Advances in Parasitology, Volume 117

David Rollinson
Visit to download the full and correct content document:
https://ebookmass.com/product/advances-in-parasitology-volume-117-david-rollinson-
2/
Advances in Parasitology, 117

FIRST EDITION

David Rollinson
Life Sciences Department The Natural History Museum, London, United
Kingdom

Russell Stothard
Department of Tropical Disease Biology Liverpool School of Tropical
Medicine, Liverpool, United Kingdom
Table of Contents

Cover image

Title page

Copyright

Contributors

Chapter One: The microscopic five of the big five: Managing

zoonotic diseases within and beyond African wildlife protected
areas

Abstract

1: Introduction

2: The ‘Microscopic Five’

3: Challenges of BTb control at the wildlife-livestock interface:

The South African case study

4: Drivers of disease: The Kruger National Park case study

5: Disease knowledge gaps and lessons learnt from African

protected areas
 6: Communities and conservation

7: Conclusions

Acknowledgements

References

Chapter Two: Improving translational power in antischistosomal

drug discovery

Abstract

1: Filling the drug pipeline for schistosomiasis

2: Evaluating the importance of S. mansoni isolate origin for

early antischistosomal drug discovery

3: The S. mansoni mouse model for drug efficacy testing

4: Infection intensity of the patent S. mansoni mouse model

5: Pharmacokinetic/pharmacodynamic (PK/PD) relationship of

selected drugs

6: Concluding remarks

Acknowledgements and funding

References

Chapter Three: Unique thiol metabolism in trypanosomatids: Redox

homeostasis and drug resistance

Abstract

1: Introduction
2: Trypanothione metabolism

3: Effector proteins of the antioxidant defence: Old and new

actors

4: Trypanothione metabolism is linked to cysteine, polyamine,

and pentose phosphate pathway

5: The role of redox active compounds and their mechanism in

parasites survival

6: Role of thiol metabolism in drug resistance

7: Anti-parasitic potential of molecules targeted against redox

metabolism

8: Unsolved questions and future prospects

References
Copyright
Contributors
Chapter One: The microscopic five of the big five:
Managing zoonotic diseases within and beyond
African wildlife protected areas
Anya V. Tobera,⁎; Danny Govenderb,c; Isa-Rita M. Russoa,†; Jo Cablea,† a School of Biosciences, Cardiff University, Cardiff,
Wales, United Kingdom
b SANParks, Scientific Services, Savanna and Grassland Research Unit, Pretoria, South Africa
c Department of Paraclinical Sciences, University of Pretoria, Onderstepoort, South Africa
† Authors contributed equally to this work
⁎ Corresponding author: email address: tobera@cardiff.ac.uk.

Abstract
African protected areas strive to conserve the continent's great biodiversity with a targeted focus on
the flagship ‘Big Five’ megafauna. Though often not considered, this biodiversity protection also
extends to the lesser-known microbes and parasites that are maintained in these diverse
ecosystems, often in a silent and endemically stable state. Climate and anthropogenic change, and
associated diversity loss, however, are altering these dynamics leading to shifts in ecological
interactions and pathogen spill over into new niches and hosts. As many African protected areas
are bordered by game and livestock farms, as well as villages, they provide an ideal study system to
assess infection dynamics at the human-livestock-wildlife interface. Here we review five zoonotic,
multi-host diseases (bovine tuberculosis, brucellosis, Rift Valley fever, schistosomiasis and
cryptosporidiosis)—the ‘Microscopic Five’—and discuss the biotic and abiotic drivers of parasite
transmission using the iconic Kruger National Park, South Africa, as a case study. We identify
knowledge gaps regarding the impact of the ‘Microscopic Five’ on wildlife within parks and
highlight the need for more empirical data, particularly for neglected (schistosomiasis) and newly
emerging (cryptosporidiosis) diseases, as well as zoonotic disease risk from the rising bush meat
trade and game farm industry. As protected areas strive to become further embedded in the socio-
economic systems that surround them, providing benefits to local communities, One Health
approaches can help maintain the ecological integrity of ecosystems, while protecting local
communities and economies from the negative impacts of disease.

Keywords
Emerging Infectious Diseases; One Health; Zoonoses; Spill over; Spill Back; Kruger National Park

1: Introduction
As we enter the sixth mass extinction, protecting the world's biodiversity has never been more critical.
Protected areas, including national parks, cover over 18.8 million km2 and are at the forefront of a global
effort to safeguard biodiversity (Chape et al., 2003). Managers of these protected areas must strike a
balance between protecting the ecological integrity of ecosystems and preventing exploitation of local
resources while promoting their use in education and recreation (Chape et al., 2003). If managed
correctly, protected areas can be beneficial to wildlife conservation and the country's economy through
promoting ecotourism and creating local employment opportunities (Cheung, 2012; Spies et al., 2018).
However, the management of protected areas is challenging, particularly in the Anthropocene era of
human mediated global change, and increased emergence and re-emergence of infectious diseases
(reviewed by Cable et al., 2017). These diseases can reduce fitness, alter wildlife population
structure/size and even alter ecosystem function (Holdo et al., 2009; Prins and Weyerhaeuser, 1987;
Scott, 1988). Therefore, to effectively manage wildlife populations and ecosystems, it is essential to
understand the threats posed by pathogens and the diseases they cause.
Of the 3881 terrestrial and marine national parks in the world, almost half are in sub-Saharan Africa,
with terrestrial parks here covering 1 million km2 (4% of the total land area; Chape et al., 2003;
Muhumuza and Balkwill, 2013). These parks aim to conserve Africa's unique and iconic ecosystems
ranging from open savannas and grasslands to dense forest. This variety of habitats supports high levels
of biodiversity, drawing numerous tourists who aspire to spot the ‘Big Five’ megafauna: African buffalo
(hereafter referred to as buffalo), lion, African elephant (hereafter referred to as elephant), rhinoceros
and leopard (Dube and Nhamo, 2019). However, hidden and often forgotten biodiversity within
protected areas includes pathogens, which modulate animal abundance, fitness and behaviour (Gómez
and Nichols, 2013). It is crucial to better understand drivers for past and current wildlife disease
outbreaks within protected areas, to find new approaches to predict and prevent future outbreaks. A
review of all infectious wildlife diseases within protected areas would be too large a task. Instead, we
focus on five diseases referred to here as the ‘Microscopic Five’, which are important at the human-
livestock-wildlife interface due to their broad host range and zoonotic potential. These interface diseases
would all benefit from a ‘One Health’ approach to management (Fawzy and Helmy, 2019; Innes et al.,
2020; Webster et al., 2016). We therefore purposefully included high profile diseases (bovine tuberculosis
(BTb), Rift Valley fever and brucellosis) as well as neglected diseases (cryptosporidiosis and
schistosomiasis) for study. Using Kruger National Park, one of the most researched parks in Africa (van
Wilgen et al., 2016), we will review the key factors that can influence outbreaks and transmission of the
‘Microscopic Five’ within and around protected areas (Fig. 1). By focusing on a select group of
pathogens within a specific park our intention is to highlight drivers of disease common among many
protected areas and the importance of considering all infectious diseases in wildlife management plans.
We will first give a brief introduction to the ‘Microscopic Five’ and then give examples of the
environmental and anthropogenic factors driving the dynamics of these diseases within and around
Kruger National Park. We will then discuss the key knowledge gaps and future challenges for managing
the ‘Microscopic Five’ and other important diseases and touch on different management approaches
followed in various parks.
 FIG. 1 The ‘Big Five’ and ‘Microscopic Five’, and the drivers of disease at the wildlife-
livestock-human interface. Arrows represent anthropogenic drivers from beyond
Kruger National Park. Created with Microsoft PowerPoint (version 2109) and Adobe Photoshop (2021).

2: The ‘Microscopic Five’

The ‘Big Five’ are undoubtedly one of the biggest attractions for tourists visiting South Africa's
protected areas (Dube and Nhamo, 2019). To conserve these and other wildlife, and to reduce
transmission of infectious diseases among wildlife, domestic animals and humans, we focus on the
lesser known ‘Microscopic Five’. These comprise zoonotic diseases caused by pathogens that have
multiple hosts, including humans, and are of particular importance at the human-livestock-wildlife
interface. Although we focus on five specific diseases, there are many more of importance within
protected areas (Table 1) but by highlighting a distinct few we aim to raise the profile of all infectious
diseases and possible drivers. The first three of the ‘Microscopic Five’ (bovine tuberculosis, brucellosis
and Rift Valley fever) are high profile or state-controlled diseases in South Africa and any outbreaks
must be reported to the World Organisation of Animal Health (OIE). All three of these diseases are
trade-sensitive diseases and may change the trading status of a country and its ability to trade on the
global market. The remaining two (schistosomiasis and cryptosporidiosis) are neglected in comparison,
particularly in wildlife. By including these in the ‘Microscopic Five’, we aim to bring greater attention to
overlooked yet highly important diseases (see WHO, 2020). In the following account, we briefly cover
each of the ‘Microscopic Five’ discussing their host specificity, transmission pathways and known
impacts on wildlife, livestock and humans.
Table 1

Some diseases of large herbivores within Kruger National Park which may pose a threat to livesto
Spatial
Transmission Transmission
Disease. Pathogen Drivers distribution
modes routes
in KNP
Bacteria
Anthrax Bacillus Vector, Ingestion of Calcium soil Northern
anthracis environme contamina content, and
ntal, direct ted drought central
contact, vegetation regions
fomites or
carcasses
Bovine Tb Micobacteria Aerosol Respiratory Wildlife/livestock South,
bovis tract interface, host central,
density moving
north
Virus
Foot and mouth Aphtovirus Aerosol, Respiratory Wildlife/livestock North,
fomites tract interface, host central,
density south
African horse Orbivirus Mosquito Cutaneous Introduced Central
sickness vector, penetratio horses,
direct n season
contact
Rift Valley fever Phlebovirus Mosquito Cutaneous Climate change, Higher in
vector, penetratio drought, south
direct n rainfall and
contact central
regions
Protozoa
Bovine Brucella abortus Direct contact Ingestion of Host density North,
brucellosis infected central,
discharges south
during
birth,
milk,
mucus
membrane
s
Cryptosporidiosis Cryptosporidium Environmental Faecal-oral Dependant on Unknown
spp. via species and
contamina host range
tion of
food and
water
 Spatial
Transmission Transmission
Disease. Pathogen Drivers distribution
modes routes
in KNP

Piroplasma
Corridor disease Theileria parva Tick vector Cutaneous Wildlife/livestock North,
penetratio interface central,
n south
Babesiosis Babesia spp. Tick vector Cutaneous Unknown
penetratio
n
Digenea
Fascioliasis Fasciola spp. Environmental Contact with Dependant on Unknown
infected species and
water host range
Schistosomiasis Schistsoma spp. Environmental Contact with Dependant on Unknown
infected species and
water host range

2.1: Bovine tuberculosis

Bovine tuberculosis (BTb) is caused by the bacterium Mycobacterium bovis and predominantly infects
bovines, such as African buffalo (Syncerus caffer caffer) and cattle (Bos taurus), yet most warm-blooded
animals including humans can be infected (Ayele et al., 2004). Transmission mainly occurs through
inhalation of infectious particles, which is particularly problematic when livestock are kept at high
densities (Ayele et al., 2004). Though thought to have spilled over from cattle to buffalo in the early
1960s in South Africa (Bengis et al., 1996), buffalo now serve as the primary maintenance host for BTb
within Kruger National Park and Hluhluwe-iMfolozi Park, spilling over into various species of wildlife
and livestock (Michel et al., 2006). Although BTb is a controlled disease within South Africa, its control
is becoming increasingly challenging due to the presence of wildlife reservoirs, difficulty in controlling
disease in communal herds and lack of practical control options in wildlife (see Section 3). The WHO
estimated 147,000 new cases of zoonotic Tb in humans in 2016 with 12,500 deaths globally but mostly in
Africa (Sichewo et al., 2019b). Humans can become infected through drinking unpasteurised milk,
eating undercooked meat and via aerosols inhaled from infected cattle (DAFF, 2016; Sichewo et al.,
2019a).

