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A SURVEY OF ECTOPARASITES ON DOMESTIC CAT (FELIS CATUS LINNAEUS, 1758) FROM RURAL AND URBAN AREA
Nur Izzah Izzati Binti Ahmad
RC 119
974 Bachelor of Science with Honours 2013 (Animal Resource Science and Management)
2013
i
ACKNOWLEDGEMENTS
First of all I would like to thanks Allah S.W.T because bless me with a good health
so that I able to complete this study. Secondly, I would like to express my
appreciation and gratitude to my supervisor Dr Mohd Azlan Jayasilan Bin Gulam
Azad, for the precious advice, support, comments and guidance throughout this study.
I also would like to thanks to my co-supervisor, Prof. Dr. Mohd Tajuddin Bin
Abdullah and special thanks to PhD student Miss Madinah Binti Adrus, for her
guidance and teaching me the right way to identify the ectoparasites.
This appreciation is also forwarded to Sarawak Society for Prevention of Cruelty to
Animal (SSPCA) for providing stray cats and to the SSPCA staffs who helping me in
handling the cats during ectoparasite sampling process.
Last but not least, high appreciation and thanks to my family, who always support
me in vary ways and also thanks to my fellows friends Noor Aisyah Binti A. Rahim,
Siti Hanisah Binti Zahuri and also Asma’a Binti Che Mohd Rosli because willing to
assist me for my field sampling.
ii
DECLARATION
No portion of the work referred to in this dissertation has been submitted in support
of an application for another degree of qualification of this or any other university or
institution of higher learning.
_____________________
Nur Izzah Izzati Binti Ahmad
Programme of Animal Resources Science and Management
Faculty of Resources Science and Technology
Universiti Malaysia Sarawak
iii
TABLE OF CONTENTS
CHAPTER TITLE PAGES
ACKNOWLEDGEMENTS i
DECLARATION ii
TABLE OF CONTENTS Iii
LIST OF TABLES v
LIST OF FIGURES vi
LIST OF ABBREVIATIONS vii
ABSTRACT viii
ABSTRAK viii
1.0 INTRODUCTION
1.1 General introduction 1
1.2 Hypothesis statement 4
2.0 LITERATURE REVIEW
2.1 Fleas 5
2.2 Mites 6
2.3 Lice 7
2.4 Tick 7
3.0 MATERIAL AND METHOD
3.1 Study areas 13
3.2 Host collection 14
iv
3.3 Examination of ectoparasites 14
3.4 Processing sample and mounting of ectoparasites 15
3.5 Identification of ectoparasite 15
3.6 Data analysis 15
4.0 RESULT
4.0 Result 17
5.0 DISCUSSION
5.1 Urban habitat 21
5.2 Rural habitat 23
6.0 CONCLUSION AND RECOMMENDATION
6.1 Conclusion 24
6.2 Recommendation 24
REFERENCES 26
APPENDICES 30
v
LIST OF TABLES
Table 1 The previous ectoparasites studies conducted in Malaysia
from 2005 to 2012
10
Table 2 The previous ectoparasites studies conducted in other
countries from 2002 to 2010
11
Table 3 The previous ectoparasites studies conducted in other
countries 2002 to 2010 (Continues)
12
Table 4 Percentage of ectoparasite infestation from two habitats 16
Table 5 Prevalence of ectoparasites based on urban and rural habitat 17
Table 6 The diversity of the ectoparasites found on domestic cats 18
vi
LIST OF FIGURES
Figure 1 Host sampling areas 14
Figure 2 Haemaphysalis sp2 in dorsal view 19
Figure 3 Haemaphysalis sp2 in ventral view 19
Figure 4 Haemaphysalis sp3 in dorsal view 19
Figure 5 Haemaphysalis sp3 in ventral view 19
Figure 6 Ctenocephalides felis 20
Figure 7 Notoedres cati in ventral view 20
Figure 8 Felicola subrostratus 20
Figure 9 Listeropheridae sp1 in ventral view 20
Figure 10 Heterodoxus spiniger in dorsal view 20
vii
LIST OF ABBREVIATIONS
Mm Millimeter
% Percent
Km Kilometer
Kg Kampung
+ve Positive
–ve Negative
N Number
viii
A Survey of Ectoparasite on Domestic Cat (Felis catus LINNAEUS, 1758) From Rural
and Urban Area.
