Lizbeth Gonzalez

Peer Review Input

All my reviewers noted that I should revise my citations, so I decided to use the number system.

Some citations are very long, so I can see why it would become a distraction for the reader. Two

out of three reviewers mentioned that they liked my use of the common and scientific names of

some of the species I talk about, so I made sure I was consistent with all the species I mentioned.

Keystone Species

A keystone species is an organism whose presence has a large, disproportionate effect on

its environment relative to its abundance (1). Just like a keystone is essential in holding an

architectural arch together, ecosystems can fall apart or look completely different without the

presence of a keystone species( 2 ) .The removal or absence of this species of plant, animal, fungi,

or bacteria might make the remaining organisms unable to survive (2).

History

Intertidal ecologist, Robert Paine,

coined the term ‘keystone’ species

in 1966 after studying tidepools

from the Pacific Northwest of

North America (8). In his

experimental plots, Paine removed

Figure 1. P. ochraceous feeding on M. californianus
the sea star, Pisaster ochraceous,

whose preferred food source is the California mussel, Mytilus californianus. He documented a

shift from diverse algal and invertebrate communities to an assemblage taken over by mussel

strands (8). Paine showed that while the California mussel is a dominant competitor for space in
Lizbeth Gonzalez

the intertidal, P. ochraceous kept the population in check, indirectly allowing other organisms to

coexist (1).

Types of Keystone Species

Ecologists designate keystone species into any of the three main categories: predators,

ecosystem engineers, or mutualists.

Predators

P. ochraceous is an example of a predator, which indirectly affects the number of plants and

other animals in a food web by feeding on a given prey species, thereby keeping their population

sizes controlled. Another example of a predator keystone species are sharks, whose presence

keeps smaller organisms from overgrazing (the process in which too many animals feed on a

given source of vegetation without enough time for plants to recover (3)) on the seagrass beds,

which serve as food and shelter for a broad spectrum of organisms.

Ecosystem Engineers

Species that structurally modify their

environment in a way that creates new

microhabitats or changes existing environments

to meet their own needs are considered

ecosystem engineers (2). The beaver is the

Figure 2. Beavers are an example of an
most distinguishable example of an ecosystem Ecosystem engineer.
engineer. They create beaver ponds by using their large incisor teeth to cut down trees and dam

small waterways (5). The body of water they accumulate serves as a safe space to create a family

lodge with a tunnel-like underwater entrance. Although many of their wolf and bear predators
Lizbeth Gonzalez

have been hunted, beavers still rely heavily on theses lodges, where they store vegetarian food

reserves and nurse their newborns in the winter. Their engineering abilities create such sturdy

beaver lodges that when it floods a new story can be built upon the original lodge (6).

Meanwhile, their dams create habitat for other species like birds and fish, reduce flooding, and

control soil erosion (a natural process in which topsoil is lost to wind, rain, and other forces

including human activity (5,7)).

Mutualists

Two or more species are called mutualists

when they both benefit from their interactions (2).

Bees and flowering plants are just one of the many

examples of mutualistic keystone species. In

collecting their primary food sources, i.e., nectar

Figure 3. Bee pollination is an
and pollen, bees unintentionally spread pollen example of a mutualistic relationship
with flowering plants.
among flowers, increasing the chances of

fertilization and more flower growth (2).

Case Study: Gray Wolves as a Keystone Species

Three to four hundred gray wolves (Canus

lupus) inhabited the Greater Yellowstone

Ecosystem in 1872, when the government

designated the area as a national park (9).

By the end of the nineteenth century, human

settlers had hunted most of their prey,

Figure 4. Grey Wolf (Canus lupus)
Lizbeth Gonzalez

which included bison, elk, deer, and moose (9). As a result, the wolves began hunting livestock

and became a menace for farmers and ranchers (9). Up until 1965, the federal government began

eradicating gray wolves, allowing bounty programs to give up to $50 incentives for wolf capture

(9). Eradication techniques involved trapping, shooting, digging up from their dens, poisoning,

and hunting with dogs until park rangers from Yellowstone national park killed the last two

remaining pups (9).

Without the gray wolf, elk populations increased dramatically, overgrazing several plant

communities (11). Among others plant species, aspen and willow populations began decreasing,

which impacted other wildlife like beavers and songbirds (12). Specifically, the overgrazing of

aspen (Populus tremuloides), caused a rapid decline in the number of seedlings and root sprouts

that grow into trees and saplings (9). The decline in plant species caused habitat loss and affected

stream bank stability, organic matter and fine sediment deposition, regulation of water

temperature by shading, and nutrient cycling (9).

Seven years after the endangered species act was passed by the US congress, gray wolves

were enlisted as an endangered species. To reintroduce them into the Greater Yellowstone

Ecosystem, fourteen Canadian wolves were kept in 1 acre acclimation pens for ten weeks before

being released. Monitoring of wolves since their reintroduction indicated that wolves target elk

over any other prey, coinciding with an increase in willow heights in some areas. Additionally, a

ten-fold increase in beaver populations (13) and a notable increase in songbird populations is

accredited to the restoration of willow trees (14).

