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\begin{document}

\title{Associated factors contributing abundance of fleas on rodents in plague
endemic area of Karatu district, northern Tanzania.}



\author[1]{Joshua Jakoniko}%
\author[2]{Apia Massawe}%
\author[1]{Elisa Mwega}%
\author[3]{Stella Kessy}%
\affil[1]{Sokoine University of Agriculture College of Veterinary Medicine and Biomedical Sciences}%
\affil[2]{Sokoine University of Agriculture}%
\affil[3]{Nelson Mandela African Institute of Science and Technology School of Life Sciences and Bio-Engineering}%


\vspace{-1em}



  
  \date{May 27, 2024}


\begingroup
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\selectlanguage{english}
\begin{abstract}
Fleas are small wingless hematophagous insect that are frequently
infesting on rodents and other small mammals while acting as reservoirs
and vectors of many rodent borne zoonotic diseases including plague
infectious disease which is threat to the public health in many part of
the world including Tanzania. 291 rodents from nine species were
captured with Sherman traps in farm land, peridomestic areas, bush land
and forest buffer zone across wet and dry season in plague and
non-plague foci villages. Captured rodents were anaesthetized and 190
fleas comprising four species were collected and morphologically
identified with available dichotomous key. Dinopsyllus lypusus were
(46.32\%), Ctenophthalmus spp (26.84\%), Xenopsylla brasiliensis
(16.32\%) and Xenopsylla cheopis (10.53\%). 38.42\% of fleas were
collected from Mastomy natalensis, 22.63\% from Lemniscomys striatus and
18.42\% from Rattus rattus. Highest flea infestation prevalence was
found on R.rattus and was strongly associated with X.cheopis and
X.brasiliensis. Specific flea index of X.cheopis on R.rattus was (01) in
plague foci and (\textless{}0.5) in non-plague foci villages. Result of
GLM final model indicated that flea abundance was significant influenced
by rodent species (p \textless{} .001), season (p= .031), habitat type
(p= .02), rodent weight (p \textless{} .001), rodent sex (p \textless{}
.001) and plague locations (p= .02). There was significance difference
in variation of flea abundance between rodent sexes (W = 9158.5, p =
.009) and a weak positive correlation between rodent's weight and
abundance of fleas (R= 0.17, p\textless{} 0.05). Despite that, specific
flea index of X.cheopis on rats in both plague foci and non-plague foci
villages were not indicating alarming condition that would require
urgent control of fleas, still society should consistently adhere to
rodent and fleas control methods in order to limit their interaction to
the society especially in farm land and peridomestic areas where human
activities are high.%
\end{abstract}%



\sloppy


\textbf{CHAPTER THREE}

\textbf{MANUSCRIPT TWO}

\textbf{3.0 Associated factors contributing abundance of fleas on
rodents in plague endemic area of Karatu district, northern Tanzania.}

\textbf{Corresponding Author:}

\textbf{Joshua Reuben Jakoniko} \textsuperscript{a, e *}

Candidate, MSc Public Health Pest Management, Department of Veterinary
Medicine and Public Health, Sokoine University of Agriculture, P.O.Box
3021, Morogoro, Tanzania.

Email: jakonikojoshuaa@gmail.com. Telephone +255765396402

\textbf{Authors}

\textbf{Professor Apia Massawe} \textsuperscript{d, e} \textbf{(PhD)}

Researcher at Institute of Pest Management Sokoine University of
Agriculture, Morogoro Tanzania and a leader of African centre of
Excellence for Innovative Rodent Pest Management and Biosensor
Technology Development, P.O. Box 3110, Morogoro, Tanzania.

Email: apiamas@yahoo.com

\textbf{Dr. Elisa Daniel Mwega} \textsuperscript{b} \textbf{(PhD)}

Lecture at Sokoine University of Agriculture (SUA). Department of
Veterinary Microbiology, Parasitology and Biotechnology, Sokoine
University of Agriculture, P.O. Box 3015, Morogoro, Tanzania.. Email:
elisamwega@gmail.com

\textbf{Stella Thadeus Kessy} \textsuperscript{c}

Researcher Nelson Mandela African Institute of Science and Technology,
School of Life Science and Bio Engineering (LISBE), P.O. Box 447,
Arusha, Tanzania.

Email: kessystella78@gmail.com

\textsuperscript{a} Department of Veterinary Medicine and Public Health,
Sokoine University of Agriculture, P.O.Box 3021, Morogoro, Tanzania.

\textsuperscript{\textbf{b}} Department of Veterinary Microbiology,
Parasitology and Biotechnology, Sokoine University of Agriculture, P.O.
Box 3015, Morogoro, Tanzania.

\textsuperscript{\textbf{c}} Nelson Mandela African Institute of Science
and Technology, School of Life Science and Bio Engineering (LISBE), P.O.
Box 447, Arusha, Tanzania.

\textsuperscript{\textbf{d}} Institute of Pest Management, Sokoine
University of Agriculture, P.O. Box 3110, Morogoro Tanzania.

\textsuperscript{e} African centre of Excellence for Innovative Rodent
Pest Management and Biosensor Technology Development (ACE IRPM\&BTD).

\textbf{3.1 Abstract}

Fleas are small wingless hematophagous insect that are frequently
infesting on rodents and other small mammals while acting as reservoirs
and vectors of many rodent borne zoonotic diseases including plague
infectious disease which is threat to the public health in many part of
the world including Tanzania. 291 rodents from nine species were
captured with Sherman traps in farm land, peridomestic areas, bush land
and forest buffer zone across wet and dry season in plague and
non-plague foci villages. Captured rodents were anaesthetized and 190
fleas comprising four species were collected and morphologically
identified with available dichotomous key. \emph{Dinopsyllus
lypusus}were (46.32\%)\emph{, Ctenophthalmus spp} (26.84\%),
\emph{Xenopsylla brasiliensis} (16.32\%) and \emph{Xenopsylla cheopis}
(10.53\%). 38.42\% of fleas were collected from \emph{Mastomy
natalensis} , 22.63\% from\emph{Lemniscomys striatus} and 18.42\% from
\emph{Rattus rattus} . Highest flea infestation prevalence was found on
\emph{R.rattus} and was strongly associated with \emph{X.cheopis} and
\emph{X.brasiliensis} . Specific flea index of \emph{X.cheopis} on
\emph{R.rattus} was (01) in plague foci and (\textless{}0.5) in
non-plague foci villages. Result of GLM final model indicated that flea
abundance was significant influenced by rodent species (\emph{p}
\textless{} .001), season (\emph{p} = .031), habitat type (\emph{p} =
.02), rodent weight (\emph{p} \textless{} .001), rodent sex (\emph{p}
\textless{} .001) and plague locations (\emph{p} = .02). There was
significance difference in variation of flea abundance between rodent
sexes (W = 9158.5, \emph{p} = .009) and a weak positive correlation
between rodent's weight and abundance of fleas (\emph{R} = 0.17,
\emph{p} \textless{} 0.05). Despite that, specific flea index
of\emph{X.cheopis} on rats in both plague foci and non-plague foci
villages were not indicating alarming condition that would require
urgent control of fleas, still society should consistently adhere to
rodent and fleas control methods in order to limit their interaction to
the society especially in farm land and peridomestic areas where human
activities are high.

