6/29/09!

Plant Oddities - ! Plant Oddities - ! the Carnivores, Parasites & Saprotrophs ! the Carnivores, Parasites & Saprotrophs !

Of all the plants, the most bizarre, the least Of all the plants, the most bizarre, the least understood, but yet the most interesting are understood, but yet the most interesting are those plants that have unusual modes of those plants that have unusual modes of nutrient uptake.! nutrient uptake.!

Carnivore: ! Parasite: Rafflesia!

Plant Oddities - ! Plant Oddities - ! the Carnivores, Parasites & Saprotrophs ! the Carnivores, Parasites & Saprotrophs !

Of all the plants, the most bizarre, the least understood, but yet the most interesting are those plants that have unusual modes of At the state, federal, and international levels, many of nutrient uptake.! these plants are extremely rare, threatened, or endangered. Many are protected internationally from collection by CITES.!

Saprotroph: Monotropa!

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Plant Oddities - ! Plant Oddities - ! the Carnivores, Parasites & Saprotrophs ! the Carnivores, Parasites & Saprotrophs ! Three aspects of these plants present problems. ! Two factors for this systematic confusion! 1. The biology of how they trap animals, recognize and penetrate other plants, and how they interact with fungi is • !the specialized roles often involve reductions in both often little understood.! vegetative and floral features!

2. The basic anatomical and morphological structures of both • !often unrelated members of these groups converge unto the vegetative and floral parts of these plants are often so modified same morphology; often related members diverge in that basic structure (homology) is little understood.! morphology!

3. Systematic relationships of 3. Systematic relationships of these plants (how to classify these plants (how to classify these organisms) are often poorly these organisms) are often poorly known and controversial.! known and controversial.!

How are these three families classified?! Plant Oddities - ! All possible combinations of the three have been proposed!! the Carnivores, Parasites & Saprotrophs ! The most influential classification system was that proposed by Arthur Cronquist Classic example of this systematic problem is a set of three families of who placed all three families in one order - the Nepenthales! carnivorous plants with various types of trapping mechanisms:!

Pitcher traps! Fly paper and steel traps!

Nepenthaceae -! -! -! Asian ! American pitcher plant! Sundews and Venus fly trap!

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How are these three families classified?! DNA sequence data indicates something quite different!! Plant Oddities - ! the Carnivores, Parasites & Saprotrophs ! The Asian and American pitcher plant families quite unrelated; the Asian pitcher plants placed with sundew family and American pitcher plants with Ericaceae! Why do plants have these unusual life styles?! Lack of basic necessity for life on land.!

+3! parasite! CO3 uptake! saprotroph! CO2! light! aquatic! tropical/temperate forest!

bogs, sand, ! desert/epiphyte! weathered bedrock! H O! nutrients! CAM! 2 parasite! C4! saprotroph! succulence! carnivore! parasite!

the Carnivores! the Carnivores!

Charles Darwin (and his grandfather) Charles Darwin (and his grandfather) was the first to painstakingly study was the first to painstakingly study carnivorous plants.! carnivorous plants.!

In his book on “Insectivorous plants”, he In his book on “Insectivorous plants”, he showed that they had adaptations to showed that they had adaptations to capture and digest animals.! capture and digest animals.!

Tom Givnish, University of Wisconsin, has refined the definition of what is a :!

• !Adaptations to lure, capture, and digest prey! • !Ability to absorb nutrients from animals!

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the Carnivores! the Carnivores!

Luring device of some type often Luring device of some type often involving color, movement, and involving color, movement, and smell ! smell !

Trapping device of some type (pitchers or drowning pools, steel traps, sticky leaves, etc.)!

the Carnivores! the Carnivores!

Luring device of some type often Luring device of some type often involving color, movement, and involving color, movement, and smell ! smell !

Trapping device of some type Trapping device of some type (pitchers or drowning pools, (pitchers or drowning pools, steel traps, sticky leaves, etc.)! steel traps, sticky leaves, etc.)!

Ability to digest animals trap, often Ability to digest animals trap, often with release of pronases and other with release of pronases and other enzymes into pool or on animal! enzymes into pool or on animal!

Mechanisms to uptake amino acids Amino acids radioactively labeled once animal is digested, often with being incorporated into the scales of specialized hairs or scales! Brocchinia (pineapple family)!

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the Carnivores! the Carnivores!

