Horizons
Volume 5
Issue 1
Article 25
12-18-2020
Size and Population Dynamics of Native Ghost Crabs, Ocypode
spp. In Response to an Invasive Ant Population in a Native Wildlife
Refuge, Oʻahu,
O ahu, Hawaiʻi
Hawai i
Haley Norma Anne Chasin
University of Hawaiʻi at Mānoa
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Part of the Marine Biology Commons
Recommended Citation
Chasin, Haley Norma Anne (2020) "Size and Population Dynamics of Native Ghost Crabs, Ocypode spp. In
Response to an Invasive Ant Population in a Native Wildlife Refuge, Oʻahu, Hawaiʻi," Horizons: Vol. 5 : Iss.
1 , Article 25.
Available at: https://kahualike.manoa.hawaii.edu/horizons/vol5/iss1/25
This Article is brought to you for free and open access by Kahualike. It has been accepted for inclusion in Horizons
by an authorized editor of Kahualike. For more information, please contact daniel20@hawaii.edu.
Size and Population Dynamics of Native Ghost Crabs,
Ocypode spp., in Response to an Invasive Ant Population
in a Native Wildlife Refuge, O‘ahu, Hawai‘i
Haley Norma Anne Chasin
Biology 400 (Marine Option Program Skill Project) (OPIHI Internship)
Mentor: Patrick Nichols, Joanna Philippoff, and Dr. Cynthia Hunter
Invasive species are harmful to ocean environments especially fragile ecosystems, like
intertidal island environments. Ant populations are among the most aggressive invaders.
Ants have been known to cause individual harm to endangered birds, plants and other
arthropods in Hawai‘i. The yellow crazy ant (YCA), Anoplolepis gracilipes, is one
of the worst invasive ants as it can form supercolonies and spew out formic acid. This
study investigates the effect of YCA on the abundance, sizes and distribution of ghost
crabs, Ocypode spp., at James Campbell National Wildlife Refuge (JCNWR), a native
wildlife restoration area. Ghost crab’s burrows were counted and measured (a proxy for
crab size), in order to see if there is a relationship between ghost crab size/number to the
density of ants (both invasive YCA and others) present. Ghost crabs are vital to sandy
intertidal regions as they are the link to land and water ecosystems and are opportunistic.
This study is important because it tell us how the ants are affecting the ghost crab populations and the ecosystem in general. This study can be used in future research to determine
conservation techniques in order to control invasive ant populations, like the YCA.
traffic from humans and risk of desiccation (Przeslawski 2004;
Nordlund et al. 2014; Pombo et al. 2017). As temperature increases or decreases it can have an effect on the embryonic
development which may reflect how a species in the intertidal
zone is distributed (Przeslowski 2004). Organisms are affected by both abiotic and biotic factors, some of which affect species in the intertidal zone more so than others (Brancho et al.
2010). Sandy intertidal zones are characterized by variations in
Introduction
Intertidal zones are areas between the ocean and land ecosystems that encompasses a more sensitive to change area compared to other terrestrial environments (Ansari et al. 2014).
The challenges of the intertidal zone come from its many hardships including, temperature extremes, wave intensity, foot
This has been an amazing opportunity to show how an invasive species changes the ecology of an
ecosystem. I learned more about research in the field and learn more about crabs and ants. I also
learned how to do better data analysis and come up with my own research question, hypothesis and
other scientific method processes. It was a great way to meet new people and develop a network of
friends in the community. Thank you to OPIHI (Our Project in Hawaii’s Intertidal) and NSF (National Science Foundation).
Horizons, Vol. 5, 2020, pp. 124–132
Copyright © 2020 by the University of Hawaiʻi at Mānoa
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Chasin
Size & Population Dynamics of Native Ghost Crabs in a Native Wildlife Refuge
sand grain size, sun exposure time, differences in the swash
zone width, temperature variations and inter/external species
interactions (Zardi et al. 2007; Przesclowski 2004). In the sandy intertidal zone species like the ghost crab (Ocypode spp.) are
affected by wind and human traffic as well as the other factors
mentioned, which may reflect fragility in their species which
could allow for better management tactics as well (Chan et al.