Key words: Ectoparasite, Flea, Rodent, Plague endemic area. Plague foci,
Specific flea index

\textbf{3.2 Introduction}

Fleas (Siphonaptera) are hematophagous insects and obligatory
ectoparasites of vertebrates that have colonized variety of habitats
from wet tropical forests to semi-arid and desert areas ( Zaj\selectlanguage{polish}ą\selectlanguage{english}c \emph{et
al} ., 2020; Zhang \emph{et al} ., 2022). They have an obligate
parasitic life with a wide range of potential hosts, primarily small
mammals and less often birds (Eads \emph{et al} ., 2020). Fleas exhibit
a holometabolous type of lifecycle and complete their life cycle in 14
days to 140 days, depending mainly on temperature and humidity (Kr\selectlanguage{ngerman}ämer
and Mencke, 2012; Ashwini \emph{et al} ., 2017; Gram and Short, 2020).

Fleas contribute signi\selectlanguage{english}ficantly in the circulation of pathogens hence
receiving considerable attention mostly because they are vector of many
zoonotic diseases (Eads \emph{et al} ., 2020). They play role in
spreading plague disease as well as other emerging pathogens that causes
zoonoses such as bartonelloses, tularemia and rickettsioses (flea-borne
spotted fever, Q fever and murine typhus), fleas also maintains and
transmits pathogens of myxomatosis and trypanosomiasis and can act as
intermediate hosts of some helminthes (Zaj\selectlanguage{polish}ą\selectlanguage{english}c \emph{et al} ., 2020;
C\selectlanguage{ngerman}ófreces \emph{et al} ., 2021). However, the most known \selectlanguage{english}flea borne
zoonotic disease is plague caused by bacteria known as \emph{Yersinia
pestis} ; a zoonotic bacteria renowned for killing millions of humans
during the Black Death in Europe in 14 century (Ditchburn and Hodgkins,
2019; Barbieri \emph{et al} ., 2021; Izdebski \emph{et al} ., 2022).
Plague is still threatening the public health in some parts of the
world, especially in African countries (Ditchburn and Hodgkins, 2019).
By doing surveillance studies on flea assemblage on rodent community in
plague endemic areas we can understand factors that influence flea
abundance and infestation potential on rodents and being aware of
significant factors that may endanger the public health and therefore we
can be in a good position to suggest effective intervention strategies
to control their spread.

Plague persistence in plague endemic areas is characterized by
coexistence of interactions among rodent communities and flea species,
whereby some species of rodents and fleas are better at maintaining,
amplifying and transmitting \emph{Yersinia pestis} (Gage and Kosoy,
2006; Antolin \emph{et al} ., 2010). Some of frequently reported fleas
and rodents species involved in plague maintenance and transmission
are\emph{Xenopsylla cheopis, Xenopsylla} \emph{brasiliensis}
and\emph{Dinopsyllus lypusus} and rodent species \emph{Lophuromys spp}
,\emph{Praomys delectorum} , \emph{Graphiurus murinus} ,
\emph{Lemniscomys striatus, Mastomys natalensis} , and \emph{Rattus
rattus} (Kilonzo\emph{et al} ., 2006; Makundi \emph{et al} ., 2008;
Eisen and Gage, 2009). A study done by Ziwa et al. (2013) has reported
the concurrence of host-vector interaction to be responsible for
transmission of plague disease. Also Makundi et al. (2015) has reported
the presence of multiple associations between domestic and peridomestic
rodent species infested with fleas to be among major contributing
factors for persistence and spread of plague in plague endemic areas.
Moreover, the roles of hosts and \selectlanguage{english}fleas, for instance in plague
maintenance or ampli\selectlanguage{english}fication, is mostly affected with change of space and
time (Kosoy\emph{et al} ., 2017).

The distributions and community structure of fleas are influenced by
numerous biotic and abiotic factors, that include, host species
diversity, sex, age, body size, immune status, host population
abundance, habitat diversity and seasonal variation of temperature and
precipitation (Lopez-Perez \emph{et al} ., 2017, 2022). Host diversity
is a relevant factor to consider since it involves variation in flea
species richness (Fantozzi \emph{et al} ., 2022), yet it is not a fixed
rule, fleas can infest hosts phylogenetically close, switching between
coexisting species within guilds (Cofreces \emph{et al} ., 2021).

Noting this uncertainty plague management in plague foci is mostly
efficient when encompassing rodents and their \selectlanguage{english}flea parasites. As
described by Garcia-Longoria \emph{et al.} (2019) host communities can
influence parasitism. Community organization of parasites is considered
to be determined mainly by their hosts because host present a habitat
for parasites, providing them with place for living, feeding and mating
(Fellin and Schulte, 2022). Thus, a host can be reflected as a
biological shelter for parasites. Unlike endoparasites, ectoparasites
are in\selectlanguage{english}fluenced not only by host characteristics, but also by
characteristics of the host environment (Krasnov \emph{et al} ., 1997).
Therefore, a habitat of the ectoparasite should not be just a particular
host, but a particular host in a particular habitat. If so, an important
determinant of parasite community structure should be a complex
host-habitat relationship. Quantifying variation of ectoparasites load
among host species and comparing these variation with other ecological
factors which are known to shape host communities is the good approach
for understanding dual nature of host-parasite interaction
(Merrill\emph{et al} ., 2020; Veitch \emph{et al} ., 2020).

