Understanding spatiotemporal patterns in green crab population dynamics is essential for predicting and managing the ecological and economic impacts of this harmful invasive species. This protocol was developed in an effort to create a standardized method for assessing green crab populations in the rocky intertidal zone of the Northwest Atlantic.
Invasive species have caused major disruptions to ecosystems worldwide. The European green crab invaded North America in the 1800s and is considered one of the world’s 100 worst invaders by the IUCN. Observations of spatiotemporal green crab population dynamics are essential for predicting and managing the ecological and economic impacts of this harmful invasive species. This protocol was developed in an effort to create a standardized method for assessing green crab population dynamics in the rocky intertidal zone of New England and Atlantic Canada. The protocol was designed to be accessible to multiple users including researchers, educators, students, and citizen scientists. Although it was designed for surveying crab populations, this protocol is easy to adapt and could be used for any number of intertidal species. The resulting data collected using this protocol has a wide range of uses, including to inform ecological research, conservation efforts, mitigation strategies, and fishery development, as well as for educational outreach purposes.
Biological invasions can potentially disrupt species interactions and ecological processes, and may have far reaching ecological1,2,3 and economic consequences4. The ability to successfully predict, mitigate, and adapt to invasions strongly depends on characterizing spatiotemporal population dynamics5. While a range of tools exist (e.g., population genetics, stable isotopes) and are emerging (e.g., eDNA) for tracking invasive species, traditional in situ monitoring techniques continue to be widely utilized for assessing invasive species distribution and abundance.
The European green crab (Carcinus maenas) is an invasive species that was first detected in North America in 1817 and has successfully invaded ecosystems worldwide6,7. Green crabs have a multitude of negative impacts on local ecosystems, including reducing native bivalve populations through predation8,9, competing with native crustaceans for food and shelter10,11,12 and destruction of eel grass habitat and subsequent changes to fish community structure12,13,14. Compounding these issues is the link between increasing temperature and increasing green crab abundance and/or range expansion15,16, which has had severe ecological and socio-economic consequences in areas such as the Gulf of Maine, where warming is occurring faster than 99% of the world’s other oceans17.
On the eastern seaboard of North America, green crabs range from Virginia to Newfoundland. They are most commonly found on wave-protected shorelines, estuaries, and embayments in depths ranging from the high tide level to 5-6 m18. Their presence in the intertidal zone makes them an ideal marine species for shoreline surveys. The most distinguishing characteristic used to identify green crabs is the pattern of five spines or ‘teeth’ on each side of the eyes and three spines between the eyes (see Appendix 1). Their carapace (dorsal side) is typically a mottled dark green and brown, but ventral color patterns can vary greatly (see Appendix 2).
There are many organizations, researchers, citizen scientist groups, and educators currently conducting green crab population monitoring. However, the lack of a standardized protocol makes it difficult to compare datasets and to ultimately understand green crab populations on both a local and regional scale. This protocol is designed to quantify spatiotemporal population dynamics of green crabs in the rocky intertidal zone in New England and Atlantic Canada. Ideally, the development of a standardized, inexpensive, and easily adaptable survey will promote long-term monitoring efforts by a wide range of users, including researchers, citizen scientists, educators, and students.
Although green crabs are the target species of interest in this protocol, data is also collected for native Jonah and rock crabs (Cancer borealis and Cancer irroratus), as well as the invasive Asian shore crab (Hemigrapsus sanguineus). These are crab species commonly found in the rocky intertidal zone in northern New England, and trends in their population distribution and abundance have ecological and economic significance. An Intertidal Crab Field Guide was developed alongside this protocol to aid in crab identification (Appendix 1) specific to northern New England. A data entry and storage platform called “Intertidal Green Crab Project” was also developed for this protocol using Anecdata19. Anecdata is a free online citizen science platform that provides web-based and mobile solutions for gathering and accessing observations, and provides a user-friendly platform to easily collect, manage, and share data.