Lizbeth Gonzalez

Importance of Keystone Species

Many keystone species provide lots of ecosystem services for humans as well as other

organisms. When organisms are given this title, it allows for wildlife conservationists to

determine where to allocate their efforts. One example of an organization that funds research on

bumble bees, an endangered keystone species, is The Xerces Society for Invertebrate

Conservation. In parts of Britain and the Netherlands, where bees have been lost, declines in

insect pollinated plant abundance have been documented (10). Several high value crops, like

blueberries, cranberries, and clover rely heavily on bumble bees, while greenhouse tomatoes are

exclusively pollinated by bumble bee s ( 10 ) .Additionally, pollination services by insect

pollinators have an economic value of around $3 billion per year in the United States alone, with

the majority being bees (10).

The rapid decline of north American bumble bees is most likely due to habitat loss,

pesticide use, climate change, or the introduction of non-native bumble bee diseases (10). Native

tall grass prairie is considered one of the bumble bees’ habitats, but over 99% of it has been lost

to land conversion in the Midwest (10). Although studies on the effects of pesticides are few,

their results consistently show negative impacts (10). The timber industry in eastern Canada used

pesticides to control for forest insects, resulting in the disappearance of bumble bees and the

decline in blueberry harvests near the sprayed area (10). Furthermore, bumble bees have

precisely timed relationships with plants that will likely be affected by climate change. Scientists

predict that bumble bee habitat range will be forced to shift because of climate change,

diminishing cold adapted species’ range and expanding warm adapted species’ range (10).

Luckily, there are a few things that anyone can do to alleviate some of the challenges that

bumble bees and other endangered keystone species are facing:

Lizbeth Gonzalez

 Planting a diverse assemblage of native plants in your garden can provide shelter, reduce
water usage, and provide food for many native species
 Avoiding the use of insecticides or herbicides to protect flowering plants
 Foster bee nests, which are often found in abandoned rodent burrows, and other shelters
 Reduce mowing in areas where bees might nest
 Learn to identify Native species and report them to a citizen scientist platform (i.e xerces
society or iNaturalist)

.
Lizbeth Gonzalez

Bibliography

1. Freeman, S. Biological science (5th ed., p. 1138). Pearson.

2. Keystone Species. (2022). Retrieved 27 September 2022, from

https://education.nationalgeographic.org/resource/keystone-species/

3. Skoulikidis, N. (2022). Mountainous areas and river systems. Retrieved 27 September

2022, from https://www.sciencedirect.com/topics/earth-and-planetary-sciences/

overgrazing

4. Magazine, H. (2022). No Sea Stars = Mussel Beach | Hakai Magazine. Retrieved 27

September 2022, from https://hakaimagazine.com/videos-visuals/no-sea-stars-mussel-

beach/

5. Build a Beaver Dam (U.S. National Park Service). (2022). Retrieved 27 September 2022,

from https://www.nps.gov/articles/buildabeaverdam.htm

6. Nature - BBC Science Focus Magazine. (2022). Retrieved 27 September 2022, from

https://www.sciencefocus.com/nature/why-do-beavers-build-dams/

7. Mulvihill, K. (2022). Soil Erosion 101 | YourEarth Solutions. Retrieved 27 September

2022, from https://www.yourearth.net/soil-erosion-101/

8. Paine, R. (1966). Food Web Complexity and Species Diversity. The American

Naturalist, 100(910), 65-75. doi: 10.1086/282400

9. Keystone Species | Learn Science at Scitable. (2022). Retrieved 27 September 2022, from

https://www.nature.com/scitable/knowledge/library/keystone-species-15786127/

10. Bumble Bee Conservation. (2022). Retrieved 27 September 2022, from

https://www.xerces.org/bumblebees
Lizbeth Gonzalez

11. Laliberte, A., & Ripple, W. (2004). Range Contractions of North American Carnivores

and Ungulates. Bioscience, 54(2), 123. doi: 10.1641/0006-

3568(2004)054[0123:rconac]2.0.co;2

12. Ripple, W., & Beschta, R. (2004). Wolves, elk, willows, and trophic cascades in the

upper Gallatin Range of Southwestern Montana, USA. Forest Ecology And

Management, 200(1-3), 161-181. doi: 10.1016/j.foreco.2004.06.017

13. The History and Current Status and Distribution of Beavers in Yellowstone National

Park. (2022). Retrieved 27 September 2022, from

http://www.npshistory.com/publications/yell/beaver-history-status.pdf

14. Avian response to willow height growth (Salix spp.) in Yellowstone’s Northern Range.

(2022). Retrieved 27 September 2022, from

https://www.semanticscholar.org/paper/Avian-response-to-willow-height-growth-(Salix-

spp.)-Hansen-Baril/aa06924b681942eed08f0574d6e0b8d918a14f78

15. Magazine, H. (2022). No Sea Stars = Mussel Beach | Hakai Magazine. Retrieved 27

September 2022, from https://hakaimagazine.com/videos-visuals/no-sea-stars-mussel-

beach/ (Starfish and mussel picture)

16. Beavers | Beaver lodge, Beaver, Beaver dam. (2022). Retrieved 27 September 2022, from

https://www.pinterest.com/pin/6614730682658992/