2006; Brancho et al. 2010).
Included in the biotic factors, is organismal competition
which may dictate the spatial distribution of the intertidal zone
(Connell and Gillanders 2007; Nordlund et al. 2014). For the
genus, Ocypode, some spatial distributions are based on gender
differences, where the males are found in the upper intertidal/dry areas and females/juveniles are found in the vegetation zone (Fellows 1975), though some indicate that juveniles
are found more towards the wet swash zone due to desiccation awareness, which may be an indication of differences in
burrow diameters as adult males tend to leave behind larger
burrows (Chan et al. 2006). It may also indicate gender-based
pyramidal formation, an important sign of colony formation
where males build burrows with pyramid-like sand pellet behind them, in order to attract females (Trott 1998; Fellows
1975). Lighter (1977) describes how Ocypode used all three
kinds of spatial dispersal patterns, randomness, aggregation
and uniformity based on sex and crowding distances. He used
a similar method to this study to characterize the sizes of crabs,
by measuring their hole diameter (Lighter 1977; Pombo et al.
2007). Species distribution and spatial variation can change
based on different biotic and abiotic factors and can be based
on species interactions as well (Brancho et al. 2010).
Species interactions can be beneficial, harmful, or neither
(Connell and Gillanders 2007). Sometimes species that are
more dominant ‘take-over’ or invade an area. If invaders are unmanageable, they can compromise the spatial distribution and/
or densities of Ocypode, interfering with their natural processes
(Bergstrom et al. 2009; Dejean et al. 2010). Due to the fragility of island-like environments, allowing many generations for
species to evolve and co-exist together, invasive species are a
detriment to these environments, like in Hawaiʻi (Reimer 2004;
Bergstrom et al. 2009). This phenomenon threatens Hawaiian
ecosystems as there are 30% endangered and rare species and
more than 1000 native Hawaiian species already extinct (Allen 2000). Therefore, it is vital that conservation efforts go into
place in order to protect the remaining species present.
Many ant species including Solenopsis invicta (Red Fire Ant),
Linepithema humile (Argentina Ant), Pheidole megacephala (Big
Headed Ant), Wasmannia auropunctata (Electric Ant) and Anoplolepis gracilipes (Yellow Crazy Ant), which have been known to
cause detrimental ecological effects around the world (Wetterer
2005; Invasive ants . . . c2018). A successful invader has low intraspecific aggression (unicolonial nests), high interspecific aggression and mutualistic relationships (Kirschenbaum 2007; Dejean
et al. 2010). Ants are invasive due to their ability to form high
125
densities or supercolonies causing ‘invasional meltdowns’ of species in the area (Abbott 2005). There are about 45 different ant
species in Hawaiʻi all of which are non-native and invasive with
about half found in urban, agricultural and natural environments
(Reimer 2004; Kirschenbaum 2007). The invasive ant species,
Anoplolepis gracilipes or the yellow crazy ant, originated from West
Africa, India or China. They are found mostly in tropical, moist
lowlands environments but not found in areas above 1200m or
in arid environments. A. gracilipes has been known to have affects
on nesting birds and native invertebrates in Seychelles as well as
endemic crabs found on Christmas Island, Australia (Wetterer
2005). A. gracilipes forms a mosaic of high-density supercolonies
that can last for more than 24 hours a day, year-round as well as
their ability to spit out formic acid into the eyes or weak areas of
birds and crabs (Abbott 2005; Gül 2017). On Christmas Island
they have had the ability to kill the native red land crabs within
24 hours (Abbott 2005), which makes YCA a substantial threat
to native Hawaiian crabs.
Ocypode spp. is an ecologically important part of the sandy
intertidal food webs. They are highly mobile and feed on a variety
of foods which making them an important balance in the food
web ecosystem. Ocypode spp. are also a key link between inland
ecosystems and the marine environment as they are the predator/
prey item for higher level consumers like birds, as well as some
of their food is obtained through terrestrial resources (Rae et al.