1. Timing of survey work
2. Pre-survey preparation
3. Site selection and description
4. Conducting survey
5. Data management and analysis
In 2019, this protocol was used to conduct monthly intertidal green crab surveys at three locations from May-November (Sandy Point, Yarmouth, ME (43°46'17.92"N, 70° 8'45.52"W), Robinhood Cove, Georgetown, ME (43°48'13.80"N, 69°44'50.97"W), and New Meadows River, West Bath, ME (43°51'17.84"N, 69°51'55.20"W)), and at one location from May-August (Damariscotta River, Walpole, ME (43°56'9.42"N, 69°34'52.75")). The data collected indicated wide variations in spatial and temporal green crab population density (Figure 3) and sex ratio (Figure 4), as well as significant differences in cumulative size frequency among sites (e.g., Damariscotta crabs were significantly smaller and Sandy Point crabs were significantly larger than the other populations sampled (Kolmogorov-Smirnov test, p < 0.05 for all comparisons), Figure 5). Shell condition changed seasonally within each site, with peak pre-molt and soft-shell phases for males in the spring and for females in the spring and fall (Figure 6).
The protocol was also used in 2018 and 2019 by Georgetown Central School 3rd and 4th grade students who surveyed the same site in Georgetown, ME each year in October. They observed a shift in the population from being dominated by 5-15 mm CW sized crabs in 2018 to 15-30 mm CW sized crabs in 2019 (Figure 7). They also observed an increase in the percent occurrence of the invasive Asian shore crab from 3.5% in 2018 to 9% in 2019 (Figure 8).
Figure 1: Tidal height zonation. Diagram of tidal height zonation along the shoreline. Surveys are conducted in the low intertidal zone (i.e., the lower third of the intertidal zone that runs parallel to the shoreline). Please click here to view a larger version of this figure.
Figure 2: Defining survey area. Image of a survey site in Yarmouth, Maine with tidal zonation displayed in yellow and survey area in green. Please click here to view a larger version of this figure.
Figure 3: Green crab density. Average monthly density (#/m2) of green crabs at Damariscotta River, New Meadows, Robinhood Cove, and Sandy Point intertidal survey sites. Error bars represent ± 1 standard error. Please click here to view a larger version of this figure.
Figure 4: Green crab sex ratio. Sex ratio by month for green crabs at Damariscotta River, New Meadows River, Robinhood Cove, and Sandy Point intertidal survey sites. Sex ratio is expressed as the percent of the population that is male. The green “0” appearing in July indicates 0% male crabs and 100% female crabs at Robinhood Cove in that month. Blue “NA” indicates no data collected at Damariscotta River from September-November. Please click here to view a larger version of this figure.
Figure 5: Green crab size frequency. Cumulative size frequency of green crabs at Damariscotta River, New Meadows River, Robinhood Cove, and Sandy Point intertidal monitoring sites for all months combined (May-November for Damariscotta River, New Meadows River, Robinhood Cove, and May-August for Damariscotta River). Please click here to view a larger version of this figure.
Figure 6: Shell condition ratio. Monthly shell condition ratio of A) female and B) male green crabs at New Meadows River intertidal monitoring site. Please click here to view a larger version of this figure.
Figure 7: Student-collected size frequency. Size frequency of green crabs surveyed by Georgetown Central School 3rd and 4th grade students in 2018 and 2019. Please click here to view a larger version of this figure.
Figure 8: Species occurrence. Percent occurrence of crab species surveyed by Georgetown Central School 3rd and 4th grade students in A) 2018 and B) 2019. Please click here to view a larger version of this figure.
This protocol describes a survey method for assessing spatial and temporal trends of crab populations in the rocky intertidal zone that is accessible to multiple users including researchers, educators, students, and citizen scientists. The benefits of this protocol include the following: it does not require specialized or expensive equipment, the methodology is approachable for a wide range of skill levels (e.g., 3rd and 4th grade students have successfully used it), and it can easily be adapted to suit the needs of the investigator. In addition, the optional use of Anecdata allows for observations to be recorded directly into a data storage platform and accessed by a broad array of users. While the species of interest here is the invasive European green crab, this protocol could be used for any number of intertidal species. However, it is important to carefully consider unique characteristics and behavior of each species targeted when generalizing this protocol.