2.2: Rift Valley fever

Rift Valley fever (RVF) is caused by a zoonotic, vector borne virus predominantly spread by Aedes
mosquitoes (Clark et al., 2018). The virus was first reported in South Africa in 1950 and subsequent
outbreaks have occurred sporadically every 7–11 years infecting mainly domestic livestock but also a
range of wild mammals and humans (Beechler et al., 2015a; Métras et al., 2015). Human infection occurs
mainly through direct contact with blood or tissue from infected animals or through consuming
unpasteurised milk but can also result from an infected mosquito bite. Symptoms vary from mild, flu-
like to severe haemorrhagic fever that can be fatal (Clark et al., 2018). Over 4000 human cases and
around 1000 deaths have been reported in the last 20 years, predominantly in Africa and Saudi Arabia
(Petrova et al., 2020). Little is known about how the pathogen is maintained during inter-epidemic
periods. One suggestion is vertical transmission from mosquitoes to their ova, which has been
demonstrated with Aedes mosquitoes under laboratory-controlled conditions (Romoser et al., 2011).
Another possibility is that it is maintained in wild animal populations (Beechler et al., 2015a; see Section
4.3.4). Commercial vaccines are available for livestock but there is currently no licensed human vaccine
(Petrova et al., 2020).

2.3: Brucellosis
Brucellosis, caused by bacteria of the Brucella genus, is ranked among the most economically important
zoonotic diseases globally. Although it is an OIE notifiable disease, outbreaks are thought to be greatly
under-reported in Africa (McDermott et al., 2013). The species of medical and veterinary importance are
Brucella abortus, Brucella melitensis and B. suis (see Ducrotoy et al., 2017). Infection in humans can lead to
a debilitating illness known as ‘Mediterranean’ or ‘undulant’ fever and is commonly misdiagnosed as
malaria (Ducrotoy et al., 2017; Godfroid et al., 2011). Human infection occurs through direct contact
with or consumption of an infected animal. Consumption of un-pasteurised milk causes most human
infections, while human to human transmission is rare (Godfroid, 2018). Several wildlife species have
been reported as seropositive for this disease and African buffalo are thought to be a reservoir for B.
abortus (see Godfroid et al., 2013). Infection can cause abortions in livestock reducing farm productivity,
however the effects of the disease on wildlife are largely unknown and may differ between species
(Gorsich et al., 2015). Vaccines are available for livestock and small ruminants but not yet for humans
(Ducrotoy et al., 2017).

2.4: Cryptosporidiosis
Cryptosporidiosis, caused by several species of the protozoan Cryptosporidium genus, can lead to severe
diarrhoea in humans and animals globally. Infectious diarrhoea is a major cause of death in children
under five in Africa and Cryptosporidium is second only to rotavirus as a contributor to this disease
(Kotloff et al., 2013; Squire and Ryan, 2017). Transmission occurs through the faecal oral route via close
contact with infected humans, animals or contaminated food and water (Innes et al., 2020). Currently
there are at least 40 recognised species with varying host specificities but the most important two
species infecting humans and livestock are C. hominus and C. parvum. The latter is the predominant
cause of diarrhoea in young calves and is the most important zoonotic species. Cryptosporidium parvum
is more genetically diverse than C. hominus with several subtypes with differing host specificities,
therefore an integrated genotyping approach has been advocated to differentiate these subtypes (Innes
et al., 2020). Cryptosporidium species have been identified in a range of wildlife, yet most studies focus on
humans and livestock (Zahedi et al., 2016). C. parvum, C. ubiquitum and C. bovis were recently identified
in wildlife within Kruger National Park in elephant (Loxodonta africana), buffalo (Syncerus caffer) and
impala (Aepyceros melampus; see Samra et al., 2011). Oocysts of Cryptosporidium spp. have also been
detected in zebra (Equus zebra), buffalo and wildebeest (Connochaetes gnou) faeces in Mikumi National
Park, Tanzania (Mtambo et al., 1997). There is currently no available vaccine for cryptosporidiosis yet
there is potential to develop one for cattle (Innes et al., 2020).

2.5: Schistosomiasis
Schistosomiasis is a waterborne, zoonotic disease of veterinary and medical importance, caused by
digenean parasites of the genus Schistosoma. Schistosomiasis is a major public health threat with an
estimated 207 million people infected and 779 million people at risk globally, with 90% of these
infections in Africa (Steinmann et al., 2006). Like all digeneans, schistosomes have an indirect lifecycle.
They require an intermediate freshwater snail host within which they reproduce asexually ultimately
producing cercariae, which are free-swimming larval stages that subsequently infect a definitive
mammalian host (Cribb et al., 2003). Definitive animal or human hosts can become infected with
schistosomiasis by entering infested waters—the water-borne larvae burrow through the skin of the new
host (Cribb et al., 2003). There are at least 12 known schistosome species in Africa of which 5 are known
to infect humans (S. haematobium, Schistosoma mansoni, S. intercalatum, S. guineensis and S. mattheei).
Schistosoma mattheei is of note as although predominantly a parasite of cattle, it has also been found in
wildlife and humans where it is known to hybridise with S. haematobium (see Pitchford, 1961). The other
species infect a wide range of domestic and wild animals including cattle, horses, buffalo, baboons,
zebra, hippopotamus and rodents (Standley et al., 2012). Traditionally, malacological monitoring
programmes have only targeted snail species known to harbour human infecting schistosomes, but a
wider approach is clearly needed as we become aware of wider host ranges (Pennance et al., 2021) that
are likely to shift with increasing environmental stressors. There is currently no vaccine for
schistosomiasis and the main control strategy for humans is preventative chemotherapy, improved
water, sanitation and hygiene and snail control (WHO, 2022).

3: Challenges of BTb control at the wildlife-livestock interface: The South

African case study
South Africa has been challenged with the control of BTb since the disease was first reported in the
country in 1880, initially focusing on livestock, and now including control in wildlife (DAFF, 2016).

3.1: Control in livestock

Early BTb surveillance included the introduction of tuberculin skin testing in cattle in 1905, followed by
its declaration as a notifiable disease in 1911 and the initiation of the Division of Veterinary Services BTb
scheme in 1969 (DAFF, 2016; Michel et al., 2019). This scheme focused on compulsory testing of
commercial cattle herds suspected to be infected, with slaughter of positive individuals, quarantine and
disinfection of farms. Initially, great progress was made, reducing prevalence to 0.04% by 1991 (1.1
million cattle tested); however, the number of tests have since declined due to budget cuts and a
decreased workforce (DAFF, 2016; Michel et al., 2019). Current prevalence in communal livestock is
variable (< 0.5% to > 15%) (Musoke et al., 2015; Sichewo et al., 2019b).
In 2021 the national cattle herd was estimated at 12 million, consisting of commercial dairy herds
(20%) and beef and dual-purpose herds (80%) (DAFF, 2021). Testing of cattle is no longer compulsory
and current control of BTb is guided by the Interim BTb Manual from the Department of Agriculture,
Forestry and Fisheries (DAFF), South Africa, which proposes the use of four testing programmes (Table
2; DAFF, 2016). All programmes are voluntary apart from the infected herd program, which can be
enforced by the Animal Diseases Act, 1984 (Act No. 35 of 1984) (DAFF, 2016). The approved test is the
cervical intradermal tuberculin (CIT) test (DAFF, 2016).

Table 2

Four levels of Bovine Tb surveillance programmes in South Africa.

Surveillance herd One off survey used by state officials to determine the prevalence of BTb within an
programme area or by a stock owner conducting a self-assessment
Maintenance To join this programme, herds are required to undergo two consecutive tests with
herd 100% negative results at least 3 months apart. These BTb free herds are then
programme tested every 2 years. If an individual tests positive, then the entire herd is
moved to the infected herd programme
Infected herd Compulsory programme for herds that have tested positive with the CIT test, as
programme well as those detected from meat and milk inspection, post-mortems or clinical
cases. These herds are placed under quarantine and kept under supervision of
a state veterinarian, who will order the slaughter of infected animals. The rest
of the herd is tested every 3 months and is only let out of quarantine once the
herd has undergone two consecutive negative tests
Diagnostic Individual cattle destined to be imported or exported. Imported cattle are kept in
testing quarantine and must undergo a compulsory CIT test. Before export, cattle
programme must also receive a comparative CIT test—a requirement for many importing
(individuals) countries
3.2: Control in wildlife
The control of BTb in wildlife is becoming increasingly important as many farms switch from livestock
to game farming, and wild buffalo reservoirs hinder control efforts in cattle (Michel et al., 2019). Bovine
Tb has been identified in 21 different wildlife species in South Africa, including most recently giraffe
(Hlokwe et al., 2019). The current control scheme is focused on domestic cattle and although some tests
have been adjusted for use in buffalo, this is not the case for other wildlife species. The Buffalo
Veterinary Procedural Notice (VPN) was published in 2017 outlining the procedures for disease testing,
movement and contingency planning for disease outbreaks in buffalo (DAFF, 2017). The buffalo VPN
states that for movement purposes, buffalo must have a negative CIT test as outlined in the manual for
cattle. Importantly, the interpretation of CIT has been based on cattle thresholds due to the lack of
species-specific cut-off values for African buffaloes. The gamma interferon test is also an effective
diagnostic tool for buffalo but is not approved by DAFF for movement purposes. There is currently no
guidance on control of BTb in other wildlife species and there are limited verified diagnostic tests in
these species (DAFF, 2017).
Kruger National Park and Hluhluwe-iMfolozi Park are the only two parks within South Africa that
contain buffalo herds maintaining BTb yet they have adopted different control approaches. Bovine Tb
was first detected in Hluhluwe-iMfolozi Park in 1986 and a test and cull disease programme was
initiated in 1999. This programme involved a mobile capture unit to corral buffalo in different areas of
the park, test them by means of the CIT test and culling positive individuals. Between 1991 and 2006,
4733 buffalo were tested, with herd prevalence ranging from 2.3% to 54.7%. Subsequent, data analysis
suggested that the programme was effective at reducing BTb prevalence, particularly in areas with
intensive test and culling operations (Le Roex et al., 2016). Kruger National Park took a different
approach to managing BTb in its buffalo population after the disease was detected in this host species in
1990. They aimed to breed disease free buffalo from Foot and Mouth Disease infected parents within the
park in order to conserve the genetic pool of Kruger buffalo in an ex-situ population (Laubscher and
Hoffman, 2012). This approach, which used dairy cows as foster parents for buffalo calves initially, and
later switched to having the buffalo mothers rear their young, was highly successful and also popular
with farmers, eventually shifting from a few government funded projects to hundreds of private buffalo
breeding farms (Laubscher and Hoffman, 2012). Additionally, Kruger National Park did extensive BTb
monitoring surveys between 1993 and 2007, to assess the spread and impact of BTb in herds, and
determine if the disease was having population level effects. Since it entered the park, BTb has been
detected in 12 spill-over species (Michel et al., 2006) and remains a concern in low density species, such
as wild dog and black rhinoceros (Higgitt et al., 2019).
With the disease currently not shown to be affecting population recruitment or growth in buffalo, the
real concern becomes spill-over to other hosts and therefore finding an effective vaccine that limits
disease severity and spill-over is a priority. Currently there is only one registered vaccine for BTb
control. The BCG vaccine is predominantly used in humans but has yielded promising results for use in
domestic cattle (Arnot and Michel, 2020). However, when trialled in wild buffalo within the Kruger
National Park, the BCG vaccine protection was insufficient and did not limit bacterial shedding (De
Klerk et al., 2010). This was thought to have resulted from priming with environmental non-TB
mycobacteria, which has been shown to reduce the protective efficacy of the BCG vaccine (Brandt et al.,
2002; De Klerk et al., 2010). Importantly similar studies in badgers in the UK found the BCG vaccine to
be effective in limiting disease severity (and therefore bacterial load; Chambers et al., 2011), meaning
that defining the clinical end point for vaccine efficacy trials is important. Another vaccination trial in
buffalo is currently underway, testing both BCG and DNA-sub-unit vaccines.