2019). Ghost crabs are also an important part of bioindication of
oil spills and other such chemical contamination (Ghost Crabs . . .
unknown copy-right date). Hawai‘i has two ghost crab species,
Ocypode ceratophthalmus (horn-eyed ghost crab) and Ocypode laevis or Ocypode pallidula (pallid ghost crab), The horn-eyed ghost
crab is found all over the world whereas the pallid ghost crab is
found on different islands throughout the pacific (Lighter 1977).
Invasive species have become one of the causes for the 91
(54%) extinctions of the 680 extinct species around the world
of which 34 (20%) of the extinctions, invasive species were the
only cause of extinction (Clavero and Garcia-Berthou 2005).
Many species have become introduced, since the first arrival
of humans over 1,000 years ago and many have also become
extinct including the important land-dwelling crab, Geoprapsus
severnsi, once common on the Hawaiian islands. The loss of the
land-dwelling crab allowed for there to be an ecological imbalance, that was once harmonious, in an ecosystem (Gonzaga
2011). This emphasizes the importance of understanding invasive species ecology as ecosystems do not want to allow more
species to go extinct as well as disrupt the normal harmony of
the ecosystem present. The effect of invasive species could influence the environment and species around them in the refuge.
Therefore, this study was focused on James Campbell
National Wildlife Refuge (JCNWR) where a restoration of four
native water birds has taken place (Hunt and Eric 2000). The
focus is to examine how the invasive ant population of A. gracilipes and other ants affects the native crab population, Ocypode spp. and the vegetation zone at JCNWR.
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126
The main objectives of this study are first to investigate
population changes over time (comparing 2019 and 2020 data
sampling seasons), second, to compare crab densities in different zones of the beach (sand vs. vegetation), and third, to determine whether or not invasive ant species are affecting crab
populations as well as the vegetation within JCNWR. Understanding the ant populations mutualistic relationship will allow
us to better monitor them and how they influence the ecosystem. We may also be able to even eradicate them from the reserve to help the species in that area and hopefully limit the
number of extinctions or catch extinctions before they occur.
Materials and Methods
Site Mapping
Our site is located on the North Shore of Oʻahu from Kahuku
to James Campbell National Wildlife Refuge (JCNWR) to Turtle Bay (Fig. 1). The site is on a sandy and rocky intertidal zone
in-front of a protected reef which allows there to be smaller
waves on the beach front. Tides were looked up using computer data bases (https://www.surf-forecast.com/breaks/Turtle
-Bay/tides/latest) and start time was set at 0830–0900.
In order to map out the different transect locations or
sample areas we walked along the beach from Kahuku to Turtle Bay and used the Global Position System (GPS) to pinpoint
the locations. We walked along the beach and found 12 sites
that were large enough to have two transects in the sandy areas
(2x12=24) and plotted them on Google Earth with the GPS coordinates (Fig. 1). Lines mark the area of the reserve boundary.
Vol. 5, Fall 2020
We wanted each area to have a patch of sand in the water where
the mole crabs survey could be conducted as well as enough
sand for the ghost crab surveys. Only 22 of the 24 surveys were
conducted due to the COVID-19 pandemic.
Ants Surveys
For the ants at each transect point (marked by GPS) we placed
a container with a smear of peanut butter and honey on the
sides of each container and SPAM at the bottom in order to
attract the ants (Fraiola K., pers. comm.). We placed the container in a shady area at each transect point with a marker for
the location away from the wind so that minimal sand went
into the container. Location (GPS point), date and time in/out
were put onto the container. We left the ants out for 30 minutes to 2 hours and upon collection, capped the bottles. After
the surveying day was done the bottles were placed in a freezer and kept frozen until counting. After freezing, we counted
how many yellow crazy ants were present using a dissecting
microscope; we also counted the number of other ants present
but did not classify them to species (Fig. 2). Due to the global
SARS-CoV-2 pandemic, ants were not able to be placed under
a microscope towards the end of the survey. Ants were visual
identified with the naked eye, so our ant identification may
not be as accurate therefore classification of the ant species
was called other or the total number of ants in the container
was used.