Nur Izzah Izzati Binti Ahmad
Animal Resources Science and Management
Faculty of Resources Science and Technology
Universiti Malaysia Sarawak
ABSTRACT
A survey of ectoparasites on domestic cats from two different habitats has been conducted in six different
locations from October 2012 to March 2013. A total 69 of domestic cats have been examined for the presence of
ectoparasites, comprising 34 from urban habitat and 35 from rural habitat. Thirty six (52.1%) domestic cats have
been discovered to be infested with at least one species of ectoparasites. Eight species of ectoparasites manage to
be collected throughout the study. Felicola subrostrata (21.7%) is the common ectoparasite recovered from the
infested host. Listophorid mites (14.5%) show the high infestation on domestic cat from rural habitat. Sarcoptes
scabiei (10.1%), Notoedes cati (8.7%), Ctenocephalides felis (7.2%) infested the both rural and urban domestic
cat with the quite similar rate of infestation. Meanwhile, Heterodoxus spiniger (2.9%) and other two species of
Ixodid tick, which both show 1.4% of infestation rate only found on the urban domestic cats. However, S. scabiei
and C. felis are the only ectoparasites species among that have been found are important in medical point of view.
There is significant different in species diversity (χ2=38.29, p=0.00), as urban habitat show the higher number of
ectoparasite species compared to the rural habitat.
Keyword: ectoparasite, domestic cats, rural, urban, significant different.
ABSTRAK
Satu kajian mengenai ektoparasit pada kucing domestic telah dilaksanakan di enam lokasi yang berbeza dari
bulan Oktober 2012 hingga bulan Mac 2013. Sejumlah 69 ekor kucing domestik telah diperiksa untuk kehadiran
ektoparasit, yang terdiri daripada 34 ekor dari habitat bandar dan 35 ekor dari habitat luar bandar. Tiga puluh
enam ekor (52.1%) kucing domestik telah dikenalpasti dijangkiti dengan sekurang-kurangnya satu spesies
ektoparasik. Lapan spesies ektoparasik berjaya dikumpul sepanjang kajian. Felicola subrostrata (21.7%) adalah
yang biasa ditemui pada tuan rumah yang dijangkiti. Hama Listophorid (14.5%), menunjukkan jangkitan yang
tinggi pada kucing domestik dari luar bandar. Sarcoptes scabiei (10.1%), Notoedes cati (8.7%), Ctenocephalides
felis (7.2%) menjangkiti kucing domestik dari habitat bandar dan luar bandar dengan kadar jangkitan yang agak
sama. Manakala, Heterodoxus spiniger (2.9%) dan dua spesies sengkenit Ixodid, yang kedua-dua spesies
menunjukkan 1.4% kadar jangkitan, hanya dijumpai pada kucing domestik dari habitat bandar.
Walaubagaimanapun, hanya S. scabiei dan C. felis adalah spesies ektoparasit dari kalangan yang telah ditemui
mempunyai kepentingan dari segi perubatan. Terdapat perbezaan yang penting dari segi kepelbagaian spesies
(χ2=38.29, p=0.00), yang mana, habitat bandar menunjukkan bilangan spesies ektoparasik yang tinggi
berbanding dengan habitat luar bandar.
Kata kunci: ektoparasit, kucing domestik, luar bandar, bandar, perbezaan yang penting
1
CHAPTER 1
INTRODUCTION
1.1 General introduction
The domestic cat (Felis catus) is the most favorite companion animal among
Malaysian. These may due to their cuteness and lovely appearance and also their
capability to control the population of the pest in their owner house. The domestic
cats can be found in all over the place that have the human presence, whether in rural
or urban habitat.
Domestic cat population serves as an importance host for the various medical
important species of ectoparasites (Borji et al., 2011) and the infested stray domestic
cats can easily spread many zoonotic diseases to the indoor domestic cats via direct
contact (Jittapalapong et al., 2008) as the ectoparasites serve as the vector for many
zoonotic disease such as Lyme disease, plague, Rocky mountain spotted fever,
tularemia (Neldes & Reeves, 2005), rabies, cat-scratch disease, ehrlichiosis and
toxoplasmosis (Jittapalapong et al., 2008). The potential of diseases transmission
from domestic cats to the human being is high as the domestic cats share the same
habitat with the human being.