Concisely, the aim of this study was to examine rodent's flea load and
their associated factors in plague endemic area of Karatu district by i)
quantifying and assessing the variation of flea's parasitological
indices (specific flea index, total flea index and flea infestation
prevalence on rodents), ii) assessing the association of abundance of
species of fleas with rodent species, plague and non-plague foci
villages, season (wet and dry season), habitat type and rodent sex, iii)
investigating flea biased parasitism on rodent's weight and sex (male
and female rodents) and lastly, iv) assessing the influence of host
characteristics (rodent species, sex and weight), habitat type, season
and plague locations on abundance of fleas. Result of this study will
help to create awareness to the public health sectors and vector control
programs by providing valuable information that will help in decision
making process regarding flea control in plague endemic area of Karatu
district, northern Tanzania.

\textbf{3.3 Material and methodology}

\textbf{3.3.1 Study area}

This study was conducted at Karatu district, Arusha region in northern
part of Tnzania. The district is bordered by Monduli district to the
east, Shinyanga region to the west, Ngorongoro district to the north,
and Manyara region to the south and southeast. The study was conducted
in four villages found in Karatu district which are Rhotia Kati
3deg19'16.4''S 35deg44'22.3''E, Kambi ya Simba 3deg17\selectlanguage{english}'05.28"\selectlanguage{english}S 35\selectlanguage{ngerman}°48\selectlanguage{english}'30.84"\selectlanguage{english}E,
Kitete 3\selectlanguage{ngerman}°13'58.5''S 35°50'30.8''E and Marera 3°18'32.2''S 35°43'26.6''E.
Rhotia Kati and Kambi ya Simba are plague foci villages while Marera and
Kitete are non-plgue foci villages (Kilonzo \emph{et al} .,
2006)\textbf{(Figure 1 below)} . Economic activities of residents in the
study area involve crop cultivation, livestock keeping and small retail
shops.

\textbf{Figure 1} : Showing map of Tanzania highlighting Rhotia and
Mbulumbulu wards where four villages were selected for rodent sampling
in this study.

\textbf{3.3.2 Selection of study area}

In this study four villages were selected depending on the information
of whether been involved or not involved in the past human plague
outbreak (Kilonzo \emph{et al} ., 2006). Rodents were sampled from four
habitats that include, farm land, bush land, peridomestic area and
forest buffer zone that were selected purposefully from each village.
Selection criteria of habitats was relying on the following factors; a
habitat characterized with bushes comprising shrubs and little dwarf
scattered trees away from settlements were considered as bush land
habitat, a land cultivated with cereal and other agricultural crops were
considered as farm land habitat, a forest buffer zone habitat was
declared by the conservation authority where they provided permission
and guidance to trap in the area. Forest buffer zone was defined as a
part of protected land adjoining conservation area and community area
with no human activities allowed in that area. Peridomestic area was
considered to be an area of land surrounding human settlement within a
range of one hundred meter. Bush land, farm land and forest buffer zone
were all considered as sylvatic areas and were selected at least 300 to
500 m away from the human settlements.

\textbf{3.3.3 Study design and sample size}

This study was repeated cross sectional design involving collection of
rodent sample across season (wet and dry season). Preliminary survey was
conducted in a study area one week before rodent sampling begun to
request permission and inform authority about this study. Sherman live
animal traps were used to capture rodents in farm land, bush land,
peridomestic areas and forest buffer zone.

Rodent sample size was estimated with formula described by Naing\emph{et
al} . (2006)

\begin{equation}
n=\frac{Z^{2}\text{\ P\ }\left(1-P\right)}{d^{2}}\nonumber \\
 \end{equation}

Whereby n= sample size; Z= test statistics at 95\% confidence
level;\emph{P} = expected prevalence of \emph{Yersinia pestis} in
rodents; (1-\emph{p} ) = probability of having no infected rodent;
\emph{d} = desired level of precision. Values in the formula are:
Z=1.96, \emph{d} = 5\% and \emph{P} = 10\%. By inserting the values into
the formula\((\frac{\par
\begin{matrix}{1.96}^{2}&amp;\times 0.1(1-0.1)\\
\end{matrix}}{{0.05}^{2}}\)= 138.29). Therefore, sample size for this
study was estimated to be 140 rodents in each season (wet and dry)
giving a total sample size of 280 rodents.

\textbf{3.3.4 Rodent trapping and processing}

A total of 120 Sherman traps baited with peanut butter and mixed with
maize flour were used to capture rodents in plague and non-plague foci
villages in farm land, bush land, peridomestic and forest buffer zone
habitats. Selection of trapping sites was made successfully with the
help of the residents and involved observation of rodent's signs such as
rodent's pathway, droppings, movements sign or observation of gnawed
seeds, and depending on the information from the residents about
presence of rodents in their surroundings. Information about this study
was clearly provided to the owner of trapping site for requesting
permission before trapping. 35 traps were set at a distance of 10 m
apart in seven transect lines containing five trapping station in all
habitats except in peridomestic areas were 35 Sherman traps were set at
2 to 5 m apart by targeting areas with rodent burrows or other rodent
signs around human houses and livestock shelters. Traps were usually
sets on the field in the evening at 5:00 pm and left overnight for at
least three consecutive days while inspecting the traps for captured
rodents every day during morning and afternoon.

Captured rodents were carefully removed from the trap using animal
handling bag and were anaesthetized using diethyl ether in a bottle
contained cotton wool. Blood sample were collected from anaesthetized
rodents for making blood smear for observation of \emph{Yersinia
pestis}coccobacilli under light microscope; organ such as lugs and
spleen were collected from the euthanized rodent for confirmation of
(\emph{pla} ) gene of \emph{Y.pestis} DNA under qPCR (results in another
manuscript). Fleas were collected from a euthanized rodent before
dissection by brushing their body thoroughly over a plastic basin
(Makundi \emph{et al} ., 2008). Rodents were brushed from the head to
the base of the tail using small shoeshine brush until all fleas stop to
fall in a basin. Animal handling bag and anaesthetizing bottle were
checked for presence of fleas and were all collected with a fine
forceps. Collected fleas were counted, recorded and preserved in a well
labeled eppendorf tubes containing 70\% ethanol and stored at room
temperature until morphological identification using dichotomous key in
the laboratory.