Ghost Crab
Ghost crabs are in the subphylum Crustacea and the genus
Ocypode. They are characterized by their white almost ghost
like appearance in addition to their burrowing behavior
Figure 1 Location (GPS points 1-24) of James Campbell National Wildlife Refuge (JCNWR) on the island of O‘ahu, Hawai‘i. Each
site was found by walking along the 6.4 Kilometers beach finding
12 sandy zones and dividing the zones into 2 for a total of 24
sites. Rocky intertidal areas in the ocean were not surveyed. Only
22 of the 24 GPS points were actually surveyed due to COVID-19.
Figure 2 Containers to collect the ants were placed in shaded
areas away from the wind for 30 minutes to 2 hours. Each container was capped and placed in a freezer until time for counting. Samples were placed under a dissection microscope to
identify if they were yellow crazy ants on the left or other ant on
the right (Pharaoh). Due to global SARS-CoV-2 pandemic, microscopes were not available towards the end of the surveying
and samples were visually identified with the naked eye.
Chasin
Size & Population Dynamics of Native Ghost Crabs in a Native Wildlife Refuge
(Pearse and Vicki 1987). Lighter (1977) used a similar method to this study to characterize the sizes of crabs, by measuring their hole’s diameter (also Pombo et al. 2007). Ocypode
make large burrows in order to defend the space surrounding their burrows (Fellows 1975). At each GPS point starting
from the swash zone, where the tidal line ends, we counted
the number of ghost crabs. When we counted the number of
ghost crabs we also looked at the sizes of their holes. Hole
sizes were measured by measuring the diameter of the shaft
of each burrow. If we could not see into the hole or there
was no darkness the hole was not counted (had to be a fresh
hole). At the swash zone three people lined up side by side
with arms spread out, we walked up the beach counted and
measured the hole diameter using a ruler until we reached
the vegetation zone, noticing what type of vegetation and
continued till the last hole was observed (Fig. 3). In the vegetation zone some of the holes could have been made by the
vegetation and wind. Measurement of the distance from the
swash zone to the last hole were taken using a transect (measured in meters to the nearest decimal point). Observations
of wind, surf and tides were taken into account along with
vegetation type.
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Results
The average number of crabs in 2019 (n= 1065; 39.444±20.363)
was not significantly different from the total number of crabs
in 2020 (n=554; 30.77778±16.491) Therefore we accepted the
null hypothesis (T-test; df=41, p=0.12406, Fig. 4).
The average number of crabs in the sand in 2019 (n=557;
20.6292963±14.6633) was not significantly different from
the average number of crabs in the sand in 2020 (n=404;
22.4444±16.8531). Therefore we accepted the null hypothesis
(F-test, df(2019)=26 and df(2020)=17; p=0.254710648; T-test;
df=33, p=0.711939178, Fig. 5).
The average number of crabs in the vegetation zone
from 2019 (n=532; 19.7037±17.0064) was significantly higher
than the average number of crabs in the vegetation zone from
2020 (n=150;8.3333±7.1702) therefore we rejected the null
Data Analysis
We determined the relative density of yellow crazy ants to other
ants over an area using a correlation analysis, density of ants
was measured in ants per second. For the average number of
crabs we used a T-test. For the ghost crabs we used density
of ghost crabs as the number of crabs per meter squared. We
compared the yellow crazy ants to ghost crab hole diameter,
number and density using correlation tests. For comparing
vegetation and abundance/hole diameter of the crabs to the
presence/absence of ants we used the program R (GLM) (Data
Analysis credit Patrick Nichols).
Figure 3 On the bottom right shows the holes that we measured using a ruler and on the left shows a live ghost crab. The
hole diameter corresponds to the size of the ghost crabs (Lighter 1977; Pombo et al. 2007).
