The Wonderful World of Plantlets, Bulbils, Cormlets, Tubercles, and Gemmae

Probably the most well known strategy that plants have for dispersal is by way of seeds. Seeds are plants in embryo, and new generations of plants are born when seeds, released from their parent plants, find suitable locations to germinate. But one of the most amazing things about plants in general is that they have the ability to reproduce in a variety of different ways, and many plant species are not limited to seeds as their only means of dispersal. A paper by Scott Zona and Cody Coyotee Howard, published in Flora (February 2022), introduces us to the intriguing world of aerial vegetative diaspores – just one of the many ways that plants have to get around.

A diaspore is a plant structure that facilitates dispersal. Seeds are diaspores, as are spores, which are produced by non-seed bearing plants like mosses and ferns. If you’ve ever planted bulbs, you’ve handled another type of diaspore. Bulbs and corms, which many spring flowering plants are grown from, form little offshoots called bulblets and cormels that, when detached from their parent structure, can grow into new individuals. These vegetative diaspores are produced below ground. Aerial vegetative diaspores, on the other hand, are formed on above ground plant parts. This clunky term encompasses a number of different structures that are often simply called bulbils, which Zona and Howard explain is used as “a catch-all term that obscures their morphological identity.”

Compiling a list of plant species that feature aerial vegetative diaspores is a difficult task when plant descriptions from various sources use a broad selection of terminology for the same or similar plant parts. To help complete this task, Zona and Howard defined five distinct types of aerial vegetative diaspores – plantlets, bulbils, cormlets, tubercles, and gemmae – and came up with a list of 252 taxa that are known to feature at least one of these structures.

plantlets on the leaf margin of Kalanchoe daigremontiana (wikimedia commons; Aurélien Mora)

Plantlets are miniature plants attached to another plant. Once mature, they have clearly visible leaves, stems, and roots (or root initials) and are non-dormant, meaning they are ready to grow on their own as soon as they’re given the opportunity. The tiny plants borne along the margins of the leaves of mother of thousands (Kalanchoe daigremontiana) is a great example of a plantlet.

A bulbil consists of a shortened stem surrounded by scale leaves modified for food and water storage. Sometimes root initials are visible at the base of the bulbil. Bulbils remain dormant until they are dispersed and conditions are suitable for growth. When bulbils start growing but remain attached to the plant, they become a plantlet. A good example of a bulbil can be found on bulbous bluegrass (Poa bulbosa).

Cormlets are comprised of stem tissue and, like plantlets and bulbils, have a single axis of polarity. They have highly reduced scale leaves and are dormant at dispersal. Bulbil bugle lily (Watsonia meriana), despite its misleading common name, is a good example of a plant that produces cormlets.

Tubercles are made up of swollen stem tissue and, like tubers (their underground counterparts), have multiple shoot buds and multiple axes of polarity (meaning there is no right side up like there is in plantlets, bulbils, and cormlets). They lack scale leaves and are dormant at dispersal. Air potato (Dioscorea bulbifera) is an example of a tubercle-producing plant. As you might guess from the common name, potato-like structures are produced aerially on this vining plant that was introduced to North America from Africa and is now invasive in Florida.

A gemma is a tiny cluster of undifferentiated cells. Gemmae are non-dormant and lack polarity. They are the smallest and least common form of aerial vegetative diaspore and can be found on Drosera pygmaea, a species of sundew native to parts of Australia and New Zealand.

Drosera pygmaea (wikimedia commons; Björn S…)

Zona and Howard’s list of plants with aerial vegetative diaspores is the most comprehensive list to date – although it is undoubtedly and understandably missing some – and includes representatives from 42 plant families and 21 plant orders. Plantlets are the most common form of aerial vegetative diaspore at 116 taxa, with bulbils coming in second at 72. Cormlets and tubercles are less common, with 25 and 16 taxa respectively. Their paper includes the full list and offers further information about many of the species listed. It’s worth taking time to explore and is a valuable resource for anyone interested in the topic. In addition, their discussion section highlights a number of questions that warrant further investigation.

Questions surrounding reproductive strategies and the dispersal of aerial vegetative diaspores are particularly interesting. Because these structures are vegetative, they are essentially clones of the parent plant, meaning there is no genetic mixing as occurs when seeds are produced. This can be an advantage when sexual reproduction isn’t possible due to lack of pollinators, environmental restrictions, or chromosomal/polyploidy anomalies. It also assures that new individuals are pre-adapted to the site, and it can help a species colonize an area quickly. This ability to rapidly colonize explains why several of the species on Kona and Howard’s list are known to be invasive in parts of the world outside of their native range.

A species that produces both seeds and aerial vegetative diaspores may have an advantage when it comes to dispersal since both types of diaspores have their strengths. Seeds can remain dormant in the soil and are likely to persist in the environment longer than vegetative diaspores, but vegetative diaspores can be produced without relying on pollinators and can establish new individuals quickly. The modes of dispersal between the two can also vary. Seeds can be dispersed by wind or carried away by animals, while vegetative diaspores often rely solely on gravity to get around. One exception is hitchhiker elephant ear (Remusatia vivipara), whose bulbils are equipped with tiny hooks that cling to animal fur and are transported in a similar manner to burs.

hooked bulbils of hitchhiker elephant ear (Remusatia vivipara) (wikimedia commons; Dinesh Valke)

When the advantages of aerial vegetative diaspores are considered, it is a wonder that we don’t see them more often. Many plants can be easily propagated by taking stem, leaf, and/or root cuttings and placing them in conditions that favor adventitious root and shoot growth. This may suggest that dormant genetic pathways for producing vegetative diaspores exist in most plants. Or maybe not. Genetic studies of species that feature these structures are needed in order to understand why they are found in some species and not others. Kona and Howard leave us with a slew of research questions like this, and it’s a topic I’ll continue to check in on.

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