4: Drivers of disease: The Kruger National Park case study

4.1: Past and present disease management
Kruger National Park first opened as the Sabi Game Reserve in 1898 (10,364 km2) as a response to
campaigns for the conservation of wild animals subjected to uncontrolled hunting and to the 1896
rinderpest epidemic (Mabunda et al., 2003). In 1926, the Sabi Game Reserve was combined with the
Singwitsi Reserve (5000 km2 region named after the Shingwedzi River) and later renamed Kruger
National Park. James Stevenson-Hamilton, who was appointed warden in 1902, was tasked with
managing the aftermath of the rinderpest epidemic which, along with previous hunting activities,
decimated the game population, leaving elephant and white rhinoceros (Ceratotherium simum) locally
extinct (Mabunda et al., 2003). The rinderpest epidemic also severely affected buffalo, eland (Tragelaphus
oryx) and greater kudu (Tragelaphus strepsiceros; hereafter referred to as kudu), whereas wildebeest and
zebra were unaffected (Stevenson-Hamilton, 1957). The first 60 years of park management (1900–1960)
focused on protecting, preserving, and propagating, aiming to increase game numbers through
introductions of large herbivores, provision of water sources and culling of predators (Venter et al.,
2008).
Colonel J.A.B Sanderberg took over from Stevenson-Hamilton as Warden in 1946 and 8 years later the
first case of anthrax was confirmed in the north of the park (Mabunda et al., 2003). This was followed by
repeated outbreaks in 1959–60, 1970 and 1990–91, and outbreaks in the central part of the park in 1993
and 1999 (Bengis et al., 2003; De Vos and Bryden, 1996). The 1959–60 outbreak lasted just 4 months and
yet within this time over 1000 mammals died: kudu, waterbuck (Kobus ellipsiprymnus) and roan
(Hippotragus equinus) being the most affected (Pienaar, 1961). Simultaneously, BTb likely entered the
park, transmitted from cattle to buffalo on the southern border, although it was not detected in the park
until 30 years later (Bengis et al., 1996). At this time, park management shifted to a ‘management by
intervention’ approach and the next 30 years (1960–1990) focused on measuring, monitoring and
manipulation (Mabunda et al., 2003). Fencing of the park was ordered by the National Department of
Agriculture in order to prevent the spread of disease to surrounding livestock, such as foot and mouth
(FMD) endemic in buffalo (Bengis et al., 2003). Fence construction started in the early 1960s with the
western boundary followed by the eastern boundary in the late 1960s, by 1980 all boundaries of the park
were enclosed. The fences (over 360 km in length and 65 km in width) restricted movement of wildlife
leading to increased numbers of large herbivores, such as elephant and buffalo, which were
subsequently controlled by culling operations and in the early 1970s, a certified abattoir was built within
the park to optimise use of the culled meat (Mabunda et al., 2003). From 1990 to 2010 management
shifted again to focus on integration, innovation and internationalisation. The severe drought of 1992–93
followed by the February floods in 2000 as well as the catastrophic wildfire in September 2001, which
killed both people and animals within the park, were indicative of the need for management to become
more adaptive to the increasingly unpredictable environment (Mabunda et al., 2003). Since 1995, Kruger
has used a strategic adaptive management approach, which involves management decisions and actions
guided by research and monitoring while learning from unexpected events or outcomes. This approach
also aims to maximise heterogeneity of the park and led to its expansion across national boundaries
creating the Greater Limpopo transfrontier conservation area (GLTFCA) spanning the Limpopo
(Mozambique), Kruger (South Africa) and Gonarezhou (Zimbabwe) National Parks. A portion of fences
of approximately 45 km was removed between Limpopo and Kruger in 2002 (Caron et al., 2016; Venter
et al., 2008).

4.2: Kruger National Park's current adaptive management approach

In the past, most management issues in Kruger National Park were focused within the park boundaries;
however, since the recognition that threats and drivers to biodiversity conservation often occur outside
of the footprint of the National Parks, management issues are extending beyond the park boundaries
and becoming more socio-economic in nature (Venter et al., 2008). The creation of the Greater Limpopo
transfrontier conservation area shifted the park from being single use for wildlife to a multi-use park,
sharing its land with communities and their livestock. The park's current strategic adaptive
management aims to increase understanding of complex ecosystems and broader societal needs of local
communities. This process is guided by setting appropriate thresholds of potential concern (TPC), a set
of adaptive management goals and endpoints that define upper and lower levels of acceptable change,
enabling management to determine how much a system can be allowed to fluctuate before it becomes a
concern and requires management action. Although TPCs prove useful for simple metrics like invasive
plants and river flows, they have proven more challenging for complex systems such as disease where
drivers and responders are not always known (Gaylard and Ferreira, 2011; Venter et al., 2008).
 Kruger National Park's 2018–28 management plan includes a disease management programme as a
supporting objective to the higher-level objective of biodiversity conservation. This programme
acknowledges endemic wildlife diseases within the park as a key component of biodiversity yet
highlights the need to prevent and mitigate the spread of disease at the wildlife-livestock-human
interface and limit the introduction or impact of novel infectious diseases (Spies et al., 2018).

4.3: Environmental drivers of disease transmission

4.3.1: Spatial heterogeneity and the north/south divide
Topography, climate, geology and the associated soil and vegetation patterns can exert a bottom-up
control on ecosystems. The combination of these abiotic factors can influence fire patterns and animal
behaviours, as well as disease dynamics (Venter et al., 2003).
Kruger National Park lies within part of the north-eastern South African lowveld, which generally has
plains of low to moderate relief with some low mountains and hills. The geology of the park can be
crudely divided into granite plains on the west and basalt plains on the east, separated by a north-south
strip of sedimentary rock (Venter et al., 2003). Rainfall in the park increases along a north to south
gradient with annual mean rainfall of 350 mm in the northeast to 750 mm in the southwest. Geology and
rainfall have influenced the difference in soil and vegetation types between the north and the south of
the park. The south generally consists of deeper and more diverse soil types with predominantly open
canopy acacia tree bushveld and savannah with a well wooded area in the southeast. In contrast, the
north tends to have less diverse, thinner soils with a higher calcium content. Vegetation is dominated by
mopane trees with rare lowveld riverine forest occurring along the rivers in the northeast and sandveld
vegetation type in the northwest (Gertenbach, 1983; Spies et al., 2018). The northern most section of the
park is unique as it contains a varied assemblage of rock formations with associated soil and vegetation
types. It also contains the only true floodplain in Kruger (Venter et al., 2003). For management purposes,
Kruger National Park has been partitioned into 35 landscapes depending on geomorphology,
vegetation, soil, climate types and associated fauna (Gertenbach, 1983; Venter et al., 2003). A social-
economic gradient exists along the northern and southern boundaries of the park. Dense peri-urban to
urban developments lie along the southwestern border, including sugarcane plantations, forestry and
the nearby city of Mbombela (previously known as Nelspruit; Fig. 2). The central and north-western
boundaries are buffered by private nature reserves and community subsistence farming, and further
north becomes more rural with large agricultural areas and poor villages with limited economic
opportunities (Spies et al., 2018). Wildlife densities also differ across the park with megaherbivore
(elephant and buffalo) densities higher in the north than the south (Fig. 2).
 FIG. 2Megaherbivore (African buffalo and elephant) density across Kruger National
Park and fence breakages (red cross) from damage causing animals (DCAs).
Elephant cause most breakages enabling diseased buffalo to escape. Foot and
mouth (FMD) veterinary control zones and nearby villages are also shown. Map
produced by the Skukuza GIS Office.

This ecological heterogeneity within the park can create spatial heterogeneity in disease dynamics. A
park wide survey of RVF in buffalo in 1998 showed significantly higher seroprevalence of buffalo herds
in the south and central regions of the park compared to the north (Beechler et al., 2015a). This was
attributed to lower rainfall and different vegetation in the north leading to less suitable breeding
habitats for mosquito vectors (Beechler et al., 2015a). Brucellosis prevalence in buffalo was significantly
associated with park section and soil type (Gorsich et al., 2015). Buffalo captured on the resource poor
granitic soils were twice as likely to be seropositive for brucellosis compared to those on the resource
rich basaltic soils (Gorsich et al., 2015). Moreover, buffalo on granitic soils had higher prevalence in the
southern section of the park compared to the central section (Gorsich et al., 2015). This was attributed to
nutrient poor vegetation in the southwestern granitic soils and general lower body condition of buffalo
in the south of Kruger National Park (Caron et al., 2003; Gorsich et al., 2015). The effect of brucellosis
infection was also dependant on the seasonal heterogeneity of the park, brucellosis infection was
significantly associated with lower body condition but only in the dry season (Gorsich et al., 2015).
Knowledge of this heterogeneity of different disease dynamics and how the landscape and environment
affect this is of great importance and can help target monitoring and management of diseases within the
park.

4.3.2: Climate change and severe weather events

Africa is considered one of the most vulnerable areas to global climate change (Serdeczny et al., 2017).
Average temperature readings from the Skukuza weather station in Kruger National Park have shown a
2 °C increase from 1977 to 2018 with a maximum temperature increase of 0.5 °C per decade (Dube and
Nhamo, 2019). Kruger National Park has suffered numerous droughts in 1966–67, 1982–83, 1991–92,
1995–96 and most recently in 2015–17 (Staver et al., 2019). The most severe drought on record in 1991–92
had a mean total rainfall of 235.6 mm compared to the 534 mm long-term mean annual rainfall and the
number of days with rain within this period (24.2) was significantly less than the mean annual total
(48.3; Zambatis and Biggs, 1995). This was followed by a severe flood event in 1996. Certain disease
outbreaks within the Kruger National Park, such as anthrax and foot and mouth disease (FMD) have
occurred after a dry period (Pienaar, 1961).
Drought not only affects resource availability and body condition, but also the behaviour and
movement of animals, including buffalo, which in turn can alter disease transmission (Cross et al., 2004;
Staver et al., 2019). Combining buffalo behavioural association data with disease models predicted that
dry conditions facilitate increased spread of BTb within buffalo populations due to increased herd
switching (Cross et al., 2004). The 2015–17 drought forced buffalo to move north to areas where the
drought was less severe (Staver et al., 2019). Movements like this could lead to the transmission of
diseases into new areas of the park and is likely to have played a role in the spread of BTb northwards
through the park (Michel et al., 2006).
Drought particularly affects the dynamics of water borne diseases such as schistosomiasis and RVF
carried by vectors (snails and mosquitoes respectively) reliant on water or moisture with populations
that fluctuate depending on climate variations (Cribb et al., 2003; Romoser et al., 2011). Floods and
prolonged wet periods have been associated with outbreaks of RVF. South Africa has experienced three
major RVF epidemics (1950–51, 1973–75 and 2008–11). Epidemiological data from the 2008 to 2011
epidemic was modelled to quantify spatial and temporal environmental factors associated with disease
incidence (Métras et al., 2015). Initial years saw the incidence of RVF increase with increased vegetation
density and presence of wetlands. However, for 2010 and 2011, of which 2010 was the longest lasting
outbreak, the strongest risk factor was temperature. For 2010, the risk of RVF increased by a hazard ratio
of 15.7 in areas between 25 and 32 °C and by 44.35 in areas over 35 °C compared with those below 25 °C
(Métras et al., 2015). This is important for Kruger National Park as the average annual temperature in
the south of the park was 32.3 °C and is set to increase (Dube and Nhamo, 2019).
Modelling the RVF outbreaks in Kruger National Park from 2008 to 2011 suggested that soil
saturation index anomalies exceeding the long-term mean by 20%, followed by a sudden rainfall event
could be a reliable predictor for outbreaks. When tested on previous outbreaks, the model successfully
predicted 90% of outbreaks more than 1 month before they occurred (Williams et al., 2016). Other
factors such as vegetation density and increased temperature are also important risk factors (Métras et
al., 2015).