Figure 4 Average total number of ghost crabs in the vegetation and sand zones combined from 2019 (n=1065 crabs;
39.444±20.363 crabs, orange) to 2020 (n=554; 30.77778±16.491
crabs, purple). The numbers of crabs total from 2019 to 2020
were not significantly different therefore the null hypothesis was not rejected (F-test, df(2019)=26 and df(2020)=17;
p=0.184777246; T-test; df=41, p=0.12406).
Figure 5 Average number of crabs in the sand from 2019
(n=557; 20.6296293±14.6633, orange) and the average number
of crabs in the sand from 2020 (n=404; 24.4444±16.3782, purple). The average numbers of crabs total from 2019 to 2020
were not significantly different therefore the null hypothesis was
accepted (F-test, df(2019)=26 and df(2020)=17; p=0.254710648;
T-test; df=33, p=o.711939178).
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Horizons
hypothesis (F-test, df(2019)=26 and df(2020)=17; p=0.00027;
T-test; df=38, p=0.003766, Fig. 6).
Data from 2020 season comparing the sand zone
(n=404; 24.4444±16.3782) and the vegetation zone (n=150;
8.33333±7.17019977) showed that there was a significant difference (F-test; df=17, p=0.0005; T-test; df=23; p=0.003373899,
Fig. 7) between the two zones, therefore we rejected the null
hypothesis.
There was not a significant difference between the number of crabs for the 2019 season in the vegetation zone (n=
532; 19.703704±17.006367) compared to the sand (n=557;
20.6296±14.6633), therefore we accepted the null hypothesis
(F-test; df=26, p=0.227438875; T-test; df=27, p=0.831197, Fig. 8).
The density of ghost crabs also did not change in response
to the total amount of ants being present including the other ant
populations (correlation; df=17, r=-0.211, p=0.385, Fig. 9).Fig. 9
Figure 6 Average number of crabs in the vegetation from 2019
(n=532; 19.7037±17.0064, orange) and the average number of
crabs in the vegetation from 2020 (n=150; 8.3333±7.1702, purple). The average numbers of crabs total from 2019 to 2020
were significantly different therefore the null hypothesis was rejected (F-test, df(2019)=26 and df(2020)=17; p=0.00027; T-test;
df=38, p=o.003766).
Figure 7 Comparison between average number of crabs in
the sand for only 2020 data (n=404; 24.4444±16.3782, blue)
and the average number of crabs in the vegetation (n=150;
8.33333±7.1702, green). The average number of crabs in the
vegetation zone was significantly smaller the the average number of crabs in the sand, therefore reject the null (F-test, df=17;
p=0.00027; T-test; df=23, p=0.003766).
Vol. 5, Fall 2020
Figure 8 Average number of ghost crabs in the vegetation
zone (n= 532; 19.703704±17.006367, green) compared to the average number of crabs in the sand (n=557; 20.6296±14.6633),
therefore we accepted the null hypothesis (F-test; df=26,
p=0.227438875; T-test; df=27, p=0.831197) for the 2019 year.
Figure 9 Density of total ants present (ants/sec) compared to
the density of ghost crabs (crabs/m2). These two values were
negatively not significant correlated (correlation; df=17, r=-0.211,
p=0.385).
Density of total ants present (ants/sec) compared to the density
of ghost crabs (crabs/m2). These two values were not significantly correlated (correlation; df=17, r=-0.211, p=0.385).
There was a negative, non-statistically significant relationship between the total density of crabs and the density of
yellow crazy ants (Correlation; df=17, r=--0.319. p=0.183, Fig.
10). There was also a negative, non-statistically significant relationship between the density of crabs in the sand and the density of yellow crazy ants (Correlation; df=17, r=-0.369, p=0.120,
Fig. 10). Though positive, there was no statistically significant
relationship between the density of crabs in the vegetation and
the density of yellow crazy ants (Correlation; df=17, r=0.107,
p=0.662, Fig. 10).
