The Origin and Continous Evolution of The Real World’s Alien Plants:

Carnivorous Plant Evolution as a Model of

Homology and Analogy

A Synthesis Paper Presented to the Faculty of

Philippine Science High School CALABARZON Region Campus

Submitted by:

Alexa Ysabel D. Liwag

BIOLOGY 3A

11 Vega

Micah Teresa Elayda

Biology III Adviser

 Plants are commonly known for their immobility and passiveness. This makes

human beings think of plants as generally non-dangerous to them despite the self-

defense mechanisms these plants posess. Although the general impression of men

on plants is that they are unfrightening, “The Savage Garden: Cultivating

Carnivorous Plants” by Peter D’Amato (1998) suggests that humans have an

unconscious fear of pretty, pulpy, and passive plants. D’Amato states that this is

evident in pop culture, esspecially in science-fiction horror films such as “The Day of

the Triffids”, “The Thing”, or even the cult classic, “Little Shop of Horrors”. That being

said, the existence of carnivorous plants is recognized as bizarre and almost

peculiar, with scientists like Charles Darwin (1875) choosing to pursue studies

involving these plants, specifically those insectivorous.

Carnivorous plants can be used as models to decribe how homology and

analogy, because of their evolutionary pattern and descent. There are homologous

traits in various carnivorous plants, which are analogous to other carnivorous plants

which evolved in different locations across the planet. There have been similar traits

among organisms which have been taxonomically classified as closely related that

are also present in plants that are not considered as related to the subject

organisms.

The Carnivorous Plant Nursery categorizes carnivorous plants according to

the possible types of traps the plant may use to capture prey. Plants with active

traps, such as Dionaea muscipula, or the venus fly trap, tend to consume small

insects through a trigger release system, which involves the plant’s reaction to

stimuli. Carnivorous plants with active traps may be found in terrestrial or aquatic

settings, with bladderworts or Utricularia sp. as a primary example. On the other

hand, plants with semi-active traps make use of secretions of sticky substances to
capture prey on leaf surfaces. The leaves then roll over to cover and digest the prey.

This mechanism in evident in sundews (Drosera sp.) and butterworts (Pinguicula

sp.). Plants may also make use of the passive trap, wherein decorative, tubular

leaves filled with digestive enzymes trap its prey by opening its hood and allowing

them to fall into the tubular leaf. Evident examples of these include pitcher plants

such as Sarracaenia and Nepenthes. These may be useful especially in determining

how each trap might have evolved as products of analogy.

The origin of carnivorous plants is still uncertain to date. Givnish (2015)

provides evidence that the earliest record of carnivorous plants, Roridula sp., dates

up to 35 to 47 million years ago. A fossil record of a fragment of a tentacled leaf

preserved in amber and found around the Baltic was linked to be an earlier species

of the carnivorous plants, Roridulaceae. These have also been found near the region

and in other Mediterranian areas, but are now more common in South Africa which

implies that the group was far more widespread. Roridulaceae are shrubs with

insect-trapping leaves whose carnivory causes an acceleration in photosynthesis.

The early Roridulaceae ancestor is one of the few undoubted fossils of

carnivorous plants. But, Givnish also describes another possible common ancestor,

Archaeamphora sp., found from Chinese sediments which date up to 112 million

years ago. It was stated in his report how there is a possibility that the

Archaeamphora sp. may not even be a close and direct ancestor of any known

carnivorous plants. Its unusual leaves may not have been traps (Brittnacher, 2013)

and the “seeds” found in the fossil may have been insect eggs (Hermanová &

Kvacek, 2010).
 Evidence from Thorogood (2017) discusses how pitcher plants and its

varieties are a product of convergent evolution. After observing the different trap

surfaces of sifferent types of pitcher plants, Thorogood suggests that “biomechanical

constraints may have driven the evolution of trap microstructures”. This phenomenon

is observed in the independent evolution of two morphologically different

superhydrophilic trap surfaces – that of Nepenthes and Sarracenia – in unrelated

plant orders. This makes it a striking example of convergent evolution, specifically,

functional convergence.