\textbf{3.3.5 Rodent identification}

Rodents were identified to genus and specie level using an established
taxonomic nomenclature of rodents identification (Happold, 2013).
Identification involved observation of rodent's fur color and
morphometric measurements that were taken in millimeters using digital
vernier caliper to measure length of hind foot (\emph{pes} ), ear
length, tail length and head to body length and also the rodent's weight
were measure in gram by using digital weighing balance.

\textbf{3.3.6 Fleas processing and identification}

Fleas were processed in the laboratory following standard procedure in
(Hastriter and Whiting, 2003) cited in (Berrizbeitia \emph{et al} .,
2017), involving passing fleas through series of reagents to make their
feature clear enough to be identified with dichotomous key. After
processing, fleas were examined under digital stereo microscope
OPTA-TECH\selectlanguage{ngerman}\textsuperscript{®} and identified to genus and specie level
following conventional dichotomous key as described in
(Harimalala\emph{et al} ., 2021; Friggens \emph{et al} ., 2020). Fleas
were then mounted on a microscope glass slides in accordance with
conventional procedures for fleas processing before identification.

Flea processing/clearing were involving puncturing of fleas at the
region between abdominal sterna II and III using thin-fine needle after
placing them on a wax-block. Punctured fleas were soaked in potassium
hydroxide (10\%) for 24 h to decolorize their dark color and make their
features visible for identification. After this process fleas were
immersed in distilled water for 30 min to wash excess potassium
hydroxide and stop decolonization process. After 30 min fleas were
removed and gently compressed on their abdomen to expel marinated soft
tissues throughout the punctured hole followed with dehydration process
in a series of ethanol solutions (70\%, 80\%, 95\% and absolute) for 30
min in each step.

For clarification of flea's exoskeleton, fleas were immersed in methyl
salicylate for 15--20 min and then transferred to xylene for a minimum
of 1 h. After then, they were mounted on a microscope glass slide in a
Canada balsam (Campbell \emph{et al} ., 2018) which finalized the
process and fleas were ready for identification. Fleas voucher specimen
of each species identified were reserved at college of veterinary
medicine and biomedical sciences at Sokoine University of Agriculture
(SUA), Morogoro Tanzania.

\textbf{3.3.7 Data analysis}

Flea data were recorded in Microsoft Excel 2016 and prepared in a
readable data set format, data set were imported in R programming
language (R Core Team version 4.2.2 of 2022) from which all statistical
analyses were conducted with a significance level (\selectlanguage{greek}α\selectlanguage{english}) of 0.05. Data were
tested for normality before analyzed and were found not normally
distributed (Shapiro-Wilk test \emph{p} \textless{}.05) leading us to
use non-parametric test for inferential statistics. Rodent's flea load
on examined rodents were evaluated by assessing total flea indices and
specific flea indices (Zimba \emph{et al} ., 2012). Percentage
prevalence of rodents infested with fleas (proportion of examined
rodents positive for fleas) were computed with 95 \% confidence interval
for proportion and multiplied by 100\%.

Chi-square \selectlanguage{greek}χ\selectlanguage{english}\textsuperscript{2} test of association was applied to
examine the relationship of abundance of species of fleas with rodent
species, plague and non-plague foci villages, season, habitat type and
rodent sex. The nature of association was further analyzed to find the
contribution of level of factors to the Chi-square result using absolute
standardized residuals and the relationship were presented in balloon
chart (Kassambara, 2022). Flea biased parasitism on rodent's sex and
weight was analyzed by assessing the variation of flea abundance between
sexes using Wilcoxon rank sum test and through evaluation of correlation
between flea abundance and rodent's weigh using Spearman rank
correlation test (rho) respectively; these factors were further analyzed
in generalized linear model (GLM) to assess their influence on flea
abundance. The GLM was used to establish the model variables that
described the influence of predictors of abundance of flea. Model was
selected based on AIC in which model with possible lowest AIC was
selected with help of step-AIC function in MASS package of R programming
language (Venables and Ripley, 2002). Predictor variables were both
numerical and categorical variables that include, host characteristics
(rodent species, sex and weight), habitats, season and plague locations.

\textbf{3.4 Results}

\textbf{3.4.1 Flea collected from rodents}

Four species of fleas (n= 190) were collected from six species of
rodents (n=73/291) from a total of nine rodent species that were
captured with Sherman animal live traps in plague and non-plague foci
villages across wet season (January-February) and dry season (July) of
2022. We found that total flea index in wet season was 0.53 (SD=1.54),
dry season was 0.78 (SD= 1.66), plague foci was 0.71 (SD= 1.81) and
non-plague foci was 0.55 (SD= 1.13). Flea species that were collected in
this study were \emph{Dinopsyllus lypusus} (46.32 \%)\emph{,
Ctenophthalmus spp} (26.84 \%), \emph{Xenopsylla brasiliensis} (16.32
\%) and \emph{Xenopsylla cheopis} (10.53 \%). Collected fleas were
evaluated to estimate total flea indices (TFI) and prevalence of flea
infestation among rodent species \textbf{(table 1)} . Apparently, there
was no flea collected from three species of rodents \emph{Grammomys spp,
Praomys delectorum} and \emph{Otomys spp} .

\textbf{3.4.2 Variation of flea infestation prevalence on rodent
community}

We found that estimated prevalence of all rodents infested with fleas
was 25 \% (CI= 20 \% - 30 \%) and was statistically significant lower
than 50 \% of infestation prevalence \emph{p} \textless{} .001.
Variations of prevalence of rodents infested with fleas were estimated
across different habitats, plague locations and season. It was found
that farm land and peridomestic area had high prevalence of fleas 30 \%
(CI= 21\% -40\%) and 26 \% (CI= 18\% -35\%) respectively compared to
forest buffer zone and bush land 24\% (CI= 14\% -38\%) and 17\% (CI=
09\% -29\%) respectively. Plague foci villages had high fleas prevalence
26\% (CI= 20\% -33\%) compared to non-plague foci villages 23\% (CI=
16\% -32\%). Dry season had high fleas prevalence 30\% (CI= 23\% -38\%)
compared to wet season 20\% (CI= 15\% -27\%).