There was a negative not statically significant correlation
between the total average sizes of ghost crabs and the total ant
density (correlation; df=16, r=-0.233, p=0.2411, Fig. 11).
There were no significant effects of vegetation type (GLM;
F=13.4, df=11, p=0.27, Fig. 12) or yellow crazy ants presence/absence (GLM; F=1.54, df=2, p=0.46, Fig. 12) on ghost crab abundances. More diversity was present with the absence of yellow
crazy ants. There was only Akiaki and Akuikuli present with
Chasin
Size & Population Dynamics of Native Ghost Crabs in a Native Wildlife Refuge
129
Figure 10 Density of the yellow crazy ant (2020) in ants per
unit time (minutes) was affecting the density of ghost crabs, in
crab per unit area (m2). There was a non-statistically significant
relationship between the total density of crabs and the density of yellow crazy ants (correlation; df=17, r=--0.319, p=0.183),
a negative, non-statistically significant relationship between
the density of crabs in the sand and the density of yellow crazy
ants (correlation; df=17, r=-0.369, p=0.120) and positive, nonstatistically significant relationship between the density of crabs
in the vegetation and the density of yellow crazy ants (correlation; df=17, r=0.107, p=0.662).
Figure 12 Comparing the ghost crab abundance, the presence and absence of yellow crazy ants and the vegetation type
(Akiaki-pink, Akiaki/Akulukuli-blue, Naupaka/Akiaki-green,
Naupaka/Akiaki/Aku-purple, Naupaka/Akiaki/Hinahina-orange,
Naupaka/Hinhina-yellow). There were no significant effects of
vegetation type (GLM; F=13.4, df=11, p=0.27) or YCA presence/
absence (GLM; F=1.54, df=2, p=0.46) on ghost crab abundances.
More diversity was present with the absent of yellow crazy ants.
Data analysis done by Patrick Nichols.
Figure 11 Density of the total ant population (ants/sec) versus
the average hole diameter (cm). The correlation showed that
the two were negatively non-significantly correlated relationship
for the average mean hole diameter between the crabs and the
total ant density (correlation; df=16, r=-0.233, p=0.2411).
Figure 13 Comparing the ghost crab hole diameter in centimeters (corresponds to the size of the crab, Pombo et al. 2007) with
the presence (right)/absence (left) of ants and the vegetation
type (Akiaki-pink, Akiaki/Akulukui-blue, Naupaka/Akiaki-green,
Naupaka/Akiaki/Aku-purple, Naupaka/Akiaki/Hinahina-orange,
Naupaka/Hinhina-yellow). There was a significant effect of other ant presence/absence on ghost crab hole diameter (GLM;
F=19.5, df=2, p<0.001). There was also a significant interaction
of vegetation type and other ant presence/absence on ghost
crab hole diameter (GLM; F=4.73, df=1, p=0.03). There was a
marginally significant effect of the interaction between vegetation type and yellow crazy ant presence/absence on ghost crab
hole diameter (GLM; F=4.74, df=2, p=0.06). Data analysis done
by Patrick Nichols.
the presence of yellow crazy ants. However, there was a significant effect of other ant presence/absence on ghost crab hole
diameter (GLM; F=19.5, df=2, p<0.001, Fig. 13). There was also
a significant interaction of vegetation type and other Ant presence/absence on ghost crab hole diameter (GLM; F=4.73, df=1,
p=0.03, Fig. 13). Meaning that both vegetation and whether
or not other ants were present had combined effects to significantly impact the diameter of burrows. Finally, there was
a marginally significant (meaning almost at p=0.05) effect of
the interaction between vegetation type and yellow crazy ant
presence/absence on ghost crab hole diameter (GLM; F=4.74,
df=2, p=0.06, Fig. 13).
Discussion
Ghost Crabs from 2019 to 2020
This study is an on-going study from Spring 2019 to Spring
2020; comparing the data showing if there were any trends for
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130
number of crabs in the vegetation and sand zones and seeing
if there are any differences in where the crabs and ants are
located along the beach and if this affects the crab’s size. The
average number of crabs from 2019 to 2020 in the sand and
total did not change (Fig. 4 and 5).