This functional convergence is also detailed in the “morphology of multicellular

glands and associated chemistry in non-carnivorous families sister to the carnivorous

Caryophyllales” (Renner and Specht, 2013). The analyses of multiple carnivorous

and non-carnivorous plants has lead Renner and Specht to conclude that some

carnivorous plants (e.g. Dionaea, Aldrovanda, Nepenthes) have evolved from non-

carnivorous plant outgroups (e.g. Plumbaginaceae) which do not appear to have

associated vasculature in glands. Moreover, the analyses also lead the scientists to

another conclusion wherein it is suggested that “Ancestral state reconstruction

suggests that sessile glands without vasculature are likely the ancestral state for the

carnivorous Caryophyllales, while stalked and pitted glands were acquired

secondarily and independently by the non-carnivorous sister families and by various

lineages of carnivores”.

In contrast to convergent evolution and analogy, divergent evolution and

applications on homology have also been evident in carnivorous plants. Thorogood

points out the morphological trap characteristics (e.g. size, shape, peristome

geometry, presence and location of wax crystal layers) and physiological traits (e.g.
attractive volatiles, nectar and pitcher fluid composition) of Nepenthes and how they

vary across the genus. He also points out how carnivorous plants rely on the

nutrients of their prey to thrive, and “strong selection pressures should act on traps to

maximize their prey intake”.

There are also evidences of divergent evolution happening at a molecular

level. Jobson (2004) demonstrates that “a dramatic molecular evolutionary rate

increase in subunit I of cytochrome c oxidase (COX) from an active-trapping lineage

of carnivorous plants is caused by positive Darwinian selection”. The evolution in

COX I of bladderworts (Utricularia) is responsible for the organisms’ radical

morphological evolution which may also be due to strong selection pressures.

Stemming everything down, these occurences may be attributed to the living

conditions in which the plants are accustomed to. The instance of the

aforementioned claim may be caused by the inadequacy of nutritional requirements

found in the habitat, as well as the availability of insects and other small digestible

organisms. Ellison and Gotelli (2001) state how the independent evolutions of

carnivorous plants are adaptations to the low-nutrient, waterlogged habitats they

grow in. In addition, they also observe how “reliance on insect-derived nutrients

increases as carnivorous structures become more elaborate (from sticky leaves of

Drosera spp. to the >1 m tall pitchers of the cobra lily, Darlingtonia californica)” which

may also be due to adaptation.

In conclusion, carnivorous plants show a very interesting proposed pattern of

descent, springing from multiple different non-carnivorous plants, and further

evolving from one another. A lack of nutrient in a plants’ areas of growth is likely to
have caused analogous groups to start evolving to gather nutrition from insects and

other small prey, with evidences dating back up to 35-47 million years ago. Because

of strong selection pressures, carnivorous plants tend to have radical morphological

and molecular evolutions. Therefore, carnivorous plants are great examples of

organisms with a traced and definite pattern of analogy and homology, and can be

used as models of this.

References

Brittnacher J. (2013) Phylogeny and biogeography of the Sarraceniaceae. Carniv

Plant Newsletter 42(3):99–106

D'Amato, P. (1998). The Savage Garden: Cultivating Carnivorous Plants.

Darwin, C. (1875). Insectivorous Plants.

Ellison, A. M., & Gotelli, N. J. (2001). Evolutionary ecology of carnivorous plants.

Trends in Ecology & Evolution, 16(11), 623-629. doi:10.1016/s0169-

5347(01)02269-8

Givnish, T. J. (2014). New evidence on the origin of carnivorous plants. Proceedings

of the National Academy of Sciences, 112(1), 10-11.

doi:10.1073/pnas.1422278112

Hermanová Z., Kvacek J. (2010) Late Cretaceous Palaeoaldrovanda, not seeds of a

carnivorous plant but eggs of an insect. J Natur Hist Mus (Prague)

179(9):105–118

Jobson, R. W., Nielsen, R., Laakkonen, L., Wikstrom, M., & Albert, V. A. (2004).

Adaptive evolution of cytochrome c oxidase: Infrastructure for a carnivorous

plant radiation. Proceedings of the National Academy of Sciences, 101(52),

18064-18068. doi:10.1073/pnas.0408092101

Renner, T., & Specht, C. D. (2013). Inside the trap: gland morphologies, digestive

enzymes, and the evolution of plant carnivory in the Caryophyllales. Current

Opinion in Plant Biology, 16(4), 436-442. doi:10.1016/j.pbi.2013.06.009

Thorogood, C. J., Bauer, U., & Hiscock, S. J. (2017). Convergent and divergent

evolution in carnivorous pitcher plant traps. New Phytologist, 217(3), 1035-

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Types of Carnivorous Plants. (2018, May 7). Retrieved from

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information/types-of-carnivorous-plants