\textbf{3.4.3 (i) Specific flea indices (SFI) on rodents in different
habitats}

Fleas collected from all rodent species were involved in the estimation
of specific animal flea indices across different habitats \textbf{(table
2)} . Generally \emph{D. lypusus} infested five species of rodents
covering (42.47\%) of all infested rodents
(n=73/291);\emph{Ctenophthalmus spp} infested four species of rodents
(32.88\%),\emph{X. brasiliensis} infested three species of rodents
(12.33\%) and\emph{X. cheopis} infested two species of rodents
(12.33\%).

\textbf{3.4.3 (ii) Variation of specific flea indices (SFI) on rodents
in plague and non-plague foci}

Rodents in non-plague foci villages were mostly infested
with\emph{Ctenophthalmus spp} and \emph{D. lypusus} \textbf{fig (2A and
2B)}below from which \emph{Ctenophthalmus spp} was mostly infested two
rodent species i.e. \emph{L.striatus} and \emph{M.natalensis}
while\emph{D.lypusus} was mostly infested \emph{G.murinus} compared to
other captured rodents species. In plague foci villages most captured
rodents were infested with \emph{X.cheopis} and
\emph{X.brasiliensis}\textbf{fig (2C and 2D)} . \emph{X.brasiliensis}
and \emph{X.cheopis}were mostly infested \emph{R.rattus} than other
rodent species.

\textbf{Fig 2} . Bar charts showing comparison of specific animal flea
index (SFI) on rodent species collected in plague and non- plague foci
villages in plague endemic area of Karatu district northern Tanzania.
The above numbers on bar chats are percentage trap success of each
individual rodent.

\textbf{3.4.4 Association of abundance of flea species with other
factors}

We found that abundance of flea species was statistically significant
associated with type of habitat (\selectlanguage{greek}\emph{χ2}\selectlanguage{english} (9) =
41.701,\emph{p\textless{}} .001), season (\selectlanguage{greek}\emph{χ2}\selectlanguage{english} (3) =
17.839,\emph{p\textless{}} .001), rodent species (\selectlanguage{greek}\emph{χ2}\selectlanguage{english} (15) =
134.35,\emph{p\textless{}} .001) and rodent sex (\selectlanguage{greek}\emph{χ2}\selectlanguage{english} (3) =
10.543,\emph{p=} .014), but was not statistically significant associated
with plague locations (plague and non-plague foci) (\selectlanguage{greek}\emph{χ2}\selectlanguage{english} (3) =
2.4006,\emph{p=} 0.49). Balloon chart was plotted to describe this
relationship by using Pearson residuals (standardized residuals) of all
level of each factor. The high value of Pearson residual was described
with the big and bright red balloon in a gradient of size and color from
dark to pale red suggesting particular level of factor had strong
association with abundance of particular flea specie and a gradient of
small balloons of white to bright blue suggesting weak or insignificant
association of abundance of particular flea specie \textbf{(fig 3)} .

\textbf{Fig3} . Balloon chart showing association of abundance of fleas
species with other factors; the abbreviations Cteno=\emph{Ctenophthalmus
spp} , Dino= \emph{Dinopsyllus lypusus} . X.br=\emph{Xenopsylla
brasiliensis} and X.cheo= \emph{Xenopsylla cheopis} .

\textbf{3.4.5 Flea biased parasitism on rodent's sex and weight}

We found statistical significant difference in flea abundance between
male and female rodents (Wilcoxon rank sum test W = 9158.5, \emph{p} =
.009) with small magnitude of effects size (r= .153) (Rules: funder
2019). The median weight of these two sexes was 40.1g (IQR= 25.3g) and
39.4g (IQR= 23g) for male and female rodents respectively and were not
statistically significant different (W = 10140, \emph{p} = .54).
Correspondingly, we found statistical significant correlation between
flea abundance and rodent's weight with Spearman's rank correlation
coefficients (\emph{R} = .17, \emph{S} = 34176 and \emph{p} \textless{}
.05) indicating weak positive correlation \textbf{(fig. 4 A)} .

\textbf{3.4.6 Factors influencing flea abundance}

A best final fitted model obtained from the GLM (Poisson family) was
statistically significant to explain the relationship of predictor
variables to the abundance of flea \selectlanguage{greek}χ\selectlanguage{ngerman}² (15) = 106.63, \emph{p}\textless{}
.001, model coefficients in \textbf{(table 3)} , predictors chart in
\textbf{fig. 4 (4B-4F)} showing the probability of fleas in different
factors.

\textbf{Figure 4} . (A) Scatter plot for Spearman's rank correlation
test of rodent's weight and flea abundance. (B) is the GLM prediction
curve of influence of rodent's weight on abundance of fleas. (C-F) are
the GLM categorical predictor variables with their influence to
abundance of fleas.

\textbf{3.5 Discussion}

\textbf{3.5.1 Flea species collected from rodents}

Four species of fleas \emph{X.brasiliensis, X.cheopis, D.lypusus}
and\emph{Ctenophthalmus spp} that were collected from rodent community
in both plague and non-plague foci villages are all considered as
potential or likely to be potential for harboring and transmitting
plague infection from rodent to rodent or rodent to human being during
the past human plague outbreak in Karatu plague endemic area and
elsewhere, however their efficiency in plague transmission is different
(Kilonzo\emph{et al} ., 2006; Eisen \emph{et al} ., 2007; Makundi
\emph{et al} ., 2008). The most efficient flea vector of plague
infectious disease is\emph{X.cheopis} that serves in circulating plague
bacilli in both enzootic and epizootic plague periods (Eisen and Gage,
2009). The high vectoring potential of \emph{X.cheopis} is depending on
its exclusive characteristic of proventricular spines that provides
attachment area for colonization of \emph{Y.pestis} leading into
formation of proventricular blockage after taking blood meal from an
infected host, a situation which make \emph{X.cheopis} to increase its
daily biting rate for sucking blood meal with multiple regurgitation on
the biting site on an attempt to unblock its proventriculus; therefore
facilitating the increase of transmission of infection compared to other
fleas (Gage and Kosoy, 2005; Korzun and Nikitin, 1997).