Ectoparasites are the organisms that live outside the other animal or organism’s body
and feed on it. The ectoparasites are comprised from many group of parasitic
2
animals. The common ectoparasites infesting the domestic cats are belongs to the
four groups which are fleas, mites, lice and ticks.
Ectoparasites are diverse and highly adaptable to their host, as they permanently or
periodically infesting their host or their host’s habitat (Nithikathkul et al., 2005).
According to Dick et al. (2003), the ectoparasites that permanently infesting their
host have special characteristic such as small in size, flatted form and also the ability
to move on their host’s body surface or within the hairs.
Besides being a vector for disease, the ectoparasites also can cause direct damage
toward the host by ingesting blood, skin, and muscle of the host (Balashov, 2007;
Changbunjong et al., 2009). The infestation of ectoparasites may cause the host to
feel uncomfortable due to mechanical damage, action of saliva and allergic reaction
at the site that have been ingested by ectoparasites (Balashov, 2007; Jittapalapong et
al., 2008; Changbunjong et al., 2009; Mendes-de-Almeida et al., 2011).
Some of the ectoparasitic arthropods are the pests that irritates the host, whether
human or domestic animal (Jittapalapong et al., 2008), besides their role as the main
vector and potentially transmit a variety of diseases (Akucewich et al., 2002;
Jittapalapong et al., 2008; Madinah et al., 2011), cause the pruritic and non-pruritic
skin disorder (Akucewich et al., 2002; Jittapalapong et al., 2008; Chanie et al., 2010)
and hypersensitivity disorder in domestic animal (Akucewich et al., 2002;
Jittapalapong et al., 2008). The high infestation of the ectoparasites can cause the
domestic animal to lose a lot of blood and that may threaten the life of the young and
3
debilitated domestic animals (Jittapalapong et al., 2008; Mendes-de-Almeida et al.,
2011).
According to the Heukelbach et al. (2012), the prevalence of the ectoparasitic-vector
species may serve as the indicator in order to measure the risk of the transmission of
the pathogen to the human that live associated with the ectoparasitic-vector borne
pathogen.
The ectoparasites can be classified into the host specific and non-host specific
ectoparasites. The host-specific ectoparasites is the ectoparasites that exclusively
infested one or a few species of host that closely related (Dick et al., 2005), for
example, Polyplacid lice is the highly host specific ectoparasite of bird and it will not
infesting the human (Nelder & Reeves, 2005).
The prevalence of ectoparasites on domestic cats and other domestic animal is
depends on the rate of proliferation of the ectoparasites which influenced by the
environmental condition, humidity and temperature of the habitat (Akucewich et al.,
2002; Jittapalapong et al., 2008; Paramasvaran et al., 2009; Mendes-de-Almeida et
al., 2011; Nuchjangreed & Somprasong, 2007). Furthermore, Jittapalapong et al
(2008) also state that socio-economy also seemed one of the important factor for the
prevalence of the ectoparasites as increasing cost of living may cause the pet owners
least concern and unaffordability to manage their pet’s health. The relationship of
domestic cats with other animal also can contribute to the prevalence of ectoparasites
as the ectoparasites can be transmitted from other animal host (Paramasvaran et al.,
2009). The advance in transportation system and relocation of infested animal also
4
can influence the prevalence and the infestation rate of ectoparasites on domestic cats
and other animal at that particular area (Nuchjangreed & Somprasong, 2007).
As there is no prior study conducted regarding to the ectoparasites of domestic cats
from different habitats, therefore this study is important in order to understand more
about the distribution of the ectoparasites of domestic cats from different habitats.
This study also can provide the information on the general health condition based on
the level of ectoparasites prevalence.
The main objectives of this study were to determine whether there are any significant
differences on the diversity of ectoparasites between urban and rural habitats and to
describe the type of ectoparasites found on domestic cats in Kuching region.
1.2 Hypothesis statement.
HO : There is no significant different on the diversity of ectoparasites
between the urban and rural area.
HA : There is significant different on the diversity of ectoparasites
between the urban and rural area.
5
CHAPTER 2
LITERATURE REVIEW
There are many organisms that belongs to different taxa such as insect, arthropod and
annelid live as the parasite on other animal. The common ectoparasite group that
infesting the mammals are fleas, mites, lice and ticks.