Many citizen scientist and education initiatives that conduct shoreline surveys for green crabs do not utilize a standard unit of measurement (e.g., quadrat) or target a specific area of the shoreline (e.g., low intertidal zone), making spatial and temporal comparisons difficult. This protocol utilizes simple techniques to target a standardized area along the shoreline. However, this is also a limitation of the protocol in that more rigorous methods or specialized equipment would yield more exact tidal height measurements within the survey area. The survey area is defined as 100 m of shoreline in the low intertidal, but the site should further be defined as a section of shoreline delimited by natural or manmade boundaries that are easy to distinguish, or by a minimum boundary distance of 300 m from the survey area. This is an important delineation if investigators are considering surveying multiple sites in close proximity.
The protocol also includes collecting detailed measurements that can be useful for researchers and managers. It is important to note that not all measurements will be appropriate for all users, such as identifying pre-molt green crabs, which takes a considerable amount of training. Furthermore, sex identification is only recommended for crabs ≥ 10 mm CW, as the sex of smaller juveniles can be difficult to determine. Although it is not part of the protocol, it is also recommended that citizen scientists, students, educators, and others who may not be familiar with the target species utilize a digital camera to document uncertainties and allow for troubleshooting outside of the survey timeframe or to upload to the Anecdata platform for review.
As ocean temperatures continue to warm23,24 and become more favorable for green crab populations in the Gulf of Maine and Atlantic Canada15,16, assessing their distribution and abundance will be critical for timely mitigation and adaptation strategies. For example, recent efforts to develop a soft-shell green crab fishery that would allow coastal communities to benefit from green crabs have relied on population monitoring data to determine when and where to target pre-molt and soft-shell crabs21,22. Bounty programs and other removal efforts may also benefit from distribution and abundance data that can aid in determining how best to allocate time and resources. Similarly, municipal shellfish management programs often conduct yearly conservation activities such as replenishing local mudflats with hatchery raised clam seed and local trends of green crab distribution and abundance could inform best practices for these conservation measures.
Finally, this protocol has proven to be an effective outreach and education tool for students and citizen scientists. In addition to the 3rd and 4th grade students who have been using the protocol for several years, it has also been used with undergraduate students from the University of Southern Maine, and community service volunteers from Idexx Laboratories. Furthermore, an adapted version of this protocol is now being used by the Gulf of Maine Research Institute’s Vital Signs Program, which partners with Maine students and citizen scientists to monitor invasive species. Ultimately, the data collected using this protocol has a wide range of uses, including to inform ecological research, conservation efforts, mitigation strategies, and fishery development, as well as for educational outreach purposes.
The authors have nothing to disclose.
I would like to acknowledge the many researchers, students, teachers, and citizen scientists who have tested and helped me improve this protocol over the past several years: Dr. Gabriela Bradt, Dr. Robert Steneck, Erica Ferrelli, Ethel Wilkerson, Susan Ayers and the Georgetown Central School 3rd and 4th grade students, Julie Upham and the West Bath Elementary 4th grade students, the GMRI Vital Signs team, and the Idexx Laboratories Community Service Volunteers. I thank Anne Hayden, Caitlin Cleaver, and Hannah Webber for providing comments and suggestions on this manuscript. I thank the following funding sources for supporting the development and execution of this protocol: NOAA Saltonstall-Kennedy Grant Program (Grant #NA18NMF4270194), Maine Sea Grant, Robert and Patricia Switzer Foundation, and the Georgetown Island Education Fund. Finally, this manuscript was greatly improved thanks to comments and edits provided by three anonymous reviewers.
1 m2 PVC quadrat (1/2" PVC) | Any hardware/home improvement store | PVC can be sourced at any hardware/home improvement store and cut into 1m lengths to form quadrat (4 1/2" PVC elbows will also be needed to connect 1 m lengths into square) | |
1/2" rebar | Home depot | 5152 | *optional (for marking low intertidal area) |
40 m Fiberglass Transect Tape | Grainger | 3LJX1 | |
5 gal bucket | Home depot | 05GLHD2 | |
Ade Advanced Optics Salinity Refractometer | Amazon | *optional | |
Clip board | Any office supply store or Amazon | ||
Uei Waterproof Digital Thermometer | Amazon | *optional | |
Vernier calipers | Bel-Art | Many companies make calipers, however our preferred brand is Bel-Art which can be sourced on Amazon |