The number of crabs found in vegetation decreased from
2019 to 2020 indicating that something was occurring in the
vegetation zone in regards to physical, chemical or the social
structures (Fig. 6, p<0.05).
Also in the 2019 season there were no significance between the number of crabs in the vegetation zone and the sand
zone but in the 2020 season there was a significantly smaller
number of crabs in the vegetation zone compared to the sand
zone (Fig. 7 and 8).
There was also the possibility of error due to our inability to count the ants under the dissection microscope because
of the SARS-CoV-2 pandemic, therefore we combined all ants
instead of grouping them into yellow crazy ants and other ants
(Fig. 9).
Vol. 5, Fall 2020
2007). Along with Anoplolepis gracilipes/Monomorium pharoanis there were a variety of ant species in the containers we collected, some of which may be more aggressive than others in
regards to aggregation. A. gracilipes/M. pharoanis was not found
in similar locations as the other species of ants, perhaps due
to their inability to aggregate with other ant species (Kirschenbaum 2007; Monomorium pharaonis . . . c1996–2020). James
Campbell National Wildlife Refuge (JCNWR) provides an excellent location for A. gracilipes to congregate as they require
large rocks or rock-lined areas for nesting grounds (Fluker and
Beardsley 1970). The presence of yellow crazy/pharaoh ants
seems to have an effect on the diversity of the different vegetation types, as in there were only two types of vegetation when
yellow crazy/pharaoh ant were present compared to five vegetation types in the absence of yellow crazy/pharaoh ants (Fig.
12). The vegetation could have changed based on certain ant
species being present, as certain ant populations alter the soil
composition (Fig. 12 and 13; Majer 1985).
Importance
Ghost Crabs
Two main species of the genus Ocypode were found at James
Campbell National Wildlife Refuge (JCNWR), Ocypode ceratophthalmus (horn-eyed ghost crab) and Ocypode laevis or Ocypode pallidula (pallid ghost crab) (Fellows 1975). Ghost crabs
are scavengers and predators which may dictate where they are
spaced as well (Rae et al. 2019). Size differences in crabs may
reflect differences in sexual and reproductive cycles (Jonah et
al. 2015). The sizes of burrows were found not to be related to
the total ant density (Fig. 11). Small differences in sizes may
be due to their gender as well as locational patterns (Fellows
1975; Lighter 1977). Smaller sized crabs tend to live lower on
the beach and larger crabs live more in the upper littoral zone,
because juveniles may desiccate if out of their burrows for too
long (Branco et al. 2010).
Average burrow size as measured by hole diameter was
not correlated with the total density of ants (Fig. 11). The
presence of YCA appeared to limit ghost crab distribution to
Naupaka/Akiaki vegetation (Fig. 12) and vegetation type had a
significant impact on ghost crab burrow sizes (Fig. 13).
Comparing Ants to Other Ants
Dr. Sheldon Plentovich indicated that some of the ant species
that looked like yellow crazy ants in our study were actually of
the species Monomorium pharoanis (a Pharaoh ant) or possibly a cardiocondyla. M. pharoanis are smaller compared to the
A. gracillepes but otherwise look similar. The yellow crazy ants
were only positively identified in the cemetery (reserve edge).
Many species of ants are successful at invading due to their
ability to have low intraspecific aggressions, high interspecific aggression and mutualistic relationships (Kirschenbaum
Ghost crabs were not found too far into each vegetation zone,
where most of the ant populations resides. Ghost crabs are a
good indication of a healthy ecosystem as they are top omnivores of the sand dune community (Brancho et al. 2010; Jonah et al. 2015). Differences in vegetation and the presence of
ants have an effect on the distribution of crab burrows (Fig. 13).