4.3.3: Water sources

Water is often a focal point for wildlife-livestock interaction, particularly rivers which run between
protected areas and communal farmlands (Kock et al., 2014). Such interaction can enable spread of
diseases between wildlife within the park and livestock on the borders (Miguel et al., 2013; Pienaar,
1961). Within Kruger National Park, five perennial rivers (Sabi-Sands, Crocodile, Olifants, Letaba and
Luhuvu) run through the park as well as several seasonal rivers, natural pans and wetlands (Mabunda
et al., 2003; Pienaar et al., 1997). The Sabi River runs parallel to the fenced south-western border between
Kruger National Park and the adjacent communal lands of Bushbuckridge. Landscape resistance maps
for cattle and buffalo resource utilisation were used to model dispersal of these animals within these
two areas. Contact risk between buffalo and cattle was significantly higher in the dry season and was
concentrated along the Sabi River at the weaker parts of the fence. Contact risk was more widespread
and closer to villages in the wet season, yet still highest along the river (Kaszta et al., 2018). Water
sources can also increase the permeability of nearby fences to wildlife movements. Interviews with
fence maintenance workers in Kruger National Park reported that fences damaged by flooding and
predation were higher in areas with rivers compared to those without. Furthermore, reports of kudu
crossing the fences were significantly higher in areas with rivers, although this was not observed for
other wildlife (elephant, buffalo, warthog (Phacochoerus africanus) and impala) in the park (Jori et al.,
2011).
Contact at water sources has been attributed to disease outbreaks, such as FMD and anthrax (Miguel
et al., 2013; Pienaar, 1961). The Limpopo River runs between the northern edge of Kruger National Park
and adjacent communal land of Pezvi in Zimbabwe, both within the GLTFCA. Satellite data from
collared individual cattle from Pezvi and buffalo from Kruger National Park showed that the two
species shared 16.9% of habitat with most contacts occurring less than 500 m from the riverbed. These
contacts increased during the dry season suggesting that contact is driven by resource availability
(Miguel et al., 2013). Incidence of FMD antibodies in cattle was higher in sites with high buffalo contact
suggesting spread of the infection from buffalo to cattle (Miguel et al., 2013). Water points within
protected areas are also a source for interaction between different wildlife species, particularly in dry
seasons when water sources are limited. The anthrax outbreaks which occurred in the northern section
of Kruger National Park between 5 June and 11 October 1960 were all associated with natural and
artificial water points where large numbers of animals aggregated around the remaining available water
sources during the dry season (Pienaar, 1961).
Water sources also provide habitats for parasite-harbouring vectors such as freshwater snails and
mosquitoes. Freshwater snails harbour a huge number of digenean parasites some of which can cause
diseases of veterinary and medical importance, such as schistosomiasis and fascioliasis. Although
freshwater snails within Kruger National Park have been surveyed several times in the past (De Kock
and Wolmarans, 1998; De Kock et al., 2002; Wolmarans and De Kock, 2006), the diversity and
distribution of digenean parasites hosted by these snails has not yet been studied. The original surveys
identified the intermediate host snails for both schistosomes (Bulinus africanus, B. globosus, Bradyidius
tropicus, B. forskali, Biompharia pfeifferi) and fasciolids (Lymnea columella, L. natalensis; see (De Kock and
Wolmarans, 1998; De Kock et al., 2002), therefore it is important to explore the hidden digenean
diversity within these snails. Within Kruger National Park, schistosome species have been detected in
several animals including baboons, zebra, warthog, giraffe, kudu, wildebeest, buffalo (S. mattheii; see
Beechler et al., 2017; Pitchford et al., 1974) and hippopotamus (S. hippopotami and S.
edwardiense; see Pitchford and Visser, 1981). Animals sampled near man made dams had higher S.
mattheei infection rates and egg outputs than those at natural water sources, suggesting perennial
exposure and transmission at these sites (Pitchford et al., 1974).

4.3.4: Reservoir hosts

As seen with the ‘Microscopic Five’, most pathogens can infect more than one host (Cleaveland et al.,
2001). This is true for 77% of known livestock pathogens and 60% of known human pathogens
(Cleaveland et al., 2001; Haydon et al., 2002). Some hosts can act as reservoir hosts, also known as
maintenance hosts (Ashford, 1997; Haydon et al., 2002; Swinton et al., 2002). Essentially, reservoir hosts
can maintain the pathogen in the absence of cases in other species, and with a high enough prevalence
that parasites can spill over into another host species. Identifying reservoir hosts is crucial to
appropriately manage a disease. The failure to identify the importance of domestic dogs as a reservoir
host for guinea worm in humans has led to the re-emergence of a disease on the brink of eradication
(Durrant et al., 2020; Galán-Puchades, 2017).
In Africa, buffalo are well-known reservoir hosts for diseases such as BTb, FMD and corridor disease
(CD; Michel and Bengis, 2012). Buffalo are a keystone species in the African savanna ecosystem and
have a high economic value due to their importance for wildlife ecotourism, live game trade and
hunting industries (Glanzmann et al., 2016). Buffalo's gregarious nature, tendency to form large herds,
roam long distances, cross park boundaries and undergo regular fission fusion events make them ideal
hosts to maintain and transmit diseases (Caron et al., 2016; Cross et al., 2005; Wielgus et al., 2021). The
population of buffalo in South Africa in 1998 was estimated at over 31,000, of which only 7.7% were
disease-free (Winterbach, 1998). The two largest populations of buffalo in South Africa are found in
Kruger National Park and Hluhluwe-iMfolozi Park both of which are infected with BTb and CD, with
the Kruger population additionally being infected with FMD (Winterbach, 1998).
Within Kruger National Park, buffalo appear to have spread BTb to numerous wildlife species
including warthogs, baboons (Papio ursinus) and lions (Panthera leo). Bovine Tb isolates from all three of
these species were genetically highly similar and, in some cases, identical to buffalo strains (Keet et al.,
2000; Michel et al., 2009). More recently, high sero-prevalence (83%) has been detected in endangered
wild dogs (Lycaon pictus), thought to be eating infected prey such as warthogs (Higgitt et al., 2019).
Although these dogs appeared to be healthy at the time of the study, little is known about how the
disease may progress in this species (Higgitt et al., 2019). Another accepted reservoir for BTb is kudu
(Michel and Mare, 2000; Renwick et al., 2007). Clinical manifestations of BTb in kudu include abscesses
in the cranial lymph nodes from which infectious discharge is secreted onto thorns and leaves while the
animal browses on vegetation (Palmer, 2013; Renwick et al., 2007).
Buffalo are also thought to play a part in the maintenance of RVF during the inter-epidemic periods.
Between 2005 and 2008, a total of 227 buffalo seronegative for RVF were monitored within the park.
During the 4 years, five of these buffalo became seropositive despite no outbreaks being detected in
other species, which suggests circulation of the virus within the buffalo population (Beechler et al.,
2015a).

4.3.5: Co-infections
Since over 80% of all known species are parasitic, co-occurrence of different parasites is the norm
(Vaumourin et al., 2015). Such co-infections can be synergistic, by which one parasite facilitates the
infection of other parasites, antagonistic where one parasite inhibits infection of other parasites, or can
have no effect on each other (Hoarau et al., 2020; Vaumourin et al., 2015). A pathogen can alter the host's
immune response making it more susceptible to others (Ezenwa et al., 2010). Co-infections of closely
related parasite species or strains can lead to hybridisation, potentially creating more virulent pathogens
as seen with certain schistosome species (Huyse et al., 2009). In South Africa the predominantly animal
schistosome species S. mattheei has become increasingly prevalent in humans, thought to be due to
hybridisation with the human species S. haematobium (see Pitchford, 1961). This was confirmed
experimentally, resulting in fertile first generation (F1) hybrids which were more infective and
developed more quickly than the parents (Pitchford, 1961; Taylor, 1970; Wright and Ross, 1980).
However, prior infection with S. haematobium or S. mansoni seems necessary for S. mattheei to become
established in humans (Pitchford, 1961).
Over the last decade a body of research has been conducted assessing the impact of co-infections on
disease dynamics within African parks and the findings are concerning (Beechler et al., 2015b, 2019;
Broughton et al., 2021; Budischak et al., 2012; Ezenwa et al., 2010; Sylvester et al., 2017). In the
Hluhluwe-iMfolozi Park, helminth infections have been shown to alter the immune response of wild
buffalo, which could make them more susceptible to BTb infection (Ezenwa et al., 2010). Nematode
infected individuals had a depressed Th1 immune response, which is important in controlling BTb
infection and other intracellular microparasite infections, whereas Th1 responses were enhanced in
hosts that were nematode resistant. Disease modelling predicted that without these nematodes, BTb
would not have established infection in the buffalo population (Ezenwa et al., 2010). In 2008, an
outbreak of RVF occurred in a buffalo breeding facility close to the southern section of Kruger National
Park. To determine the effect of existing BTb infection on the dynamics of RVF outbreak, BTb positive
and BTb negative individuals were monitored for RVF before and during an outbreak. Bovine Tb
positive individuals had a twofold greater risk of RVF infection than BTb negative individuals. Bovine
Tb infection also worsened the clinical effects of RVF with pregnant co-infected individuals six times
more likely to abort than those with just RVF infections (Beechler et al., 2015b). Scaled-up models of
these data also showed that the presence of BTb increases the risk of RVF infection for the entire herd,
not just those infected with BTb. These findings were mirrored in free-ranging buffalo within the park
(Beechler et al., 2015b). Bovine Tb can also alter the composition of parasites within a host. Buffalo
within Kruger National Park that acquired BTb infections showed significant increases in both
taxonomic and functional parasite richness, as well as shifts in composition associated with the loss of
nematodes and gain of schistosomes (Beechler et al., 2019). Co-infections may also reduce the efficacy of
diagnostic tests. British calves experimentally infected with F. hepatica and M. bovis reacted less strongly
to the single intradermal comparative cervical tuberculin test (SICCT) than those infected with M. bovis
alone (Claridge et al., 2012).
Another important co-infection that could threaten the conservation of another ‘Big Five’ species is
BTb and Feline Immunodeficiency Virus (FIV) in lions (Sylvester et al., 2017). Feline Immunodeficiency
Virus is endemic to these keystone predators, yet BTb was not reported in Kruger National Park's lions
until 1996 (Keet et al., 2010). As FIV can cause lymphocyte deficiencies, infected lions may be
predisposed to infection with BTb. Within Kruger National Park, lions positive for FIV were more likely
(although this was not significant with a sample size of 56) to be infected with M. bovis than those
negative for FIV (Sylvester et al., 2017). A more recent study concluded that total gastrointestinal
parasite burden and richness was significantly higher in FIV positive lions (Broughton et al., 2021). Co-
infections of the ‘Microscopic Five’ and other diseases in Kruger's wildlife need greater attention as this
could greatly alter the dynamics of diseases previously thought to be benign. Co-infections leading to
hybridisations of human and animal specific schistosome species could also hinder control efforts as the
WHO currently focuses on treating human infection by mass drug administration to school age children
and little is known about whether hybrids could be more resistant to preventative chemotherapy or
whether it could change the age profile of infection. This could delay the WHO's target to eliminate
schistosomiasis as a public health problem by 2030 (Stothard et al., 2020; WHO, 2022).