Plants who may accidentally kill (drown in this case) So it is not surprising that carnivores show up in groups that have “pre-adaptations” to the carnivory life style.! What are not carnivores?! animals and even be able to utilize their amino acids; leaf What are not carnivores?! “pitcher” in this case is simply an adaptation to collect Shown here are two species (A and B) of Brocchinia water as an epiphyte.! that are carnivores in the pineapple family.!

They are closely related to other species in the genus that impound water or are ant-fed, but not carnivorous.!

Billbergia! Bromeliaceae!

the Carnivores! the Carnivores!

Carnivorous plants are found in 19 genera, all but 2 are dicots. They are classified Carnivorous plants are centered in 3 nutrient poor bedrocks around the world.! into 9 groups, although many groups individually exhibit several kinds of carnivory modes.! Southeastern United States coastal plain: the ancient erosional product of the Appalachian uprise and with boggy peatlands!

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the Carnivores! the Carnivores!

Carnivorous plants are centered in 3 nutrient poor bedrocks around the world.! Carnivorous plants are centered in 3 nutrient poor bedrocks around the world.!

Southeastern United States coastal plain: Southeastern United States coastal plain: the ancient erosional product of the the ancient erosional product of the Appalachian uprise and with boggy Appalachian uprise and with boggy peatlands! peatlands!

Western Australia - a Precambrian Western Australia - a Precambrian bedrock, highly leached, and nutrient bedrock, highly leached, and nutrient poor! poor!

Guayana Highlands of southern Venezuela and adjacent areas of Brazil and Colombia - the higher elevation “tepuis” are rain drenched and extremely nutrient poor!

the Carnivores! Types of Carnivores!

Carnivorous plants are centered in 3 nutrient poor bedrocks around the world.! Passive traps! Southeastern United States coastal plain: the ancient erosional product of the ! pitfall ! Appalachian uprise and with boggy ! !Sarraceniaceae - American pitcher plants! peatlands! ! !Nepenthaceae - Asian pitcher plants! ! !Bromeliaceae - “pineapple” pitchers! ! !Cephalotaceae - Australian pitcher plant! Western Australia - a Precambrian bedrock, highly leached, and nutrient poor!

Guayana Highlands of southern Venezuela and adjacent areas of Brazil and Colombia - the higher elevation The tepui region was made famous by Sir “tepuis” are rain drenched and Conan Doyle as the “Lost World” of extremely nutrient poor! “Jurassic” and “Arachniphobia” fame!

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Types of Carnivores! Types of Carnivores!

Passive traps - pitfall ! Passive traps - pitfall ! ! !! ! !!

Heliamphora! Sarraceniaceae!

Woody pitcher plants restricted to tepuis of South America!

Sarracenia! Sarraceniaceae!

pitcher plants restricted to coastal plains of SE U.S.A. with S. purpurea (above) distributed to the north!

Types of Carnivores! Types of Carnivores!

Passive traps - pitfall ! Passive traps - pitfall ! ! !! ! !!

Sarracenia purpurea! Sarraceniaceae! Pitcher plants often have an alluring leaf flap, then downward projecting hairs, then a slippery slope Pitcher plants have unusual drooping 5-merous of wax, and finally a drowning pool. Codine like flowers with peltate style and 5 stigmatic lobes and compounds stupefy the insects before digestive many stamens! enzymes are released.!

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Types of Carnivores! Types of Carnivores!

Passive traps - pitfall ! Passive traps - pitfall ! ! !! ! !!

Flowers have an unusual pollination syndrome Flowers have an unusual pollination syndrome geared for outcrossing! geared for outcrossing!

Types of Carnivores! Types of Carnivores!

Passive traps - pitfall ! Passive traps - pitfall ! ! !! ! !!

Insects are attracted by sight of the “cobra” Darlingtonia (Sarraceniaceae) - the Cobra lily tongue and nectar produced there. Once in the restricted to northern California and Oregon! pitcher, the insects slip into the drowning pool.!

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Types of Carnivores! Types of Carnivores!

Passive traps - pitfall ! Passive traps - pitfall !

Nepenthes is a large genus of pitcher Nepenthes is a large genus of pitcher plants in Asia and a few in African plants in Asia and a few in African rainforests! rainforests!

The pitcher is a modified leaf drip tip, a common feature in rainforest leaves!

Types of Carnivores! Types of Carnivores!

Passive traps - pitfall ! Passive traps - pitfall !

Nepenthes is a large genus of pitcher Brocchinia is one of two genera of plants in Asia and a few in African Bromeliaceae, the pineapple family, rainforests! that are carnivorous. It is the only example of a genus with both The pitcher is a modified leaf drip carnivorous and non-carnivorous tip, a common feature in rainforest species.! leaves!

