrients that are highly acidic levels that favour carnivorous plants. (Among
readily available in animal tissues. It is this environmen- other effects, high acidity reduces decomposition rates so
tal stress that gives carnivorous plants a selective advan- few nutrients are available.)
tage. By diverting resources into making specialised struc- Carnivores are not picky in terms of temperatures –
tures, such as prey-trapping leaves, the plants benefit by tropical to boreal habitats are all populated by carnivorous
harvesting nitrogen and other nutrients from captured plants. Marine or brackish habitats are the only major
prey. In contrast, in a nutrient rich environment, the ecosystem type not penetrated by these hungry botanicals,
carnivorous approach is not a valuable strategy. In fact, no doubt because of the high availability of nutrients in
carnivorous plants are so specialised for low nutrient such conditions.
conditions that they cannot survive in nutrient-rich Such a wide variety of habitats means that carnivorous
plants are widely distri-
buted, more than most
a b people suspect. They are
found on every continent
(except Antarctica), and
their global centres of diver-
sity include southeastern
Asia (Nepenthes), southeast-
ern USA (Sarracenia), west-
ern Australia (Drosera and
Utricularia), northern South
America (Heliamphora), and
southern Mexico/central
America (Pinguicula).

New research
Traditional botanical studies
are still active (new species
are being described each
year), and ecologists are
broadening (and complicat-
ing!) our understanding of
carnivorous plants. We are
learning that, instead of
capturing anything small
Figure 3. (a) A cultivar bladderwort flower, Utricularia calycifida ‘Asenath Waite’. (b) Aquatic bladders enough to fit into their
of Utricularia intermedia. leaves, some carnivorous

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C a r n i v o r o u s p l a n t s