4.4: Anthropogenic drivers of disease transmission: Wildlife-livestock-

human interface
Interactions between wildlife and livestock can drive the spread of diseases (Kock et al., 2014). Such
interaction can be linear, across a fence, or focal, at a shared water hole, where pathogens from an
infected animal or population can spill over into a vulnerable population via direct or indirect (vector)
contact. This spill-over can be bi-directional from livestock to wildlife or vice versa (Bengis et al., 2002).
Humans can also be involved with pathogens spilling over from animals to humans known as sylvatic
(from wildlife) or urban (domestic animals) zoonoses (Figueiredo, 2019). Bovine Tb likely first entered
Kruger National Park via transmission from cattle to buffalo in the south-western corner of the park
(Bengis et al., 1996). An outbreak of BTb was then recorded in cattle in the communal rangeland in
Mpumalanga Province on the western border of the park in 2012, suggesting spill back of the disease
from buffalo to cattle (Musoke et al., 2015).

4.4.1: Permeability of wildlife fences

Wildlife fences are commonly used in protected areas to prevent the spread of disease, such as foot and
mouth (FMD), between wildlife and livestock. A veterinary fence of note is the ‘Red Line’ in Namibia,
erected in 1960 with the purpose of separating the FMD endemic north from the rest of the country. This
1250 km fence extends from east to west bisecting the entire country (Miescher, 2012) with unintended
impacts on animal migration (Gadd, 2012) and socio-economic factors in the country. South Africa has
used a combination of fencing and zoning to improve FMD control, splitting the country into an
infected zone, buffer zone and FMD free zone (Fig. 2). This involved erecting a 750 km fence separating
the western and southern boundary of Kruger from neighbouring communal land, private farms and
private game reserves (Jori et al., 2011). The fences between the park and private reserves have since
been removed to allow more space for wildlife and the combined area makes up the infected zone.
Adjacent to this is a 10–20 km wide buffer zone which is split into two sections, one with vaccination
and one without vaccination but increased surveillance (Jori et al., 2009; Kaszta et al., 2018).
Although Kruger National Park's fences were placed as a barrier between infected and buffer zones,
they are highly permeable and, between 1996 and 2006, 1676 buffalo escaped across the park fence
bordering the Mpumalanga Province (Jori et al., 2009). Semi-structured interviews of fence workers
along 357 km of the western and southern border fence reported that higher numbers of kudu and
impala were seen crossing over into communal land than buffalo (Jori et al., 2011). Impala are also
capable of transmitting the FMD virus, they can experience both clinical and sub-clinical infections and
may maintain the South African Territory (SAT) serotype within local populations (Vosloo et al., 2009).
The same workers also reported that the main causes of fence damage were elephant and humans,
followed by predator and prey conflict, flooding and animals digging under the fence (Jori et al., 2011).
Elephant will break fences to access water, desirable trees, notably the Marula (Sclerocarya birrea) or
agricultural crops. Such fence breakages occur more in the northwest of the park, likely due to greater
elephant densities (Fig. 2). There are also more rivers traversing from west to east, through the fences
creating weak spots. These areas are also where livestock and wildlife may share water resources,
particularly in the dry winter months which show peaks of fence breaking activity.
Between 2000 and 2007, five outbreaks of FMD occurred in cattle near the western boundary of the
park (Jori et al., 2009). The most recent outbreak of FMD in Kruger occurred in 2013/14 in cattle in the
Mpumalanga Province, adjacent to the park's western border within the inspection zone with
vaccination. Most of these outbreaks were attributed to contact between wildlife and livestock, mainly
buffalo and cattle (Blignaut et al., 2020). The reliance on fences to act as a barrier against diseases may
exacerbate the problem as this may encourage farmers to relax vaccinating their livestock, or state
officials to not conduct regular animal inspections for early detection of outbreaks.

4.4.2: Edge effects

Edge effects describe the impacts of interactions between habitat patches or fragments and the
surrounding matrix (Suzán et al., 2012). In the case of Kruger National Park, the wildlife reserve is the
habitat patch surrounded by a matrix of communal farms and villages. Such edges can facilitate the
emergence of infectious diseases into new areas.
C. parvum was first detected in Kruger in 2008 with low prevalence in elephant, buffalo and impala
(4.2%, 1.4% and 1.9% respectively; Samra et al., 2011). For all three host animals, prevalence was
significantly higher in areas close to the western park boundary than in the park centre (Samra et al.,
2011). This suggests transmission from livestock or humans inhabiting the bordering farms and villages
to wildlife within the park. A subsequent molecular study within the same area detected a 2.8%
prevalence of C. ubiquitum in impala and C. bovis in buffalo again near the western park boundary
(Samra et al., 2013). Calves from the bordering communal farmlands of Bushbuckridge were also tested,
revealing a prevalence of 4% for C. andersoni and 4% for C. bovis. Farmers from the area reported buffalo
and impala as the most seen wild species outside the park boundary and 6.2% of these farmers reported
bringing their cattle into the park to drink (Samra et al., 2013). This may suggest spill over from either
livestock and or humans to wildlife or vice versa. Cryptosporidium was later confirmed in children from
the same communal grazing area with a 5.6% prevalence, but predominantly infected with the
anthroponotic species C. hominus (see Samra et al., 2016).

4.4.3: Transfrontier conservation areas

Many protected areas across the globe are clustered along international borders that are usually fenced
preventing the natural migration of large mammals. A management decision was taken to remove a
number of these fences within southern Africa to create one large, protected area known as
Transfrontier Conservation Areas (TFCA) and Transfrontier Parks (TFP) with the aim of conserving
biodiversity and enabling the movement of large mammals, aiding socio-economic development, and
promoting a culture of peace (Hanks, 2003). There are at least 13 TFCAs and TFPs within southern
Africa, six of which include South Africa (Lunstrum, 2011). The Greater Limpopo Transfrontier
Conservation Area (GLTFCA), created in 2002, spans the Limpopo (Mozambique), Kruger (South Africa)
and Gonarezhou (Zimbabwe) National Parks and includes conservancies, wildlife ranches and
communal farmland, covering a total area of 85,000 km2 (Caron et al., 2016; Ferreira, 2004). These areas
have many conservation benefits particularly for animals with large home ranges, such as elephant,
buffalo, and wild dog, that need more space to disperse or hunt (Caron et al., 2016; Cook et al., 2015;
Davies-Mostert et al., 2012). However, there is also now increased potential for pathogen spread
between wildlife, livestock and humans within these areas.
In 2009, a strain of BTb related to buffalo in Kruger National Park was identified in buffalo in
Zimbabwe. Telemetry studies revealed that sub-adult female buffalo were moving long distances
between the national parks and even out of the GLTFCA (Caron et al., 2016). One 2.5-year-old collared
female walked 95 km over 6 days during which she crossed into Zimbabwe and Mozambique and
visited a buffalo herd in the Limpopo National Park (Mozambique) as well as a commercial cattle
ranching area (Caron et al., 2016).

4.4.4: Neighbouring game farms and private reserves

South Africa's economy is rapidly transitioning from livestock-based to wildlife-based agriculture and
ecotourism, with the wildlife industry becoming the fastest growing agricultural sector (Saayman et al.,
2018). Kruger National Park is bordered by several private game reserves and farms. Certain private
wildlife reserves on the western border of the park have removed their fences to allow free movement of
animals, creating the Greater Kruger National Park Complex (GKNPC; Hlokwe et al., 2019). These
reserves are under the jurisdiction of the state veterinary office Bushbuckridge East (Orpen) of the
Mpumulanga veterinary services (Hlokwe et al., 2019). Although this creates larger areas for the wildlife
to roam it also increases the likelihood of interspecific and intraspecific interaction, which can be a
driver for disease transmission, particularly problematic when the disease status of some animals is
unknown. In 2013 a novel M. bovis strain was identified in a blue wildebeest (Connochaetes taurinus) that
had been culled after escaping a private game reserve in the GKNPC (Hlokwe et al., 2014). One or more
translocations of untested blue wildebeest likely brought in this new strain (Hlokwe et al., 2014). In 2014
a different M. bovis strain, not previously detected in South Africa, was found in a female giraffe (Giraffa
camelopardalis) within the same nature reserve (Hlokwe et al., 2019). A kudu on a game farm just south
of Kruger National Park tested positive for an M. bovis strain dissimilar to strains previously isolated
within Kruger wildlife or cattle in that region (Bengis et al., 2001). As kudu can easily cross fences and
are a good candidate reservoir host, they could pose a threat of introducing new strains to the park. The
risk of transmission of M. bovis and other infectious diseases from wildlife is increased by the fact that
testing, or even disease risk assessment, before translocation is only mandatory for buffalo and no other
wildlife species or cattle (Hlokwe et al., 2014, 2019). Thus, the translocation of untested animals (both
wild and domestic) from these private farms and reserves could pose a threat to wildlife in Kruger
National Park and surrounding livestock and humans.

4.4.5: Human-wildlife conflicts and illegal wildlife trade

Protected areas may be thought of as a haven for wildlife with tourists and staff often the only human
inhabitants, yet Kruger National Park, for instance, is bordered by seven municipalities in which
approximately two million people reside (Swemmer et al., 2017). The Limpopo Province surrounding
the north of the park is one of the poorest in the nation with a 67% poverty rate (Warchol and Johnson,
2009). It is therefore important to understand how people and their activities can influence disease
dynamics within and around parks and this must be factored into management decisions. A major
threat to the biodiversity and conservation is the illegal wildlife trade, driven by inequality, poverty,
food insecurity and increasing human populations (Bezerra-Santos et al., 2021; Lindsey et al., 2013). The
illegal wildlife trade is also a significant risk factor for the spread of zoonotic pathogens (Bezerra-Santos
et al., 2021). The trade directly threatens one of Kruger National Park's ‘Big Five’, the white rhinoceros,
poached for their horns to be sold on the Asian market (Annecke and Masubelele, 2016).
Hunting of other game including buffalo and kudu for bush meat is also on the rise in Kruger. What
was once a local subsistence practice, the bush meat trade is now a commercialised, global enterprise
and is no longer sustainable (Warchol and Johnson, 2009). Interviews with communities neighbouring
Kruger National Park suggested that bush meat was poached from the park and readily marketed by
local merchants (Warchol and Johnson, 2009). Several social and cultural factors influenced this
including the affordability of bush meat and the use in traditional weddings, as well as the perception
that parks were not benefiting local communities. There were also reports of game rangers in the park
being complicit with poachers (Warchol and Johnson, 2009). Although there are no studies in Kruger
National Park assessing the role of poaching and bush meat in the spread of zoonotic diseases to
humans or their livestock, these activities have been linked to parasitic outbreaks in humans across the
globe (Bezerra-Santos et al., 2021). Risk assessments are needed to determine the threat of zoonotic
disease spread through the trade and consumption of bush meat from Kruger National Park. Poaching
can also exacerbate the problem of permeable fences as fences are cut or damaged by poachers entering
the park which can lead to animals escaping from the park.