Similarly, \emph{D.lypusus} and \emph{Ctenophthalmus spp} are conceived
as enzootic plague vectors and they also facilitate rapid transmission
of the diseases during plague outbreak (Eisen and Gage, 2009;
Ziwa\emph{et al} ., 2013; Enscore \emph{et al} ., 2020).
\emph{D.lypusus} and\emph{X.brasiliensis} were conveyed as competent
vectors for transmission of plague disease during 2007 plague outbreak
in Mbulu and (1996/7) in Karatu plague endemic area (Kilonzo \emph{et
al} ., 2006; Makundi \emph{et al} ., 2008). Furthermore,
\emph{D.lypusus} were reported in East Africa as a plague vector and an
important enzootic flea in the maintenance of \emph{Y.pestis} (Devignat,
1949; Arap\emph{et al} ., 1977; Kilonzo 1992). This suggested that these
flea species \emph{X.brasiliensis, X.cheopis} and \emph{D.lypusus} are
important fleas in the transmission and maintenance of plague bacilli in
plague endemic foci of Karatu district.

\textbf{3.5.2 Specific flea index (SFI)}

Since oriental rodent fleas (\emph{X.cheopis} ) received high public
health concern regarding their efficiency in amplification and
transmission of \emph{Y. pestis} from rodent to rodent and from rodent
to human (Eisen and Gage, 2009; Maestas and Britten, 2017), we have
evaluated their specific flea index as one of the potential factor that
may cause risk for epizootic plague among rodents and eventually to the
society (Gage, 1999; Eisen \emph{et al} ., 2012, 2020). It has been
reported that a specific flea index \textgreater{}1 for
\emph{X.cheopis}on rats represents a potentially dangerous situation
with respect of increased plague risk to human in the event of an
outbreak of plague (Gage, 1999). Similarly an outbreak of human plague
is more likely to happen if the specific flea index is \textgreater{}5
(Singchai \emph{et al} , 2003).

Based on result obtained in this study, we found that specific flea
index of \emph{X.cheopis} was equal to one (1.0 SFI) in plague foci
villages and less than 0.5 (\textless{}0.5 SFI) in non-plague foci
villages. Despite that, these indices do not really call for urgent
control of fleas in these areas, it provide the basic information to
understand how plague foci villages require more attention to prevent
situation from exceeding the risk level. The high specific flea index in
plague foci villages is probably due to high abundance of host of
oriental rodent fleas i.e. \emph{Rattus rattus} and less
frequently\emph{Mastomys natalensis} as compared to non-plague foci
villages.

\textbf{3.5.3 Associated factors influencing flea abundance on rodents}

Basically we found that flea abundance on rodent community were
influenced by variation of season of the year (wet and dry season),
plague and non-plague foci villages, variation of habitat types and host
characteristics. Among all these factors, variation of season of the
year has the major impact on flea abundance as it determine distribution
of food and breeding cycle of their rodent host (Leirs \emph{et al} .,
1996). Change in temperature and humidity of the year in response to
changing season (wet and dry season) is significantly controlling the
growth and survival of immature fleas (Ziwa \emph{et al} ., 2013).
Different studies have reported the influence of seasonal change on
abundance of rodents and fleas to have the impact on maintenance and
transmission of plague disease (Njunwa \emph{et al} ., 1989;
Makundi\emph{et al} ., 1994; Eads \emph{et al} ., 2016; Shuai \emph{et
al} ., 2022).

Variation of wet and dry season is generally affecting growth,
reproduction and survival of immature stage of fleas (Krasnov \emph{et
al} ., 2001; Ngeleja \emph{et al} ., 2017). Low abundance of fleas
during wet season could be explained by the fact that, the survival of
immature fleas in rodent burrows is affected by soil moisture which is
controlled by precipitation outside the burrow. Excessive wet conditions
in rodent burrows contained with organic matter at a relative humidity
\textgreater{}95\% can promote growth of vicious fungi that reduces
larval and egg survival (Eisen, 2009; Ben Ari \emph{et al.} , 2011). The
increase in flea abundance during dry season was probably due to the
increase in the abundance of their rodent host as well as the supportive
weather with moderate warm and moist condition during early dry season.
Reportedly, rodent's flea abundance is mainly affected by ambient
temperatures, precipitation, and relative humidity, where as warm-moist
weather is providing the explanation for higher flea indices (Ben
Ari\emph{et al.,} 2011; Wale \emph{et al} ., 2023).

Plague foci villages was significantly increasing abundance of fleas
compared to non-plague foci villages. The increase of flea abundance
could be encouraged by high proportion of abundance of rodents in plague
foci that presented many potential biological habitats and source of
blood meal for fleas. The increase in abundance of rodents influences
the increase in abundance of fleas and subsequently enhances the
distribution and composition of flea communities (Krasnov \emph{et al}
., 2002; Eads \emph{et al} ., 2016).

Habitat type was another factor that influenced abundance of fleas in
our study area. Different habitats have different characteristics that
are important for determining distribution and abundance of rodent host
of fleas. As reported by Brinkerhoff, (2008) and Laudisoit et al.
(2009), the change of habitats is affecting the composition of rodent
species and their fleas, making it an important factor in ecological
surveillance of abundance of flea species on rodents. In this study we
found that abundance of fleas was significantly increasing in farm lands
as compared to other habitats. Farm land habitat was observed to
encourage rodent colonization as it promote availability of food such as
maize and wheat seeds left after harvest during dry season. Habitually
rodents prefer to reside in areas with adequate foods where they can
make burrows and nest to protect them and their young from predation.
Habitats with high food availability and less disturbance and control
facilitate the increase in abundance of rodents and their fleas.

Moreover, we found that different rodent species have different
influence on flea abundance. \emph{Rattus rattus} was significantly
influencing abundance of fleas as compared to other rodents. The
characteristics of individual rodent host have been reported to affect
responsible mechanism for flea acquisition (Kiffner \emph{et al} .,
2013). Behavior and tendency of \emph{R.rattus} to live in human
habitats facilitates the interactions and sharing of fleas among them
due to existence of small range of this habitat. Movements
of\emph{R.rattus} from one house to another and from nearby farms and
livestock shelters is also predisposing this specie to encounter many
fleas which are subsequently shared to other \emph{R.rattus} . In
addition, human habitats provide suitable environment with availability
of food and warm condition supporting growth and reproduction of both
rodent and fleas especially when the control measures are abandoned
leading to increase flea index among rodents in this habitat. Tactics
employed by fleas and many other ectoparasites to colonies microhabitats
of the host species (such as rodent's burrows) or wait until a suitable
host is present helps parasites to infest host easily especially a new
born or when new host has visited the burrow (Bitam \emph{et al} .,
2010).