species specialise in specific prey types. Nepenthes are Not all carnivores appreciate foragers, however. It is
particularly adept in this dimension – Nepenthes albomar- possible that the leaf-curling of some Pinguicula around
ginata may specialise only on foraging termites, while the prey may be, in part, to jealously protect the prey from
‘most scatological’ award must surely go to Nepenthes lowii thieving kleptoparasites. Might it be that the dramatic leaf
(Figure 4). This plant may not be strictly carnivorous at all. curling seen in so many Drosera species is not intended to
It produces edible exudates on its pitcher lids that are irre- help digest the prey, but rather is primarily to protect the
sistible to sunbirds, and as the birds feast they excrete into food items from theft?
the pitchers, much to the plant’s coprophagous satis- Laboratory research has of course been active, with
faction. Another innovative species is Nepenthes particularly active investigations directed towards under-
ampullaria. This plant is noteworthy for producing standing the molecular mechanisms of nutrient uptake,
clusters of ground pitchers that carpet the ground. These the importance of carnivory for growth, and the nature of
pitchers may function in part to capture detritus raining the digestive juices secreted by carnivores.
from the forest canopy. Even research into the history of carnivorous plant
Of particular research interest is the notion that non- botany has been active, if in a particularly bawdy direction.
plant accomplices may be cooperating with the carnivores. The logic behind the coining of the common name ‘Venus
For example, the digestion in many of the pitcher plant flytrap’ has been essentially inexplicable. The explanation
genera may be performed, at least in part, by bacteria or concocted by its discoverers – New World botanists and
other inquiline fauna (lodgers), such as larval flies, their cohorts in England – was that its flowers were as
mosquitoes and even tadpoles. Such organisms help digest beautiful as the goddess, Venus, was. The plant’s small,
the prey and excrete useful nitrogenous compounds into drab, white and green flowers do not corroborate this
the pitcher fluid. Some scavenging organisms, such as unlikely explanation. Furthermore, the poorly Latinised
spiders (Misumenops nepenthicola and Thomisius version of this name (by botanists who certainly could do
nepenthephilus) or diving ants (Camponotus schmitzi), better) added to the confusion (Dionaea muscipula means
plunge into their friendly Nepenthes pitchers to retrieve ‘Aphrodite’s mousetrap’ – suggesting that it is a catcher of
large prey items. Perhaps without the intervention of these mammalian prey and not of insects). Recently Nelson and
opportunistic feeders, nutrient overload could occur, which McKinley (1990) revealed the smutty secret. Those puri-
would damage the pitcher through over-feeding. The mutu- tanical naturalists of Botany’s past envisioned – in the
alism between ants and Nepenthes bicalcarata (Figure 1a) plant’s quivering red lobes, sensitive behaviour and attrac-
is so advanced that the tendril supporting the pitcher is tive qualities – similarities to female genitalia. The
hollow and inflated, providing excellent nesting cavities for Goddess of Love was invoked, and the jest was immor-
the ant allies. talised with the Latin name. (The official explanation
Another kind of mutualism is exemplified by the case of about the attractive flowers was simply a cover story to
Capsid bugs (e.g., Cyrtopeltis and Setocornis species), protect the delicate sensibilities of the drawing room
which safely navigate the adhesive glands of Drosera and ladies.) Later, William Bartram (a participant in the
Byblis sundews to eat the prey captured on the sticky conspiracy) was so bold as to waggishly write in his Travels
leaves. The bugs apparently produce excrement that … of this ‘sportive vegetable’ that seduced incautious
contains useful nitrogenous compounds. Indeed, when insects with its ‘incarnate lobes’.
deprived of these bugs, some carnivorous Byblis species are
completely incapable of translating captured prey into
Carnivores in captivity
absorbable nutrients. Roridula (a genus of South African
plants) was temporarily removed from the list of carnivo- Carnivorous plants are not easy to grow for the casual
rous species when it was noted that its glandular leaves horticulturist. Even Darwin noted that he was unsuccess-
were resinous and not mucosal. It is apparently carnivo- ful with his Venus flytraps. However, with just a small
rous after all, as long as you are willing to allow the mutu- amount of research, representative species of most of the
alism with assassin bugs. carnivorous genera can be grown with only minor pain.
Horticultural interest in carnivorous
plants rises and falls – the first noteworthy
peak was in the Victorian era when
Nepenthes and orchids both were grown
(and more often killed) by the voracious
plant hunters and their financing nurseries
in Great Britain. The tide of interest is high
once again, and this time the internet has
helped increase communication among scat-
tered, isolated enthusiasts. In addition, the
membership numbers in carnivorous plant
organisations are rising. The most promi-
nent group is the International Carnivorous
Plant Society (ICPS), which produces a fine
quality journal* called Carnivorous Plant
Newsletter, which includes comments on
cultivation, conservation, new carnivorous
cultivars and even peer reviewed scientific
papers, such as new taxon descriptions.

* The author is coeditor for Carnivorous Plant

Figure 4. Nepenthes lowii – an avian lavatory – cultivated in California. Newsletter!

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hoping to capture the genetic variety of carniv-

orous plants are too late – the cake has already
been consumed, and only the crumbs remain.
The fate of the remaining carnivorous plant
communities are not at all certain – will these
fragmented populations throughout the world
survive the present era of anthropogenic extinc-
tions? Will they survive the effects of global
climate change? We will see (or rather, our chil-
dren will). These plants, which require clean
and unspoiled habitats to survive, are the
fabled canaries in the coal mine – their fate is
our own.