5: Disease knowledge gaps and lessons learnt from African protected

areas
Research underpins Kruger National Park's Strategic Adaptive Management approach but going
forward we need to plug key knowledge gaps with both traditional and novel research techniques. Most
disease-related research within parks have focused on pathogens of economic importance with simple
(direct) life cycles such as FMD and BTb. Pathogens with complex life cycles, such as digeneans, have
been largely neglected. Increased knowledge of these parasites including their genetic diversity and
distribution is hugely important, especially with mounting evidence on the impact of co-infections
(Ezenwa et al., 2010).
There is a global lack of knowledge about the transmission of Cryptosporidium oocysts within the
environment (Innes et al., 2020). African protected areas would provide a good study system to better
understand this transmission and determine exactly how these parasites enter, and possibly accumulate
in parks. It is also crucial to determine whether the species detected negatively affect the fitness of
infected wildlife and whether infection with Cryptosporidium species could increase the pathogenicity of
existing diseases (Beechler et al., 2015a). There is an apparent lack of empirical data for the more
neglected ‘Microscopic Five’ (schistosomiasis and cryptosporidiosis) and for other similar diseases. The
recent pandemic has further hindered the control efforts of neglected diseases, but also highlighted the
importance of monitoring them (Ung et al., 2021). It is imperative that on the ground surveys and
collection of empirical data is carried out to understand disease epidemiology in parks. This data can be
easily and relatively cheaply collected through traditional epidemiological methods, such as serological
surveys and questionnaires, which can be used to analyse the risk of these diseases to people, their
livestock and wildlife within the park.
For some of the more high-profile diseases such as BTb and brucellosis, much empirical data has
already been published and can be assimilated together in mathematical models, which can aid the
adaptive management process. Where it is difficult to acquire sufficient empirical data needed for
traditional modelling of disease dynamics, a Bayesian approach has been suggested as it allows for
diseases to be modelled in complex systems with limited data (Kosmala et al., 2016). Using this
approach to assess the dynamics of BTb in lions in Kruger National Park has revealed that the pathogen
is primarily transmitted from buffalo to lion, while lion to lion transmission is low (Kosmala et al., 2016).
Longitudinal studies are necessary to determine trends over time. One resource that can help are
biobanks, where samples and corresponding data from a variety of wildlife species have been frozen for
decades, over 20 years in the case of Kruger National Park. Such resources should be used to aid our
understanding of long-term trends in disease, which can then be incorporated into mathematical
models for forecasting. Other protected areas should strive to create their own biobanks from collected
samples and data as this can be a valuable resource for disease management as well as species
conservation.
The increasing switch from livestock farming to game farming poses some important questions
regarding the transmission of zoonotic infectious diseases within and around protected areas. Many
national disease control measures, including those for BTb and brucellosis in South Africa, are focused
on vaccination and monitoring of livestock while wildlife is largely neglected. In South Africa disease
testing for BTb is only mandatory for livestock/buffalo but not for other wildlife (DAFF, 2016; Michel et
al., 2019). The translocation of wildlife between game farms and reserves around Kruger National Park
has already brought new strains of BTb, which could spill over into the park (Hlokwe et al., 2014).
Moreover, this could have health implications as the industry for game meat expands globally,
prompting the need to adopt hazard identification and critical control points protocols for safe food
production.
Landscape characteristics influence dynamics of several diseases within African protected areas (Dion
et al., 2011). Kruger National Park's heterogeneous landscape has led to marked differences in disease
dynamics between the northern and southern regions of the park. A handful of studies have assessed
spatial epidemiology within Kruger National Park (Dion and Lambin, 2012; Dion et al., 2011), but this
cannot be achieved without disease prevalence and host-parasite abundance data (Schwabl et al., 2017).
Landscape genetics/genomics has been strongly advocated as a method to overcome these issues
(Schwabl et al., 2017). The method combines spatial and molecular data to assess correlations between
gene flow or local adaptation and landscape features (Schwabl et al., 2017; Storfer et al., 2018). Previous
applications of landscape genetics/genomics have focused on assessing barriers to gene flow within
vulnerable populations to aid conservation efforts (Corlatti et al., 2009; Wasserman et al., 2013). More
recently it has been applied to track parasite transmission (Biek and Real, 2010). However, its use for
parasites with complex lifecycles and multiple hosts is currently limited (Sprehn et al., 2015). Landscape
genomics would be particularly useful in predicting the spread of diseases maintained by important
reservoir species such as buffalo. Tracking the movement or restriction of buffalo gene flow across the
landscape may help to predict the movement of diseases such as BTb and FMD. This approach will be
more complex for pathogens that require an intermediate host as part of their life cycle, such as
schistosomes. In this case landscape genomics could be applied to determine the effect of intermediate
and definitive hosts on gene flow and local adaptation of parasites (Sprehn et al., 2015).
Preventing emerging infectious diseases from entering the park is part of Kruger National Park's 10-
year management plan. Edges of the park, especially those adjacent to human settlements and
farmlands, are important areas for introduction of diseases as seen for M. bovis and Cryptosporidium spp.
(see Samra et al., 2011). Routine monitoring and surveillance of potential diseases is key to the early
detection and prevention of disease outbreaks (Karimuribo et al., 2012; Webster et al., 2016). Surveillance
and monitoring can be simple and should ensure optimal use of the resources available (Karimuribo et
al., 2012). Some innovative examples have made use of available technology such as mobile phone apps
to collect epidemiological data (Aanensen et al., 2009), this opens the possibility of a more participatory
and citizen science approach where locals and tourists could become a part of disease surveillance
programmes. A successful citizen science project was carried out in the Serengeti National Park in
Tanzania, where over 1 million images from a large-scale camera trapping survey were classified by
over 28,000 volunteers (Swanson et al., 2015). Each image was circulated to multiple volunteers and
responses were aggregated, using an algorithm to produce a quality data set with 97.9% agreement with
a subsample analysed by experts (Swanson et al., 2015, 2016). Similar methods could be useful for
notifying mangers of sick or dead animals which could then be monitored for disease. South African
National Parks (SANParks) are developing a surveillance system and training conservation staff to
identify basic disease syndromes and a system for reporting sick or dead animals (Spies et al., 2018).
Involving local people in conservation and disease management is another important strategy as
many drivers of disease dynamics around Kruger National Park are anthropogenic in nature.
Community Based Natural Resource Management has become popular in Africa and in some protected
areas, yet it is important that both parks and communities benefit equally (see Section 6). Community
participation played a key part in the global eradication of rinderpest in 2011 (reviewed by Roeder et al.,
2013), whereby locals were trained in vaccination and sero-monitoring of cattle which enabled 80% herd
immunity in rural areas of Africa. This was successful as local people benefitted from the vaccination of
their livestock and training empowered local communities (Roeder et al., 2013). Social studies are
important to understand how communities perceive protected areas and why human-wildlife conflicts
may occur.
Integrating the needs of the environment, animals and humans underpins the fast growing ‘One
Health’ concept, which is advocated for the control of zoonotic diseases (Webster et al., 2016). There are
several governmental and non-governmental organisations in sub-Saharan Africa that are using the
‘One Health’ approach to monitor and control zoonotic diseases at the human-livestock-wildlife
interface (reviewed by Rwego et al., 2016). As many protected areas have borders which create such an
interface, it is important that park managers work with existing institutions and agencies working in
this field and share data and management strategies. Vaccine research and development is advocated in
many One Health frameworks and there is potential for their development or improvement in each of
the Microscopic Five, however parasitic diseases with complex life cycles and immune invasive
behaviour as seen in schistosomes pose a greater challenge (Driciru et al., 2021).
Although different authorities are responsible for wildlife, livestock and human health, the
intersectoral impact of zoonotic diseases means these authorities all need to take equal responsibility for
monitoring and preventing outbreaks and spread of such diseases. Lack of communication and
 g p g p
cooperation between these sectors can hinder control efforts and the ‘One Health’ approach advocates
working across disciplines to achieve effective control of zoonoses (Randolph, 2020; Webster et al., 2016).
Nevertheless, new funding is essential to coordinate this joint effort. Zoonotic diseases and their
ecological drivers should be brought into the curriculum for student doctors along with knowledge
sharing and joint training between doctors and veterinarians. The Wits rural facility, a campus of the
University of Witwatersrand located near the southwest border of Kruger National Park, aims to
achieve this through a practical training campus where student veterinarians and medics receive
multidisciplinary training right at the human-livestock-wildlife interface.
This transdisciplinary ‘One Health’ approach is being spearheaded by three intergovernmental
organisations the World Health Organisation (WHO), the Food and Agricultural Organisation (FAO)
and the World Organisation for Animal Health (OIE), which formed a Tripartite alliance in 2010
(recently joined by the UN Environment Program (UNEP) forming a Quadripartite; UNEP, 2022; WHO,
2017). While these organisations are important for coordinating global responses to pandemics and
promoting communication and collaboration between nations, participation is voluntary, and they
cannot legally enforce policy. It is therefore the responsibility of local and national government
authorities to create new policies and enforce them. Ministries of health, agriculture, wildlife,
environment, trade and tourism must collaborate and work towards a joint commitment to long term
monitoring and prevention of zoonotic disease outbreaks (Randolph, 2020). Some African countries
have already set up collaborations between different ministries (Rwego et al., 2016).
Initial disease monitoring and prevention should start at the local level but for this to be successful,
coordination is needed at the regional, national and global level. Local communities, farmers and
consumers should also practice behaviours to prevent the spread of diseases, however community
engagement and training is needed to facilitate this. Improvements in policy and regulation in testing
and movement of animals, farming practices and the wildlife trade are also needed and must come from
local and national governments (Randolph, 2020). It is important to bridge the gap between the
management of diseases within protected areas and the surrounding communities and farms. Public
health officials should work with park managers to assess potential threats to human health as well as
threats to wildlife. This has been demonstrated in Uganda Bwindi Impenetrable National Park (BINP)
where the Ugandan wildlife authority, Bwindi community hospital and Kayonza Government Health
Centre, medical and veterinary officers are working together with an NGO called Conservation Through
Public Health (CTPH) to protect gorillas in the park and surrounding communities from disease (Rwego
et al., 2016).
A major barrier to this global co-operation is the lack of political will and lack of funding particularly
in African countries. Public-private partnerships may help alleviate these issues for example the NGO
African Parks has secured long term contracts with governments of 11 countries to manage 19 parks in
central and southern Africa and has mobilised a large funding base (African parks, 2022). Conflicts
between the local population and park authorities in addition to human wildlife conflict can also hinder
the above actions and efforts must be made to rebuild relationships and perceptions which may have
been damaged throughout history. The aftermath of the recent Covid-19 pandemic also poses
challenges and has led to large declines in funding for national parks which rely heavily on tourism for
income (Lindsey et al., 2020). Yet this creates a good opportunity to reflect and re-assess. Parks should
consider diversifying income streams to not be so reliant on international tourism for income, given its
vulnerabilities to shocks such as pandemics and economic recession. The pandemic has proven that
humans are not just threats to biodiversity, but important custodians of nature, and finding diverse
ways to fund their continued involvement as scientists, conservation managers and rangers is important
to ensure a healthy planet for people and nature (Bates et al., 2021).