\textbf{3.5.4 Flea biased parasitism on rodent's sex and weight}

We observed that sex biased fleas infestation was frequently recorded on
male rodents than females; male rodents were significantly increasing
flea infestation load. A study conducted by Moore and Wilson (2002) has
shown that arthropods (especially sticking ectoparasites) obviously
exhibit male-biased parasitism because most of them wait for the host to
visit their area rather than them searching for the host. An animal that
explore many habitats and visit many ecological niches has high
probability to encounter many parasites frequently compared to less
explorative animal with small movements within its ecological niche. Our
result is arguably with different studies (Gaines and McClenaghan, 1980;
Bitam \emph{et al} ., 2010 and Buchholz and Dick, 2017) which also
observed a similar relationship with males having high flea infestation
than female because of their large home range and wide dispersal area as
compared to female rodents. The little difference in weight between male
and female rodents obtained in this study could or could not influence
flea sex biased parasitism on rodents. Sex biased parasitism should not
be taken as a general rule since in other study elsewhere it was found
that in some host-parasite relationship both male and female rodent
hosts were found equally infested with fleas (Wirsing \emph{et al} .,
2007; Kiffner \emph{et al} ., 2011)

Similarly, weight of rodents in the general community was significantly
influencing the increase in abundance of fleas. Under natural phenomenon
weight of rodents is equivalent to the size and age of respective rodent
specie. The change of these parameters affects the physiological and
behavioral status of the individual rodent in particular ecological
habitat whereby the increase of these parameters could facilitate
rodents to become more explorative with large home range and dispersion
in searching for food, mates or better shelter and in the meanwhile they
keep increasing the frequency of encountering many fleas.
Correspondingly, large rodents are easy target for ectoparasites
compared to small one, they can also tolerate many fleas, while the
youngest one would perform anti-parasitic grooming frequently
(Wilson\emph{et al} , 2002; Hawlena \emph{et al} ., 2008).

\textbf{3.6 Conclusion and recommendation}

This study was intended to contribute knowledge about rodent's flea in
plague and non-plague foci villages of plague endemic area of Karatu
district, northern Tanzania. In this study we found that plague foci
villages was leading with high abundance of rodents and fleas with
reservoir potential for plague disease compared to non-plague foci
villages. Also the result of specific flea index for \emph{X.cheopis} on
rats in both plague and non-plague foci villages did not exceed the
risky level i.e. (\textgreater{}1 potentially dangerous situation and
\textgreater{}5 urgent situation as an outbreak of human plague is more
likely to happen), from which the society is advised to keep
implementing rodent and flea control methods in their surroundings that
help in reducing the interaction of these plague agents with human
society especially in farm lands and peridomestic areas. Furthermore
this study recommends that, for effective prevention of emergence of
plague disease and other flea borne zoonotic diseases, it is advisable
to make regular surveillance of abundance of rodents and fleas in
potential plague endemic areas while monitoring strategies placed for
plague prevention and their implementation in the society in an attempt
of improving them or replacing them with a modern one.

\textbf{Acknowledgements}

We are extremely grateful to all the people whose extensive efforts made
this study successful. We would like to express our sincere gratitude to
Africa Centre of Excellence for Innovative Rodent Pest Management and
Biosensor Technology and Development (ACE IRPM\&BTD) for funding this
research work. We also acknowledge Mr Omary Milango a field assistant
and rodent technician of Institute of Pest Management at Sokoine
University of Agriculture, Morogoro Tanzania who assisted in rodent data
collection and Miss Miriam Jophrey Bukuku and Mr Mwita Wambura of
department of microbiology, parasitology and biotechnology at SUA who
assisted in flea processing and identification.

\textbf{Conflict of interest statement}

No one declared conflict of interest in this study.

\textbf{Ethical approval}

This study was approved by the Institutional Review Board of Sokoine
University of Agriculture, Tanzania (Ref. No. SUA/DPRTC/R/186/15).

\textbf{Data availability statement}

Authors agree to deposit the data associated with this study in a
Mendeley data repository and make it publicly available once the
manuscript is accepted for publication under the Journal of Ecology and
Evolution. Data reference link: Jakoniko, J., Massawe, A., Mwega, E.,
Kessy, S., (2023),~``Contributing factors for abundance of fleas on
rodents in plague endemic area. Karatu District, Northern
Tanzania'',~Mendeley Data,~v1http://dx.doi.org/10.17632/mydn68pyzz.

\textbf{ORCID}

\emph{Joshua R. Jakoniko} https://orcid.org/0009-0003-5462-9840

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\textbf{Table1} . Fleas collected from rodents, showing TFI and
percentage prevalence of infestation\selectlanguage{english}
\begin{longtable}[]{@{}lllll@{}}
\toprule
\textbf{Rodent species} & \textbf{No. of rodents examined (n \%)} &
\textbf{No. of rodents infested with fleas (prevalence)} & \textbf{No.
of fleas collected (n \%)} & \textbf{TFI}\tabularnewline
\midrule
\endhead
\emph{Arvicanthis niloticus} & 70 (24.05) & 12 (0.17) & 27 (14.21) &
0.39\tabularnewline
\emph{Grammomys spp} & 12 (4.12) & 0 & 0 & 0\tabularnewline
\emph{Graphiurus murinus} & 2 (0.69) & 1 (0.5) & 3 (1.58) &
1.5\tabularnewline
\emph{Lemniscomys striatus} & 43 (14.78) & 13 (0.3) & 43 (22.63) &
1\tabularnewline
\emph{Lophuromys flavopunctatus} & 13 (4.47) & 4 (0.31) & 9 (4.74) &
0.69\tabularnewline
\emph{Mastomys natalensis} & 122 (41.92) & 31 (0.25) & 73 (38.42) &
0.6\tabularnewline
\emph{Otomys spp} & 3 (1.03) & 0 & 0 & 0\tabularnewline
\emph{Praomys delectorum} & 1 (0.34) & 0 & 0 & 0\tabularnewline
\emph{Rattus rattus} & 25 (8.60) & 12 (0.48) & 35 (18.42) &
1.4\tabularnewline
\textbf{Total} & \textbf{291} & \textbf{73 (0.25)} & \textbf{190} &
\textbf{0.65}\tabularnewline
\bottomrule
\end{longtable}