2.1 Fleas
Fleas from order Siphonaptera are small wingless insect which both male and female
adult largely infesting and feed on the mammal’s blood (Taylor et al., 2007). The
complete duration of fleas life cycle is highly influenced by the environmental
condition, as the life cycle can be completed in less than two weeks or extent for
several months (Beck & Pfister, 2004).
Fleas are the predominant ectoparasites to infest the domestic cat (Akucewich et al.,
2002; Jittapalapong et al., 2008; Xhaxhiu et al., 2009; Chin et al., 2010; Borji et al.,
2011; Mendes-de-Almeida et al., 2011). The flea infestation can directly affect their
host by irritating them and beside feeding on blood they can cause dermatitis and
allergic reaction (Borji et al., 2011; Nuchjangreed & Somprasong, 2007) due to
chemical components of flea saliva (Beck & Pfister, 2004). Furthermore, the fleas
6
bite can create a wound which may develop a crust of dried exudates (Taylor et al.,
2007).
The fleas also play an important role in transmitting serious disease such as
bartonellosis (cat scratch disease) to both human and animal (Jittapalapong et al.,
2008) and may be a vector for many type of zoonotic diseases (Nelder & Reeves,
2005) and intermediate host for the helminths species such as Hymenolepis diminuta,
H. nana and Dipylidium caninum (Heukelbach et al., 2012). The long period of fleas
infestation can lead to the chronic blood spoliation, this resulted from the blood
ingesting by fleas, as it capable to ingest large amount of blood per day (Mendes-de-
Almeida et al., 2011).
2.2 Mites
Mites which is potential intermediate host for anoplocephalid cestodes is belongs to
order Acariformes, is small size animal which less than 1 mm long and using the hair
or setae as the sensory organ as most of them are blind and they also have a short life
cycle which less than four weeks or even eight days (Taylor et al., 2007). Taylor et
a.l (2007) also state that mites infestation which can occur via direct contact are
called acariasis and the severe dermatitis resulting from the acariasis are known as
mange. Most of the mites spend their entire live inhibit the host skin and feed on
their host blood, lymph, skin debris or sebaceous secretion (Taylor et al., 2007).
7
2.3 Lice
Lice are the small wingless insect with length about 0.5-8 mm and most of the
species are blind with only few species have photosensitive eyes spot (Taylor et al.,
2007). Taylor et al. (2007) also stated that this parasite is highly host specific which
feed on epidermal debris, parts of feathers, sebaceous secretions and blood as have
potential in transmission of anaplasmosis disease. Heavy lice infestation is usually
called pediculosis (Taylor et al., 2007).
2.4 Ticks
Tick is large and long-lived ectoparasite which feed on the vertebrate (mammals and
birds) blood (Taylor et al., 2007). According to Nuchjangreed and Somprasong
(2007), the surrounding temperature can give the great impact on the rate of growth,
development reproduction and survival of certain species of tick.
Tick is a vector for virus, bacteria and protozoa (Nuchjangreed & Somprasong., 2007;
Taylor et al., 2007; Xhaxhiu et al., 2009; Mendes-de-Almeida et al., 2011) as it play
the important role in transmission of disease (Akucewich et al., 2002; Dantas-Torres
et al., 2009) such as ehrlichiosis disease (Jittapalapong et al., 2008). Beside the
hazardous of the disease that it carries the tick saliva also harmful as tick saliva
contain toxin which can cause the tick paralysis (Nuchjangreed & Somprasong.,
2007).
There are many ectoparasites studies have been conducted in Malaysia, and largely
those studies are more focusing on the ectoparasites of the wild small mammals. The
studies of ectoparasites on small mammals conducted by Chuluun et al. (2005),
Mariana et al. (2005), Mariana et al. (2008), Paramasvaran et al. (2009), Mariana et
8
al. (2011) and Madinah et al. (2011) at different locations in Malaysia show that
Laelaps echidninus, L. nuttali and Leptotrombidium deliense are the common
ectoparasites species found infesting wild small mammals in Malaysia. Beside the
study of ectoparasites on small mammals, the study of ectoparasites on other host
species, such as domestic cats, domestic dogs and human which conducted by
Norhidayu et al (2012), Kernif et al (2012) and the Chin et al (2010) respectively.