6: Communities and conservation

Many areas of southern Africa are still inhabited by indigenous tribes, initially hunter gatherers, who's
knowledge of wildlife and land management has been passed down over centuries (Suzman, 2001). In
the early 1900s many of these indigenous people were cleared from their homelands to make way for
national parks and those who remained within parks had little control over the area's resources, creating
conflicts between parks and local communities (Suzman, 2000). Although many parks are now shifting
Another random document with
no related content on Scribd:
chair. But if there be objection, or another proposed, a question is
put by the Clerk. 2 Hats., 158. As are also questions of adjournment.
6 Grey, 406. Where the House debated and exchanged messages and
answers with the King for a week without a Speaker, till they were
prorogued. They have done it de die in diem for fourteen days. 1
Chand., 331, 335.
[In the Senate, a President pro tempore, in the absence of the
Vice-President, is proposed and chosen by ballot. His office is
understood to be determined on the Vice-President’s appearing and
taking the chair, or at the meeting of the Senate after the first recess.]
Where the Speaker has been ill, other Speakers pro tempore have
been appointed. Instances of this are 1 H., 4. Sir John Cheyney, and
Sir William Sturton, and in 15 H., 6. Sir John Tyrrel, in 1656,
January 27; 1658, March 9; 1659, January 13.
Sir Job Charlton ill, Seymour chosen, 1673,
February 18.
Not merely pro tempore. 1 Chand.,
Seymour being ill, Sir Robert Sawyer
169, 276, 277.
chosen, 1678, April 15.
Sawyer being ill, Seymour chosen.
Thorpe in execution, a new Speaker chosen, 31 H. VI, 3 Grey, 11;
and March 14, 1694, Sir John Trevor chosen. There have been no
later instances. 2 Hats., 161; 4 Inst. 8; L. Parl., 263.
A Speaker may be removed at the will of the House, and a Speaker
pro tempore appointed. 2 Grey, 186; 5 Grey, 134.

SEC. X.—ADDRESS.

[The President shall, from time to time, give to the Congress

information of the state of the Union, and recommend to their
consideration such measures as he shall judge necessary and
expedient. Const., II, 3.]
A joint address of both Houses of Parliament is read by the
Speaker of the House of Lords. It may be attended by both Houses in
a body, or by a committee from each House, or by the two Speakers
only. An address of the House of Commons only may be presented by
the whole House, or by the Speaker, 9 Grey, 473; 1 Chandler, 298,
301; or by such particular members as are of the privy council. 2
Hats., 278.

SEC. XI.—COMMITTEES.

Standing committees, as of Privileges and Elections, &c., are

usually appointed at the first meeting, to continue through the
session. The person first named is generally permitted to act as
chairman. But this is a matter of courtesy; every committee having a
right to elect their own chairman, who presides over them, puts
questions, and reports their proceedings to the House. 4 Inst., 11, 12;
Scob., 9; 1 Grey, 122.
At these committees the members are to speak standing, and not
sitting; though there is reason to conjecture it was formerly
otherwise. D’Ewes, 630, col. 1; 4 Parl., Hist., 440; 2 Hats., 77.
Their proceedings are not to be published, as they are of no force
till confirmed by the House, Rushw., part 3, vol. 2, 74; 3 Grey, 401;
Scob., 39. Nor can they receive a petition but through the House. 9
Grey, 412.
When a committee is charged with an inquiry, if a member prove
to be involved, they cannot proceed against him, but must make a
special report to the House; whereupon the member is heard in his
place, or at the bar, or a special authority is given to the committee to
inquire concerning him. 9 Grey, 523.
So soon as the House sits, and a committee is notified of it, the
chairman is in duty bound to rise instantly, and the members to
attend the service of the House. 2 Nals., 319.
It appears that on joint committees of the Lords and Commons,
each committee acted integrally in the following instances: 7 Grey,
261, 278, 285, 338; 1 Chandler, 357, 462. In the following instances
it does not appear whether they did or not; 6 Grey, 129; 7 Grey, 213,
229, 321.

SEC. XII.—COMMITTEE OF THE WHOLE.

 The speech, messages, and other matters of great concernment,
are usually referred to a committee of the Whole House, (6 Grey,
311,) where general principles are digested in the form of resolutions,
which are debated and amended till they get into a shape which
meets the approbation of a majority. These being reported and
confirmed by the House, are then referred to one or more select
committees, according as the subject divides itself into one or more
bills. Scob., 36, 44. Propositions for any charge on the people are
especially to be first made in a Committee of the Whole. 3 Hats., 127.
The sense of the whole is better taken in committee, because in all
committees every one speaks as often as he pleases. Scob., 49. They
generally acquiesce in the chairman named by the Speaker; but, as
well as all other committees, have a right to elect one, some member,
by consent, putting the question. Scob., 36; 3 Grey, 301. The form of
going from the House into committee, is for the Speaker, on motion,
to put the question that the House do now resolve itself into a
Committee of the Whole to take into consideration such a matter,
naming it. If determined in the affirmative, he leaves the chair and
takes a seat elsewhere, as any other member; and the person
appointed chairman seats himself at the Clerk’s table. Scob., 36.
Their quorum is the same as that of the House; and if a defect
happens, the chairman, on a motion and question, rises, the Speaker
resumes the chair, and the chairman can make no other report than
to inform the House of the cause of their dissolution. If a message is
announced during a committee, the Speaker takes the chair and
receives it, because the committee cannot. 2 Hats., 125, 126.
In a Committee of the Whole, the tellers on a division differing as
to numbers, great heats and confusion arose, and danger of a
decision by the sword. The Speaker took the chair, the mace was
forcibly laid on the table; whereupon, the members retiring to their
places, the Speaker told the House “he had taken the chair without
an order, to bring the House into order.” Some excepted against it;
but it was generally approved, as the only expedient to suppress the
disorder. And every member was required, standing up in his place,
to engage that he would proceed no further in consequence of what
had happened in the grand committee, which was done. 3 Grey, 128.
A Committee of the Whole being broken up in disorder, and the
chair resumed by the Speaker without an order, the House was
adjourned. The next day the committee was considered as thereby
dissolved, and the subject again before the House; and it was decided
in the House, without returning into committee. 3 Grey, 130.
No previous question can be put in a committee; nor can this
committee adjourn as others may; but if their business is unfinished,
they rise, on a question, the House is resumed, and the chairman
reports that the Committee of the Whole have, according to order,
had under their consideration such a matter, and have made
progress therein; but not having had time to go through the same,
have directed him to ask leave to sit again. Whereupon a question is
put on their having leave, and on the time the House will again
resolve itself into a committee. Scob., 38. But if they have gone
through the matter referred to them, a member moves that the
committee may rise, and the chairman report their proceedings to
the House; which being resolved, the chairman rises, the Speaker
resumes the chair, the chairman informs him that the committee
have gone through the business referred to them, and that he is
ready to make report when the House shall think proper to receive it.
If the House have time to receive it, there is usually a cry of “now,
now,” whereupon he makes the report; but if it be late, the cry is “to-
morrow, to-morrow,” or “Monday,” &c., or a motion is made to that
effect, and a question put that it be received to-morrow, &c. Scob.,
38.
In other things the rules of proceeding are to be the same as in the
House. Scob., 39.

SEC. XIII.—EXAMINATION OF WITNESSES.

Common fame is a good ground for the House to proceed by

inquiry, and even to accusation. Resolution House of Commons, 1
Car. 1, 1625; Rush, L. Parl, 115; 1 Grey, 16–22, 92; 8 Grey, 21, 23, 27,
45.
Witnesses are not to be produced but where the House has
previously instituted an inquiry, 2 Hats., 102, nor then are orders for
their attendance given blank. 3 Grey, 51.
When any person is examined before a committee, or at the bar of
the House, any member wishing to ask the person a question, must
address it to the Speaker or chairman, who repeats the question to
the person, or says to him, “You hear the question—answer it.” But if
the propriety of the question be objected to, the Speaker directs the
witness, counsel, and parties to withdraw; for no question can be
moved or put or debated while they are there. 2 Hats., 108.
Sometimes the questions are previously settled in writing before the
witness enters. Ib., 106, 107; 8 Grey, 64. The questions asked must
be entered in the journals. 3 Grey, 81. But the testimony given in
answer before the House is never written down; but before a
committee, it must be, for the information of the House, who are not
present to hear it. 7 Grey, 52, 334.
If either House have occasion for the presence of a person in
custody of the other, they ask the other their leave that he may be
brought up to them in custody. 3 Hats., 52.
A member, in his place, gives information to the House of what he
knows of any matter under hearing at the bar. Jour. H. of C., Jan. 22,
1744–5.
Either House may request, but not command, the attendance of a
member of the other. They are to make the request by message of the
other House, and to express clearly the purpose of attendance, that
no improper subject of examination may be tendered to him. The
House then gives leave to the member to attend, if he choose it;
waiting first to know from the member himself whether he chooses
to attend, till which they do not take the message into consideration.
But when the peers are sitting as a court of criminal judicature, they
may order attendance, unless where it be a case of impeachment by
the Commons. There, it is to be a request. 3 Hats., 17; 9 Grey, 306,
406; 10 Grey, 133.
Counsel are to be heard only on private, not on public bills, and on
such points of law only as the House shall direct. 10 Grey, 61.

SEC. XIV.—ARRANGEMENT OF BUSINESS.

The Speaker is not precisely bound to any rules as to what bills or

other matter shall be first taken up; but it is left to his own
discretion, unless the House on a question decide to take up a
particular subject. Hakew., 136.
 A settled order of business is, however, necessary for the
government of the presiding person, and to restrain individual
members from calling up favorite measures, or matters under their
special patronage, out of their just turn. It is useful also for directing
the discretion of the House, when they are moved to take up a
particular matter, to the prejudice of others, having priority of right
to their attention in the general order of business.
[In the Senate, the bills and other papers which are in possession
of the House, and in a state to be acted on, are arranged every
morning and brought on in the following order:]
[1. Bills ready for a second reading are read, that they may be
referred to committees, and so be put under way. But if, on their
being read, no motion is made for commitment, they are then laid on
the table in the general file, to be taken up in their just turn.]
[2. After 12 o’clock, bills ready for it are put on their passage.]
[3. Reports in possession of the House, which offer grounds for a
bill, are to be taken up, that the bill may be ordered in.]
[4. Bills or other matters before the House, and unfinished on the
preceding day, whether taken up in turn or on special order, are
entitled to be resumed and passed on through their present stage.]
[5. These matters being dispatched, for preparing and expediting
business, the general file of bills and other papers is then taken up,
and each article of it is brought on according to its seniority,
reckoned by the date of its first introduction to the House. Reports
on bills belong to the dates of their bills.]
[The arrangement of the business of the Senate is now as follows:]
[98]

[1. Motions previously submitted.]