Catopsis is the only other Like many pitcher plants, Nepenthes carnivorous member of the exudes nectar and a sweet smell! Bromeliaceae!

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Types of Carnivores! Types of Carnivores!

Passive traps - pitfall ! Passive traps - pitfall !

Cephalotus - the Australian pitcher - is so is restricted to the unusual looking that its systematic placement nutrient poor summits of the tepuis. was unknown until recent DNA evidence placed When grown in the greenhouse with it near the family Oxalidaceae or sorrels. ! nitrogen added, the leaves green up and the pitcher opens up.!

Oxalis!

Types of Carnivores! Types of Carnivores!

Passive traps! Passive traps!

! pitfall ! ! pitfall ! ! !Sarraceniaceae - American pitcher plants! ! !Sarraceniaceae - American pitcher plants! ! !Nepenthaceae - Asian pitcher plants! ! !Nepenthaceae - Asian pitcher plants! ! !Bromeliaceae - “pineapple” pitchers! ! !Bromeliaceae - “pineapple” pitchers! ! !Cephalotaceae - Australian pitcher plant! ! !Cephalotaceae - Australian pitcher plant!

! lobster pot! ! lobster pot! ! !Sarraceniaceae (Sarracenia psittacina)! ! !Sarraceniaceae (Sarracenia psittacina)! ! !Lentibulariaceae ()! ! !Lentibulariaceae (Genlisea)!

! flypaper! ! !Byblidaceae - rainbow plant! ! !Droseraceae ()! ! !Dioncophyllaceae! ! !Roridulaceae!

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Types of Carnivores! Types of Carnivores!

Passive traps - flypaper! Active traps! ! !! ! flypaper ! (Byblidaceae) - the rainbow plant - ! !Droseraceae () - sundews! has modified leaves with sticky hairs. ! !Lentibulariaceae () - butterwort! Light hitting the glandular hairs causes a rainbow effect which seems to attract ! steel trap! insects. ! ! !Droseraceae (Dionaea) - Venus fly trap! ! !Droseraceae () - water wheel! However, neither the leaves nor hairs show any movement and the mode of ! mouse trap! carnivory is thus considered passive.! ! !Lentibulariaceae () - bladderwort!

Types of Carnivores! Types of Carnivores!

Active traps - flypaper! Active traps - flypaper! Drosera (sundews) have modified leaves with Pinguicula (butterwort) has modified leaves sticky tentacles. These are alluring, sticky, and with sticky buttery top surfaces. Leaves curl move to further trap the insects.! to assist in capture.!

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Types of Carnivores! Types of Carnivores!

Active traps - steel trap! Active traps - steel trap! Dionaea (Venus fly trap) has modified leaves acting as steel traps. Two trigger hairs must be The Venus fly trap has been the inspiration of touched to snap trap shut. One species, some great films.! endangered, restricted to the Carolina bogs.!

Film kiosk at University of Wisconsin in 1975!

Types of Carnivores! Types of Carnivores!

Active traps - mouse trap! Active traps - mouse trap!

Utricularia (bladderwort) along with However, Utricularia (bladderwort) has Pinguicula (a flypaper trap) belong to the modified underwater structures (bladders) Lentibulariaceae. ! with a trap door that when triggered sucks in aquatic organisms.!

bladder!

Utricularia cornuta! trap door with trigger hairs! Beaked bladderwort!

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the Parasites! the Parasites! There are at least 8 groups of parasites, but some are so reduced and bizarre (even Parasites are plants that gain some or all of their carbon, nutrient and water their DNA is strange) that we do not know where they should be classified.! needs from other living plants (off roots, stems, or leaves).!

Holoparasites -! Non photosynthetic (non-green) plants that are obligate parasites!

Hemiparasites -! Photosynthetic (green) plants that are facultative parasites!

Cuscuta - dodder! Comandra - toadflax!

the Parasites! the Parasites!

Order Santalales - Loranthaceae (mistletoes)! Order Santalales - Santalaceae (sandalwood family)!

The hemi-parasitic sandalwood Most mistletoes are epiphytic (grow on family is largely Old World and branches of other plants). However, most often important wood sources epiphytes are not parasitic as they only (sandalwood and gopher wood).! use the host plant for support.!

Mistletoes are found in both temperate and tropical climates, but most diverse in the tropics.!