The study of the ectoparasites on the domestic animal have been conducted
worldwide as the domestic animal live close to human and the ectoparasites of
domestic animal also have the potential to be infested human being (Jittapalapong et
al., 2008). The Chin et al (2010), reported the infestation of cat’s fleas
Ctenocephalides felis on six university student in Kuala Lumpur. C. felis or its
common name cat’s fleas is the predominant fleas found on the domestic cats
(Akucewich et al., 2002; Balashov, 2007; Jittapalapong et al., 2008; Chanbunjong et
al., 2009; Xhaxhiu et al., 2009; Mendes-de-Ameida et al., 2011; Borji et al., 2011).
This ectoparasite species also have been found on the domestic dogs in Brazil
(Dantas-Torres et al., 2009; Heukelbach et al., 2012) and also on sheep and goat in
Nigeria (Ofukwu & Akwuobu, 2010).
Beside C. felis, C. canis (Nithikathkul et al., 2005; Balashov, 2007; Nuchjangreed &
Somprasong 2007; Xhaxhiu et al., 2009) and Rhipicephalus sanguineus
(Nithikathkul et al., 2005; Balashov, 2007; Nuchjangreed & Somprasong 2007;
Xhaxhiu et al., 2009; Dantas-Torres et al., 2009; Bahrami & Delpisheh, 2010;
Klimpel et al., 2010) also commonly discovered on the domestic dogs. Beside
infesting the domestic dogs, R. sanguineus also have been reported to infest domestic
cats (Akucewich et al., 2002; Mendes-de-Ameida et al., 2011) and sheep and goat
9
(Ofukwu & Akwuobu, 2010). The previous study regarding to the ectoparasites
found on the various host are summarized in Table 1 to Table 3.
10
Table 1: The previous ectoparasites studies conducted in Malaysia from 2005 to 2012
Host(s) Result Technique(s) Effort Study
Rodent Three species of mites, two species of chiggers and two
species of lice. Combing 11 Chuuluun et al (2005)
2
Bats and small
mammals
Seven species of mites, six species of chiggers and five
species of ticks. Combing 272 Mariana et al (2005)
2
Birds, bats and small
mammals
Eleven species of mites, five species of chiggers and five
species ticks. Combing 161 Mariana et al (2008)
2
Rodent and shrews Nine species of mites, four species of chiggers, four species
of tick and one species of fleas. Combing 201
Paramasvaran et al
(2009)1,2
Human One species of fleas. Hand-
removing 6 Chin et al (2010)
1
Bats, tree shrew,
rodent and myriapod
Eight species of mites, eight species of chiggers and five
species of ticks.
Combing and
using forceps 140 Mariana et al (2011)
2
Small mammals Eight species of mites, three species of and three species of
ticks Combing 16 Madinah et.al (2011)
2
Cat One species of mites. Combing 40 Jeffery et.al (2012)1
Cat One species of lice. Combing 102 Norhidayu et al (2012)1
Dog Two species of ticks Combing 212 Kernif et al (2012)2
Footnotes:
1, Urban habitat; 2, Rural habitat
*Note = the effort of the studies was measured by the number of the host that have been examined.
11
Table 2 : The previous ectoparasites studies conducted in other countries from 2002 to 2010
Host(s) Result Technique(s) Effort Study
Cat One species of mites, one species of lice four
species of ticks and three species of fleas.
Combing, skin scraping and
ear swab 200 Akucewich et al (2002)
b,1
Cat, dog , cattle, horse,
goat, sheep and pig
Ten species of lice, twelve species of mites and
seventeen species of tick Balashov (2007)
i,1,2
Cat One species of mites, one species of lice and
two species of fleas. Combing and skin scraping 575 Jittapalapong et al (2008)
a,1
Cat, dog and cattle One species of fleas, two species of tick and
three species of lice.
Combing, brush the animal
coat and collect using forceps 111 Chanbunjong et al (2009)
a,2
Cat and dog Two species of mites, one species of lice, two
species of tick and three species of fleas. Combing and skin scraping 207 Xhaxhiu et al (2009)
c,1
Cat Two species of mites, one species of lice, one
species of tick and one species of fleas. Combing 292
Mendes-de-Ameida et al
(2011)d,1
Cat One species of mites and one species of fleas. Combing 52 Borji et al (2011)e,1
Dog One species of tick and one species of fleas Hand-removal 100 Nithikathkul et al (2005)a,1
Dog One species of tick Combing and using forceps 16 Nelder and Reeves (2005)f,1
Dog Three species of mites, one species of lice, two
species of ticks and three species of fleas. Use prophylactic insecticide 83
Nuchjangreed and Somprasong
(2007)a,1
Footnotes:
a, Thailand; b, Florida; c, Albania; d, Brazil; e, Iran; f, United State; g, Nigeria; h, Ethiopia; i, Russia; 1, Urban habitat; 2, Rural habitat
*Note = the effort of the studies was measured by the number of the host that have been examined.