References
Clarke C (1997) Nepenthes of Borneo. Natural
Figure 5. Another of the few remaining prime habitats for Sarracenia being History Publications: Kota Kinabalu, Borneo.
destroyed in Florida. Clarke C (2001) Nepenthes of Sumatra and peninsular
Malaysia. Natural History Publications: Kota
Kinabalu, Borneo.
Conservation
D’Amato P (1998) The savage garden: cultivating carnivorous
Carnivorous plants are greatly threatened by human activ- plants. Ten Speed Press: Berkeley, California.
ity, for, while they are not directly targeted by humans as a Hartmeyer S (1998) Carnivory in Byblis revisited II: the phenom-
desirable resource, they are strongly damaged by our inci- enon of symbiosis on insect trapping plants. Carnivorous Plant
dental activities. Humans have tried a number of methods Newsletter, 27, 110–113.
to ‘improve’ or ‘reclaim’ their habitats. Wetlands are Juniper B E, Robins R J and Joel D (1989) The carnivorous plants.
drained (Figure 5), forests are slashed and burned, lands Academic Press: London.
are fertilised, natural wildfires are suppressed – all of these Kitching R L (2000) Food webs and container habitats: the natural
activities are harmful to carnivorous plant habitats. history and ecology of phytotelmata. Cambridge University
Simultaneously, nutrient-rich pollutants are allowed to Press: Cambridge.
seep from industrial or agricultural sources into biologi- Lloyd F E (1942) The carnivorous plants. Chronica Botanica: New
cally rich areas dismissed as ‘wastelands’. The nutrient York, USA.
flux from these sources alters the soil and water chemistry Meyers-Rice B A (2001) Rare Sarracenia poaching and the ICPS.
so much that the carnivorous species are poisoned while Carnivorous Plant Newsletter, 30, 43–50.
the non-carnivorous native and non-native species over- Nelson E C and McKinley D L (1990) Aphrodite’s Mousetrap.
whelm the ecosystem. The extirpation of the remarkable Boethius Press: Aberystwyth, Wales.
Aldrovanda vesiculosa (an aquatic version of the Venus (Further reading suggestions are listed on our website at
flytrap) from most of its range in Europe and Japan is one www.iob.org/biologist.asp)
example of this process.
Frequently, the effects of pollutants kill the sphagnum
Websites
moss in wetlands. (The importance of sphagnum moss in
these environments cannot be overstated – the moss domi- www.sarracenia.com
nates the wetland biomass, and creates the hydrological The author’s web site, includes an extensive ‘Frequently Asked
and habitat structure that defines the ecosystem.) When Questions’ area and photographic gallery.
nutrient levels rise, the sphagnum rapidly dies and the
www.carnivorousplants.org
entire structure of the habitat collapses.
The International Carnivorous Plant Society, which publishes
The effects of habitat conversion by development, agri-
Carnivorous Plant Newsletter.
cultural fragmentation of once continuous plant ranges,
altered hydrology, pollution, modified fire regimes, inva- www.labs.agilent.com/bot/cp_home
sive species, and poaching by the nursery trade and private Jan Schlauer’s taxonomic database of all carnivorous plant Latin
enthusiasts all conspire to make conservation of carnivo- names, cultivar names, and synonyms. Includes illustrations.
rous plant species extremely challenging and, perhaps, not
particularly successful. The poorly understood effects of Barry Rice completed his PhD in Arizona in 1995. He works
imminent global climate change do not make the future as an invasive species scientist for The Nature Conservancy, a
look very encouraging. nonprofit conservation organization. His interest in carnivo-
The horticultural community has become increasingly rous plants originated in 1985 when he decided to grow Venus
interested in conservation, and organisations like the ICPS flytraps he found captive in a corner flower shop. His interest
and the British CPS have conservation grant programmes. has continued, and while he is still active in horticulture, most
However, the relationships between horticulturists and of his spare time is spent doing fieldwork, plant photography,
conservation groups are not always easy – numerous and coediting the International Carnivorous Plant Society’s
poaching events have poisoned the good will of many journal. He is currently working on promoting a number of
conservation workers. Still, innovative programmes are carnivorous plant conservation initiatives.
being implemented. PO Box 72741
Field studies and laboratory research continue to reveal Davis, CA 95617
new information about these plants. However, this botani- USA
cal legacy is disappearing. In the USA, the vast majority of [email protected]
the wetlands have been destroyed. Conservationists

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