Santalanum - sandalwood!

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the Parasites! the Parasites!

Order Santalales - Santalaceae (sandalwood family)! Order Santalales - Balanphoraceae and other fungal mimics!

The hemi-parasitic sandalwood family is largely Old World and often important wood sources Species so reduced and (sandalwood and gopher wood).! so fungus-like, that many have only recently been recognized as flowering plants.! Two genera occur in the Great Lakes region. Comandra is Restricted to dark, wet known to have the greatest tropical forest floors.! number of host plant species.!

Comandra! Geocaulon! toadflax! earthstem!

the Parasites! the Parasites!

Order Rafflesiales - Rafflesiaceae [order is yet to be placed in classification]! Order Rafflesiales - Hydnoraceae [order is yet to be placed in classification]! Holoparasite restricted to vines of the grape family in Paleotropics. Vegetative parts of plant is mycelia- like and within the host. Largest flower in world only emerges from vine.!

Anatomy is so bizarre, many structures seem to have no Very reduced family homology with floral parts.! morphologically with a peculiar southern South America and southern African distribution.!

Rafflesia!

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the Parasites! the Parasites!

Cuscuta - the dodders (Cuscutaceae)! Cuscuta - the dodders (Cuscutaceae)! Dodder vegetative and The family is totally floral features are holo-parasitic and similar to that of the comprises only twining morning glory Cuscuta, the dodders.! family which is not parasitic. ! As twining parasites, they attach to the host stems and penetrate DNA evidence shows into the vascular that Cuscuta is placed in tissue.! the middle of the Convolvulaceae family. It is essentially a twining morning glory that is non-green and parasitic.! Convolvulus - morning glory!

the Parasites! the Parasites!

Scrophulariaceae and Orobanchaceae (order Lamiales)! Scrophulariaceae (order Lamiales)!

The snapdragon (Scrophulariaceae) and the broomrape The snapdragon family contains both non-parasites (e.g., (Orobanchaceae) families have long been considered to be snapdragon) and hemi-parasites (e.g., Indian paintbrush, closely related within the mint order (Lamiales) - along with lousewort, and false foxglove).! the carnivorous Lentibulariaceae (butterwort, bladderworts).!

Indian paintbrush! Scrophulariaceae!

Broomrape! Orobanchaceae!

Castilleja! Pedicularis! Aureolaria!

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the Parasites! the Parasites!

Orobanchaceae (order Lamiales)! Scrophulariaceae and Orobanchaceae (order Lamiales)!

The broomrape family contains only holo-parasites DNA evidence now shows that the hemi-parasites of (e.g., broomrape, beech-drops, and squaw-root) - Scrophulariaceae should be classified with the holo-parasites many with species-specific host relationships.! of Orobanchaceae, and not with other members of the Scrophulariaceae which are not parasitic.!

Indian paintbrush! Scrophulariaceae! Orobanche!

Closely related !!

Broomrape! Orobanchaceae! Epifagus! Conopholis!

the Saprotrophs! the Saprotrophs!

The saprotrophs are non-green plants that form haustoria (stem/root-fungal) Not surprisingly, the most common occurrences of saprotrophs occur in the connections with fungi. The fungi provide water, nutrients, and carbon source for families or closely related families of those plants with strong mycorrhizal the plants. This relationship is often referred to as a parasitic-fungal life style. associations - a common feature of many green, photosynthetic plants.! Most common in low nutrient, acidic, low-light forests.!

Coralorhiza! Voyria! Coral-root! Gentianaceae! Orchidaceae!

2 tropical forest saprotrophs!

Monotropa! Triuris! Indian pipe! Triuridaceae! Monotropaceae/! Ericaceae!

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the Saprotrophs! the Saprotrophs! The blueberry family (Ericaceae) has traditionally been separated from the DNA evidence now shows that both the Pyrolaceae and Monotropaceae are shinleaf family (Pyrolaceae) and the Indian-pipe family (Monotropaceae) independently placed within the Ericaceae. That is, certain members of the because the latter two exhibit increasing dependence on the fungal Ericaceae s.l. (sensu lato - or in the broad sense) are now adapted to the association. The Monotropaceae becoming obligate saprotrophs. ! extreme mycorrhizal dependency.!

Mycorrhizal dependency!

Bear-berry! Bear-berry! Pinesap! Ericaceae! Ericaceae! Monotropaceae!

Shinleaf! Shinleaf! Pyrolaceae! Pinesap! Pyrolaceae! Monotropaceae!

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