1. What are carnivorous plants? 4 criteriaHabitatPaper #1 – Phylogeny and structural evolution
• What are paracarnivorous plants?
Digestive mutualismsPaper #2 - RoridulaOther paracarnivores
3. SummaryEvolution of carnivorous and paracarnivorous plants
Outline
“I See You”, man-eating tree of Central/South America.
What are carnivorous plants? “Flesh-eating”; 4 criteria (Lloyd 1976).
2. Trap and killtraps: sticky, pitfall, strategic hairs, waxesdeath: usually drowning, suffocating, some eaten alive
4. Absorb nutrients
3. Digest with endogenous enzymes
Adaptation or coincidence?(i.e. herbivore defense: amber)
more difficult to observe:
• Attract animal prey (old name “insectivorous”)
visual or chemical cuesrange from deceptive mimicry rewards
?
Typical habitat: sunny, moist, nutrient poor
Ombrotrophic – “cloud fed”StagnantAnaerobicAcidic
Givnish, 1984. Carnivory in the bromeliad Brocchinia reducta, with a cost/benefit model for the general restriction of carnivorous plants to sunny, moist, nutrient-poor habitats.
Rare: <0.2% of all plants
Typical habitat: sunny, moist, nutrient poor
Decreased decompositionDecreased nitrification by bacteria
Increased denitrification by bacteria(nitrogen is lost to the atmosphere)
StagnantAnaerobic
Acidic
Atypical habitat: nutrient poor, semi-arid or epiphytic,
Drosophyllum lusitanicum
Nepenthes veitchii
Pinguicula ramosa
Early carnivorous plants? Archaeamphora longicervia
Li, Hongqi, 2005. Early Cretaceous sarraceniacean-like pitcher plants from China.
Hess, 2005. Evidence of Zoophagy in a Second Liverwort Species, Pleurozia purpurea
Structural evolution: Paper #1 - Albert et al. 1992
rbcL - ~1000 bp
Rubisco - highly conserved
13 carnivorous genera in 4 orders
(Note: Bignoniales has been collapsed into Lamiales)
4 basic trap types• Flypaper traps F
• Pitcher traps P
• Bladder traps B
• Snap traps S
Structural adaptations: Flypaper traps F
Order Ericales
Order CaryophyllalesOrder Lamiales
(Pinguicula sp.)
Structural adaptations: Pitcher traps P
Order Ericales
Order Poales1 species:
Brocchinia reducta (not in Albert 1992)Order Oxalidales
Order Caryophyllales
Epiascidiation: inrolling and fusion of leaves (pseudo-stomach?)
Phytotelma: fluid collected in a plant structure (rainwater, digestive enzymes)
aka pitfall traps – floral mimicry, nectar lure, slippery surface wax, downward pointing hairs
Structural adaptations: Bladder traps B (1 genus, ~250 spp.)
order Lamialesfamily Lentibulariaceae
Utricularia
Structural adaptations: Snap traps S
Aldrovanda vesiculosa (not in Albert 1992)
Dionaea musipula
order Caryophyllales family Droseraceae
Evolution of carnivorous plants: from Albert 1992 - present
Genlisea
Triphyophyllum
Aldrovanda
Brocchinia
Stevens, P. F., 2008. Angiosperm Phylogeny Website.
1992: 13 genera 4 orders 4 trap types2009: 14 genera 5 orders 5 trap types
Structural adaptations: Lobsterpot traps L (1 genus, 21 spp.)
order Lamialesfamily Lentibulariaceae
Genlisea(not in Albert 1992)
Flypaper traps FPitcher traps PBladder traps BSnap traps SLobsterpot traps L
Structural adaptations: Lobsterpot traps L Genlisea
(not in Albert 1992)
Caryophyllales – family Dioncophyllaceae, Triphyophyllum peltatum (1 sp.)
Evolution of carnivorous plants: from Albert 1992 - present: 5 orders
Ericales: 2 genera, 29 spp.
F
P
Caryophyllales: 6 genera, 298 spp.
F SP Poales: 1 sp.
P
Oxalidales: 1 sp.
P
Lamiales: 4 genera, 252 spp.
F
B
L
Flypaper traps FPitcher traps PBladder traps BSnap traps SLobsterpot traps L Stevens, P. F., 2008. Angiosperm Phylogeny Website.
1992: 13 genera 4 orders 4 trap types2009: 14 genera 5 orders 5 trap types
FS
P
B L
P
P
PF
F
Flypaper traps FPitcher traps PBladder traps BSnap traps SLobsterpot traps L
Evolution of carnivorous plants: from Albert 1992 - present: 5 orders
S
B L
Flypaper traps FPitcher traps PBladder traps BSnap traps SLobsterpot traps L
Convergent trap types
Evolution: Oxalidales: Cephalotus follicularis (1 sp.)
F
P
P
P
PF
F
S
B L
Ericales: Darlingtonia
Flypaper traps FPitcher traps PSnap traps SBladder traps BLobsterpot traps L
Evolution of carnivorous plants: from Albert 1992 - present
Caryophyllales: 6 genera, 298 spp.