\textbf{Table2} . Rodent specific flea indices (SFI) across different
habitats\selectlanguage{english}
\begin{longtable}[]{@{}llllllll@{}}
\toprule
\begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
\textbf{Habitats}\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
\textbf{Rodent species}\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
No. rodents examined\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
No. rodents infested (prevalence)\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
\textbf{No. fleas collected and (SFI)}\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
\textbf{No. fleas collected and (SFI)}\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
\textbf{No. fleas collected and (SFI)}\strut
\end{minipage} & \begin{minipage}[b]{0.12\columnwidth}\raggedright\strut
\textbf{No. fleas collected and (SFI)}\strut
\end{minipage}\tabularnewline
\midrule
\endhead
\begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{X. brasiliensis}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{X. cheopis}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{D. lypusus}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{Ctenophthalmus spp}\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Bush land}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\emph{A. niloticus}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
27\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
4 (0.15)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
6 (0.22)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
3 (0.11)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{Grammomys spp}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
5\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{Lemniscomys striatus}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
7\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{Mastomys natalensis}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
12\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
5 (0.42)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
4 (0.33)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
1 (0.08)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{Otomys spp}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
2\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Farm land}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\emph{A. niloticus}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
10\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
2 (0.2)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
4 (0.4)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{Grammomys spp}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
1\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{Lemniscomys striatus}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
29\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
10 (0.34)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
4 (0.14)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
19 (0.66)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
12 (0.41)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{L. flavopunctatus}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
2\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
1 (0.5)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
2 (1)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{M. natalensis}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
45\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
13 (0.29)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
3 (0.07)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
3 (0.07)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
16 (0.36)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
15 (0.33)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Forest buffer zone}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\emph{Grammomys spp}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
5\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{G. murinus}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
2\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
1 (0.5)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
3 (1.5)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{L. flavopunctatus}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
11\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
3 (0.27)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
6 (0.55)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
1 (0.09)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{M. natalensis}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
27\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
7 (0.26)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
6 (0.22)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
8 (0.3)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
3 (0.11)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{P. delectorum}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
1\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\textbf{Peridomestic}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
\emph{A. niloticus}\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
33\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
6 0.18)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
5 (0.15)\strut
\end{minipage} & \begin{minipage}[t]{0.12\columnwidth}\raggedright\strut
9 (0.27)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{Grammomys spp}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
1\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{L. striatus}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
7\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
3 (0.43)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
7 (1)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
1 (0.14)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{M. natalensis}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
38\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
6 (0.16)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
8 (0.21)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
6 (0.16)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{Otomys spp}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
1\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\emph{Rattus rattus}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
25\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
12 (0.48)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
18 (0.72)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
17 (0.68)\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
0\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\textbf{Total}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\textbf{291}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\textbf{73 (0.25)}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\textbf{31}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\textbf{20}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\textbf{88}\strut
\end{minipage} & \begin{minipage}[t]{0.10\columnwidth}\raggedright\strut
\textbf{51}\strut
\end{minipage}\tabularnewline
\bottomrule
\end{longtable}

\textbf{Table 3} . Summary of the best fitting coefficients of GLM
(Poisson family) describing the influence of predictor variables
(parameters) on abundance of fleas.\selectlanguage{english}
\begin{longtable}[]{@{}lllll@{}}
\toprule
\begin{minipage}[b]{0.17\columnwidth}\raggedright\strut
\textbf{Parameters}\strut
\end{minipage} & \begin{minipage}[b]{0.17\columnwidth}\raggedright\strut
\textbf{Estimate}\strut
\end{minipage} & \begin{minipage}[b]{0.17\columnwidth}\raggedright\strut
\textbf{Std. error}\strut
\end{minipage} & \begin{minipage}[b]{0.17\columnwidth}\raggedright\strut
\textbf{Z value}\strut
\end{minipage} & \begin{minipage}[b]{0.17\columnwidth}\raggedright\strut
\emph{p}\textbf{-value}\strut
\end{minipage}\tabularnewline
\midrule
\endhead
\begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Intercept\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
-2.48\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.41\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
-6.05\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\textless{} .001\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
\textbf{Rodent species} \emph{Grammomys} \emph{spp}\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
-14.96\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
573.36\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
-0.03\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
0.98\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\emph{G.murinus}\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
1.40\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.72\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
1.96\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.05\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\emph{L. striatus}\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.56\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.29\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
1.91\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.06\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\emph{L. flavopunctatus}\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.47\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.50\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.95\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.34\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\emph{M. natalensis}\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.24\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.25\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.93\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.35\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\emph{Otomys spp}\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
-14.91\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
1206.53\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
-0.01\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.99\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\emph{P. delectorum}\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
-15.46\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
2103.36\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
-0.01\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.99\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\emph{R. rattus}\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
1.34\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.29\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
4.66\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\textless{} .001\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
\textbf{Season} Wet season\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
-0.41\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
0.19\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
-2.15\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
0.03\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
\textbf{Habitats} Farm land\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
0.86\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
0.31\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
2.79\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
0.005\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Forest habitat\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.58\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.38\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
1.50\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.13\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
Peridomestic\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.39\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.32\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
1.21\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.23\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\textbf{Rodent Weight}\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.017\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
0.004\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
4.20\strut
\end{minipage} & \begin{minipage}[t]{0.17\columnwidth}\raggedright\strut
\textless{} .001\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
\textbf{Rodent sex} Male\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
0.53\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
0.15\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
3.46\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
\textless{} .001\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
\textbf{Plague location} Plague foci villages\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
0.40\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
0.18\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
2.20\strut
\end{minipage} & \begin{minipage}[t]{0.19\columnwidth}\raggedright\strut
0.03\strut
\end{minipage}\tabularnewline
\bottomrule
\end{longtable}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/FIGURE-1/FIGURE-1}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/FIGURE-2/FIGURE-2}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/FIGURE-3/FIGURE-3}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/FIGURE-4/FIGURE-4}

\end{center}
\end{figure}

\selectlanguage{english}
\FloatBarrier
\end{document}