Spatial distribution of the different colonies of ants may play
a role in species competition and how the ants are affecting
the ecosystem as some ants are more aggressive than others
therefore affecting the ecosystem in different ways (Fluker and
Beardsley 1970; Majer 1985; Green et al. 2011). Understanding
the distribution and densities of certain ants (ie. yellow crazy/
pharaoh ant) could allow us to successfully locate the ants in
order to be able to eradicate them. It is better to understand the
ant population now before they cause massive destruction to
the ecosystem, as has been the case on Christmas Island with
the yellow crazy ant as well as in other ecosystems with other
ant species (Wetterer 2005; Green et al. 2011). Although the
pharaoh ants are not as costly to the ecosystem as the yellow
crazy ants, they still cause damage and may be affecting the
vegetation types or vegetation zone in general (Fig. 12 and 13).
Finding yellow crazy ants in the reserve allows us to monitor
them before they increase enough to have an impact on nesting bird populations in the reserve (Fluker and Beardsley 1970;
Boland et al. 2011).
Yellow crazy ants (similarly with the pharaoh ant) prefer
low wetland areas, JCNWR is a key place for these species to
reside and flourish. The yellow crazy ant look-alike, the pharaoh ant, also forms polydymonus nests, which are made up
of multiple colonies with more than one queen. Humans can
easily spread the pharaoh ants through scattering of rubbish.
It is possible that there were some burrows that we’re too
Chasin
Size & Population Dynamics of Native Ghost Crabs in a Native Wildlife Refuge
small to see or blown-out by the wind to be found through
visual observations. Differences in grain size and temperature
may also reflect the density distribution of the ghost crabs
which tend to hide in the burrows during the day to resist
predation or desiccation (Jonah et al. 2015; Pombo et al 2017).
Some of the beaches had bigger grains of sand, while others
had finer sand which would affect their burrow sizes as well
as their spatial distribution as beaches with smaller grain sizes
tend to have more species (Fellows 1975; Pombo et al. 2017).
Areas with smaller grain sizes tended to have higher density
and mean size of crabs. Although grain size was not measured,
this could explain why different vegetation had a range of burrow sizes as different vegetation types could promote smaller
or larger grain sizes (Fig. 13; Wang et al. 1998; Pombo et al.
2017). The slope or gradient of the beach may also have an
effect on spatial distribution (Pombo et al. 2017).
Future studies could examine whether ants prefer certain kinds of food bait in traps, as the pharaoh ants have been
shown to prefer honey (Kirschenbaum 2007; Monomorium
pharaonism . . . c1996–2020). This would enable us to specifically isolate the species of interest and hone in on their effects
on the environment. It would also be interesting to add ant
traps up the beach in different orientations and see if there
are more yellow crazy ant hatchlings near the rocky areas
in order to find out more about their reproductive success
as well as their hierarchal structure (Kirschenbaum 2007).
Some successful eradication procedures have been done on
Christmas Island using a fipronil bait and allowing for the
workers to bring the poisoned food back to the queen (Boland
et al. 2011).
We know very little about the behavior and burrowing
techniques of crabs or their larval life stages which may play
into where the crabs prefer to reside (Chan et al. 2006; Brancho et al. 2010; Jonah et al. 2015). Most work on ants has mostly been done on just a single species affecting an ecosystem,
but the ant community as a whole can affect or reflect the dynamics of the ecosystem and how they are changing it (Green
et al. 2011). As more species of ants invade we are going to
most likely see changes in community structure (Reimer
2004).
Changes in community composition may reflect an
ecosystem imbalance which can be addressed by improving
management. Humans can reduce threats to ecosystems by
being more mindful of the environment and preventing the
spread of invasive species (Clavero and García-Berthou 2005).
Ecosystems in Hawai‘i are particularly vulnerable because
many species have arrived or evolved here in the absences of
predators (Reimer 2004). Sandy intertidal zones are vital to
humans as they provide resources, shelter from storms, and
are of economic importance (Jonah et al. 2015). Understanding this fragile strand ecosystem is important as it is the link
between land and sea and a connection between humans and
the ocean.
131
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