SP
F
F
P
P
P
PF
F
S
B L
Evolution: Caryophyllales - Nepenthes
N. rajafrogs, lizards and ratsup to 3 liters
Evolution: Caryophyllales – Drosophyllum “dewy pine”
Evolution: Caryophyllales – Drosera “sun-dews”
Thigmotropism - oriented growth of an organism in response to mechanical contact
Evolution of carnivorous plants: from Albert 1992 - present
Lamiales: 4 genera, 252 spp.
F
B
L
Flypaper traps FPitcher traps PBladder traps BSnap traps SLobsterpot traps L
F
P
P
P
PF
F
S
B L
Evolution: Lamiales – Pinguicula “butterworts”
Evolution of carnivorous plants: from Albert 1992 - present
Ericales: 2 genera, 29 spp.
F
P
Flypaper traps FPitcher traps PBladder traps BSnap traps SLobsterpot traps L
F
P
P
P
PF
F
S
B L
Supplemented carnivory: Ericales – Sarracenia and Roridula
supplemented bydigestive mutualismConiine in trap nectar:
same neurotoxin as poison hemlock
endogenous enzymes
Ellison et al. 2003.
Phytotelma inquiline food web
Roridula
Other Ericaleans with
digestive mutualisms:
1. What are carnivorous plants? 4 criteriaHabitatPaper #1 – Phylogeny and structural evolution
• What are paracarnivorous plants?
Digestive mutualismsPaper #2 - RoridulaOther paracarnivores
3. SummaryEvolution of carnivorous and paracarnivorous plants
Outline
“I See You”, man-eating tree of Central/South America.
What are paracarnivorous plants?(aka proto-carnivorous, subcarnivorous, pseudo-carnivorous, part-time carnivorous)
1. Attract 2. Trap/kill 3. Endogenous enzymes
Givnish, 1984
digestive mutualisms
?
What are digestive mutualisms? Darlingtonia californica (1 sp.)
no endogenous enzymes detected
digestive mutualisms:
“digestion by proxy”(exogenous enzymes)
(Bacteria)
Slime mites: Sarraceniopus darlingtoniae
Midge larvae: Metriocnemus edwardsi
(Fungi)
Spiders
Fashing, 2004. Biology of Sarraceniopus darlingtoniae (Histiostomatidae: Astigmata), an obligatory inhabitant of the fluid-filled pitchers of Darlingtonia californica (Sarraceniaceae).
enzymaticpartners:
bacteriafungi
animals
+ +enzymes rewards
gut bacteria animalsmycorrhizal fungi plants
Paracarnivores with digestive mutualisms: Habitat similarity - sunny, moist, nutrient poor
Paracarnivorous plants treated as carnivorous in Albert 1992
Proboscidea
Heliamphora
Darlingtonia
digestive mutualism
digestive mutualism
Paracarnivores with digestive mutualisms: Habitat similarity
Guiana highlands, Venezuela
Paracarnivorous plants with digestive mutualisms: Heliamphora
D’Amato, Peter, 1998. The Savage Garden
(no endogenous enzymes found)
Guiana highlands, Venezuela
frog digestive mutualism suspected, no experimental data
Paracarnivore or true carnivore? Brocchinia reducta
Givnish, 1984. Carnivory in the bromeliad Brocchinia reducta, with a cost/benefit model for the general restriction of carnivorous plants to sunny, moist, nutrient-poor habitats.
Guiana highlands, Venezuela
Paracarnivore or true carnivore? Brocchinia reducta
Isley, Paul,1987. Tillandsia.
digestive mutualism(pseudoscorpions)
4. Absorb?
but endogenous enzymes?
Givnish, 1984.
Digestive enzymes
Plachno et al. 2006. Fluorescence labeling of phosphatase activity in digestive glands of carnivorous plants. Plant Biology.
weak endogenous enzymes
Endogenous enzymes found in carnivorous plants?
•amylase (starch)•chitinase (also fungal and arthropod defense, barley seeds...)•esterase•lipase •peroxidase•phosphatase (also respiration, dephosphorylation...)•protease (i.e. nepenthacine “similar to pepsin”)•ribonuclease
Dolling WR, Palmer JM. 1991. Pameridea (Hemiptera: Miridae): predaceous bugs specific to a highly viscid plant genus Roridula.
Pameridea: 2 sp. P. roridulae, P. marlothi
(+ pollination mutualism?)
Paracarnivory - Paper #2: Roridula: 2 sp. R.gorgonias, R. dentata
digestive mutualism
larger prey microhabitat
gut bacteria animalsmycorrhizal fungi plants
+ +enzymes rewards
?
Paracarnivore or true carnivore? Roridula
glue
Anderson et al., 2007. Density-dependent outcomes in a digestive mutualism between carnivorous Roridula plants and their associated hemipterans
Paracarnivore or true carnivore? Roridula
Anderson et al., 2002. It takes two to tango but three is a
tangle: mutualists and cheaters on the carnivorous plant Roridula.
but endogenous enzymes?