12
Table 3 : The previous ectoparasites studies conducted in other countries 2002 to 2010 (Continues).
Host(s) Result Technique(s) Effort Study
Dog Two species of tick and one species of fleas. Combing 202 Agbolade et al (2008)g,1
Dog One species of lice, two species of ticks and
one species of flea. Hand-removal 41 Dantas-Torres et al (2009)
d,1
Dog One species of mites, two species of lice
and four species of tick. Combing 802
Bahrami and Delpisheh
(2010)e,1
Dog One species of lice and three species of
ticks
Combing euthanized
animal 46 Klimpel et al (2010)
d,1
Dog Three species of ticks and one species of
fleas. Combing 15 Heukelbach et al (2012)
d,1
Sheep and goat Four species of mites, two species of lice,
one species of tick and one species of keds.
Skin scraping and hand-
removal 1600 Chanie et al (2010)
h,1
Sheep and goat
Two species of mites, two species of lice,
six species of tick and one species of flea. Skin scraping
1040 Ofukwu and Akwuobu
(2010)g,2
Footnotes:
a, Thailand; b, Florida; c, Albania; d, Brazil; e, Iran; f, United State; g, Nigeria; h, Ethiopia; i, Russia; 1, Urban habitat; 2, Rural habitat
*Note = the effort of the studies was measured by the number of the host that have been examined.
13
CHAPTER 3
MATERIAL AND METHODS
3.1 Study areas
The domestic cats were caught in the two different habitats in Kuching region which
are urban and rural between October 2012 to March 2013. The urban area refers to
the place or area that are well develop infrastructurally and have many kind of
facilities such as shop premises, market, school, wide road and clinic and managed
by the District Council. Whereas the rural area refers to the place or area that is
located quite far (30-40km) from the city and lack with urban facilities. The urban
area also has the higher human population than rural area. The urban domestic cats
were caught from Stutong, Kota Samarahan and Satok. While, the rural domestic cats
were caught from Kampung Empila, Kampung Niup and Kampung Mang.
14
Figure 1: Host sampling sites (Google map, 2013)
3.2 Host collection
All domestic cats were caught by hand. The cat food pellet was use as bait, as the
domestic cats do not like being approach by the stranger. The cat food pellet makes
the domestic cats calm and makes the sampling process easier.
3.3 Examination of ectoparasites
The fur of the domestic cats were combed with a fine tooth comb on the white cloth.
All the ectoparasites from the white cloth was collected using the moistened end of a
sharpened wooden applicator stick and placed in the sample tube that contain 70%
ethanol. The sample tube labeled with sufficient information, such as location, cat’s
sex and date.
15
3.4 Processing sample and mounting of ectoparasites
The content of the sample tube was examined under stereo microscope in order to
identify and separate the ectoparasites. The ectoparasites were sorted based on
morphology. The mites were mounted directly in the Hoyer’s medium and heated
with bunsen burner for clearing. Whereas, for the bigger ectoparasites (lice and
fleas), were placed in the lactophenol for a week at room temperature before being
mounted in Hoyer’s medium. The ticks were the only ectoparasite that was not being
mounted in Hoyer’s medium, because the ticks are bigger enough to be examined
and identified by using the stereo microcope.
3.5 Identification of ectoparasites
The prepared slides were examined under compound microscope with 40x, 100x,
400x and 100x magnification in order to identify the mounted ectoparasite.
Ectoparasites were identified to species level where possible by using the published
keys and taxonomic drawings and reference by Taylor et al. (2007).
3.6 Data analysis
The data was analyzed by using the chi-square test in order to determine whether
there is significance of diversity of ectoparasites based on habitats. The chi-square
test was performed using PAST (Paleontological Statistics software package for
education and data analysis) version 2.04 (Hammer et al., 2001).