Paracarnivore or true carnivore? Roridula
Plachno et al. 2006.
Roridula
Plachno et al. 2006.
Anderson B, 2005. Adaptations to foliar absorption of faeces: a pathway in plant carnivory.
(+ absorption via cuticular gaps)
Paracarnivore or true carnivore? Roridula
Paracarnivore or true carnivore? Lamiales: Byblis
+ suspected digestive mutualism (also hemipteran feces), currently not studied?
Plachno et al. 2006.
Other fecally supplemented carnivores – Nepenthes lowii
Clarke, Charles, 1997.Nepenthes of Borneo.
At least 2 digestive mutualists(no experimental data)
3.sunbirds4.inquiline crane fly larvae
mimicry?
(tendril domatium)
Other fecally supplemented carnivores – Nepenthes bicalcarata
digestive mutualisms?
Paracarnivores or true carnivores? Catopsis and Paepalanthus
Catopsis berteroniana
Paepalanthus bromelioides(no endogenous enzymes detected)
(no endogenous enzymes detected)
Radhamani, et. al., 1995. Defence and carnivory: Dual role of bracts in Passiflora foetida.
Paracarnivores or true carnivores? Passiflora and Capsella
questionably detected protease...
Barber, J.T., 1978. Capsella bursa-pastoris seeds: Are they "carnivorous"?
(no enzyme data)
Darnowski, et. al., 2006. Evidence of protocarnivory in triggerplants (Stylidium spp.; Stylidiaceae).
Paracarnivores or true carnivores? Stylidium
absorption not detected
questionably detected protease...
Floral traps? Or herbivore defense? Plumbago (no enzyme or absorption data)
Spomer, G.G.,1999. Evidence of protocarnivorous capabilities in Geranium viscosissimum and Potentilla arguta and other sticky plants.
Puya raimondii and Pisonia grandis: avian-specific paracarnivores?!
L?
Rees et al., 1978. Puya raimondii (Pitcairnioideae, Bromeliaceae) and birds: an hypothesis on nutrient relationships.
Puya raimondii and Pisonia grandis: avian-specific paracarnivores?!
“The fate of carcasses and seeds attached to carcasses”
22 bird carcasses were monitored daily.
Scavengers usually found the carcasses within a day, and 91% were dismembered. The mean time to dismemberment was 3.3 days
“...no seeds tolerated prolonged immersion in thesea for 12 d or more (as might be experienced by seedsattached to a dead bird floating on the sea)”
“Although birds killed by entanglement add nutrients to the soil, the amounts would be trivial compared with the massive inputs from guano, failed eggs and dead chicks.”
“Seeds attached to carcasses did not have improved germination or survival.”
Burger, A, 2005. Dispersal and germination of seeds of Pisonia grandis, an Indo-Pacific tropical tree associated with insular seabird colonies
F?
Summary
Carnivorous Non-carnivorous• Convergent evolution
• Alternative nutrient pathways, cost/benefits of carnivory
• Phytotelmata inquiline food web dynamics (discrete boundaries, easily manipulated)
• Coevolution, stability, assemblage of digestive mutualisms, the role of fungal and
bacterial digestive mutualists
The end
Over 600 species of plants in 13 families have evolved adaptations to enable them to attract, trap, kill, digest and absorb animal prey. Although Darwin referred to these plants as “insectivorous”, they have been known to catch small frogs, fish, and even rats. These plants may use visual or chemical attractants to lure their victims, and occasionally even offer nectar as bait.
What are carnivorous plants?
Where do they grow?Carnivorous plants grow in nutrient poor habitats that are usually sunny
and water logged. A common misconception is that they only grow in the tropics - many species are native to North America, and Utricularia can be found in Siberian bogs. Carnivorous plants are found where plant growth is limited by a lack of nutrients, especially nitrogen, which is necessary for DNA synthesis and abundant in plant and animal tissue.
Brocchinia reducta
Native to highly leeched soils in the Guiana highlands, the leaves of Brocchinia are rolled into a tube shape that collects rainwater, and crawling insects slip on the waxy surface, fall in, and drown. This method of “pitfall” trap is found in true carnivorous plants, which fill this fluid with digestive enzymes. Rather than enzymes, Brocchinia is thought to rely on predators such as pseudoscorpions, as well as microorganisms, to digest their prey.m
Roridula gorgonias
Scientists have long speculated whether or not Roridula should be considered a true carnivore. This is because although Roridula readily traps insects on its sticky hairs, plants in the wild rely on a mutualism with an insect, Pameridia, which patrols the plant and feeds on the trapped prey. Roridula appears to derive most of its nitrogen not from the prey directly, but from the nitrogen-rich feces of Pameridia, and so is often considered a “paracarnivorous” plant.
Carnivorous or Paracarnivorous?
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