[2. Reports of committees previously made.]
[3. Bills from the House of Representatives, and those introduced
on leave, which have been read the first time, are read the second
time; and if not referred to a committee, are considered in
Committee of the Whole, and proceeded with as in other cases.]
[4. After twelve o’clock, engrossed bills of the Senate, and bills of
the House of Representatives, on third reading, are put on their
passage.]
 [5. If the above are finished before one o’clock, the general file of
bills, consisting of those reported from committees on the second
reading, and those reported from committees after having been
referred, are taken up in the order in which they were reported to the
Senate by the respective committees.]
[6. At one o’clock, if no business be pending, or if no motion be
made to proceed to other business, the special orders are called, at
the head of which stands the unfinished business of the preceding
day.]
[In this way we do not waste our time in debating what shall be
taken up. We do one thing at a time; follow up a subject while it is
fresh, and till it is done with; clear the House of business gradatim as
it is brought on, and prevent, to a certain degree, its immense
accumulation toward the close of the session.]
[Arrangement, however, can only take hold of matters in
possession of the House. New matter may be moved at any time
when no question is before the House. Such are original motions and
reports on bills. Such are bills from the other House, which are
received at all times, and receive their first reading as soon as the
question then before the House is disposed of; and bills brought in
on leave, which are read first whenever presented. So messages from
the other House respecting amendments to bills are taken up as soon
as the House is clear of a question, unless they require to be printed,
for better consideration. Orders of the day may be called for even
when another question is before the House.]

SEC. XV.—ORDER.

[Each House may determine the rules of its proceedings; punish

its members for disorderly behavior; and, with the concurrence of
two-thirds, expel a member. Const., I, 5.]
In Parliament, “instances make order,” per Speaker Onslow. 2
Hats., 141. But what is done only by one Parliament, cannot be called
custom of Parliament, by Prynne. 1 Grey, 52.

SEC. XVI.—ORDER RESPECTING PAPERS.

 The Clerk is to let no journals, records, accounts, or papers be
taken from the table or out of his custody. 2 Hats., 193, 194.
Mr. Prynne, having at a Committee of the Whole amended a
mistake in a bill without order or knowledge of the committee, was
reprimanded. 1 Chand., 77.
A bill being missing, the House resolved that a protestation should
be made and subscribed by the members “before Almighty God, and
this honorable House, that neither myself, nor any other to my
knowledge, have taken away, or do at this present conceal a bill
entitled,” &c. 5 Grey, 202.
After a bill is engrossed, it is put into the Speaker’s hands, and he
is not to let any one have it to look into. Town. col., 209.

SEC. XVII.—ORDER IN DEBATE.

When the Speaker is seated in his chair, every member is to sit in

his place. Scob., 6; Grey, 403.
When any member means to speak, he is to stand up in his place,
uncovered, and to address himself, not to the House, or any
particular member, but to the Speaker, who calls him by his name,
that the House may take notice who it is that speaks. Scob., 6;
D’Ewes, 487, col. 1; 2 Hats., 77; 4 Grey, 66; 8 Grey, 108. But
members who are indisposed may be indulged to speak sitting. 2
Hats., 75, 77; 1 Grey, 143.
[In Senate, every member, when he speaks, shall address the Chair
standing in his place, and, when he has finished, shall sit down. Rule
3.]
When a member stands up to speak, no question is to be put, but
he is to be heard, unless the House overrule him. 4 Grey, 390; 5
Grey, 6, 143.
If two or more rise to speak nearly together, the Speaker
determines who was first up, and calls him by name, whereupon he
proceeds, unless he voluntarily sits down and gives way to the other.
But sometimes the House does not acquiesce in the Speaker’s
decision, in which case the question is put, “which member was first
up?” 2 Hats., 76; Scob., 7; D’Ewes, 434, col. 1, 2.
 [In the Senate of the United States, the President’s decision is
without appeal. Their rule is: When two members rise at the same
time, the President shall name the person to speak; but in all cases
the member who shall first rise and address the Chair shall speak
first. Rule 38.]
No man may speak more than once on the same bill on the same
day; or even on another day, if the debate be adjourned. But if it be
read more than once in the same day, he may speak once at every
reading. Co., 12, 115; Hakew., 148; Scob., 58; 2 Hats., 75. Even a
change of opinion does not give a right to be heard a second time.
Smyth’s Comw., L. 2, c. 3; Arcan. Parl., 17.
[The corresponding rule of the Senate is in these words: No
member shall speak more than twice, in any one debate, on the same
day, without leave of the Senate. Rule 39.]
But he may be permitted to speak again to clear a matter of fact, 3
Grey, 357, 416; or merely to explain himself 2 Hats., 73, in some
material part of his speech, Ib., 75; or to the manner or words of the
question, keeping himself to that only, and not traveling into the
merits of it, Memorials in Hakew., 29; or to the orders of the House
if they be transgressed, keeping within that line, and not falling into
the matter itself. Mem. Hakew., 30, 31.
But if the Speaker rise to speak, the member standing up ought to
sit down, that he may be first heard. Town., col. 205; Hale Parl., 133;
Mem. in Hakew., 30, 31. Nevertheless, though the Speaker may of
right speak to matters of order, and be first heard, he is restrained
from speaking on any other subject, except where the House have
occasion for facts within his knowledge; then he may, with their
leave, state the matter of fact. 3 Grey, 38.
No one is to speak impertinently or beside the question,
superfluous, or tediously. Scob., 31, 33; 2 Hats., 166, 168; Hale Parl.,
133.
No person is to use indecent language against the proceedings of
the House; no prior determination of which is to be reflected on by
any member, unless he means to conclude with a motion to rescind
it. 2 Hats., 169, 170; Rushw., p. 3, v. 1, fol. 42. But while a
proposition under consideration is still in fieri, though it has even
been reported by a committee, reflections on it are no reflections on
the House. 9 Grey, 508.
No person, in speaking, is to mention a member then present by
his name, but to describe him by his seat in the House, or who spoke
last, or on the other side of the question, &c., Mem. in Hakew., 3;
Smyth’s Comw., L. 2, c. 3; nor to digress from the matter to fall upon
the person Scob., 31; Hale Parl., 133; 2 Hats., 166 by speaking,
reviling, nipping, or unmannerly words against a particular member.
Smyth’s Comw., L. 2, c. 3. The consequences of a measure may be
reprobated in strong terms; but to arraign the motives of those who
propose to advocate it is a personality, and against order. Qui
digreditur a materia ad personam, Mr. Speaker ought to suppress.
Ord. Com., 1604, Apr. 19.
[When a member shall be called to order by the President or a
Senator, he shall sit down; and every question of order shall be
decided by the President, without debate, subject to an appeal to the
Senate; and the President may call for the sense of the Senate on any
question of order. Rule 40.]
[No member shall speak to another or otherwise interrupt the
business of the Senate, or read any newspapers while the journals or
public papers are being read, or when any member is speaking in any
debate. Rule 38.]
No one is to disturb another in his speech by hissing, coughing,
spitting, 6 Grey, 332; Scob., 8; D’Ewes, 332, col. 1, 640, col. 2,
speaking or whispering to another, Scob., 6; D’Ewes, 487, col. 1; nor
stand up to interrupt him, Town., col. 205; Mem. in Hakew., 31; nor
to pass between the Speaker and the speaking member, nor to go
across the House, Scob., 6, to walk up and down it, or to take books
or papers from the table or write there, 2 Hats., 171.
Nevertheless, if a member finds that it is not the inclination of the
House to hear him, and that by conversation or any other noise they
endeavor to drown his voice, it is his most prudent way to submit to
the pleasure of the House, and sit down; for it scarcely ever happens
that they are guilty of this piece of ill-manners without sufficient
reason, or inattentive to a member who says anything worth their
hearing. 2 Hats., 77, 78.
 If repeated calls do not produce order, the Speaker may call by his
name any member obstinately persisting in irregularity; whereupon
the House may require the member to withdraw. He is then to be
heard in exculpation, and to withdraw. Then the Speaker states the
offense committed; and the House considers the degree of
punishment they will inflict. 2 Hats., 167, 7, 8, 172.
For instances of assaults and affrays in the House of Commons,
and the proceedings thereon, see 1 Pet. Misc., 82; 3 Grey, 128; 4
Grey, 328; 5 Grey, 382; 6 Grey, 254; 10 Grey, 8. Whenever warm
words or an assault have passed between members, the House, for
the protection of their members, requires them to declare in their
places not to prosecute any quarrel, 3 Grey, 128, 293; 5 Grey, 280; or
orders them to attend the Speaker, who is to accommodate their
differences, and report to the House, 3 Grey, 419; and they are put
under restraint if they refuse, or until they do. 9 Grey, 234, 312.
Disorderly works are not to be noticed till the member has finished
his speech. 5 Grey, 356; 6 Grey, 60. Then the person objecting to
them, and desiring them to be taken down by the Clerk at the table,
must repeat them. The Speaker then may direct the Clerk to take
them down in his minutes; but if he thinks them not disorderly, he
delays the direction. If the call becomes pretty general, he orders the
Clerk to take them down, as stated by the objecting member. They
are then a part of his minutes, and when read to the offending
member, he may deny they were his words, and the House must then
decide by a question whether they are his words or not. Then the
member may justify them, or explain the sense in which he used
them, or apologize. If the House is satisfied, no further proceeding is
necessary. But if two members still insist to take the sense of the
House, the member must withdraw before that question is stated,
and then the sense of the House is to be taken. 2 Hats., 199; 4 Grey,
170; 6 Grey, 59. When any member has spoken, or other business
intervened, after offensive words spoken, they cannot be taken notice
of for censure. And this is for the common security of all, and to
prevent mistakes which must happen if words are not taken down
immediately. Formerly they might be taken down at any time the
same day. 2 Hats, 196; Mem. in Hakew., 71; 3 Grey, 48; 9 Grey, 514.
Disorderly words spoken in a committee must be written down as
in the House; but the committee can only report them to the House
for animadversion. 6 Grey, 46.
[The rule of the Senate says: If the member be called to order by a
Senator for words spoken, the exceptionable words shall
immediately be taken down in writing, that the President may be
better able to judge of the matter. Rule 37.]
In Parliament, to speak irreverently or seditiously against the
King, is against order. Smyth’s Comw., L. 2, c. 3; 2 Hats., 170.
It is a breach of order in debate to notice what has been said on the
same subject in the other House, or the particular votes or majorities
on it there; because the opinion of each House should be left to its
own independency, not to be influenced by the proceedings of the
other; and the quoting them might beget reflections leading to a
misunderstanding between the two Houses. 2 Grey, 22.
Neither House can exercise any authority over a member or officer
of the other, but should complain to the House of which he is, and
leave the punishment to them. Where the complaint is of words
disrespectfully spoken by a member of another House, it is difficult
to obtain punishment, because of the rules supposed necessary to be
observed (as to the immediate noting down of words) for the security
of members. Therefore it is the duty of the House, and more
particularly of the Speaker, to interfere immediately, and not to
permit expressions to go unnoticed which may give a ground of
complaint to the other House, and introduce proceedings and mutual
accusations between the two Houses, which can hardly be
terminated without difficulty and disorder. 3 Hats., 51.
No member may be present when a bill or any business concerning
himself is debating; nor is any member to speak to the merits of it till
he withdraws. 2 Hats., 219. The rule is, that if a charge against a
member arise out of a report of a committee, or examination of
witnesses in the House, as the member knows from that to what
points he is to direct his exculpation, he may be heard to those points
before any question is moved or stated against him. He is then to be
heard, and withdraw before any question is moved. But if the
question itself is the charge, as for breach of order or matter arising
in the debate, then the charge must be stated, (that is, the question
must be moved,) himself heard, and then to withdraw. 2 Hats., 121,
122.
 Where the private interests of a member are concerned in a bill or
question he is to withdraw. And where such an interest has
appeared, his voice has been disallowed, even after a division. In a
case so contrary, not only to the laws of decency, but to the
fundamental principle of the social compact, which denies to any
man to be a judge in his own cause, it is for the honor of the House
that this rule of immemorial observance should be strictly adhered
to. 2 Hats., 119, 121; 6 Grey, 368.
No member is to come into the House with his head covered, nor
to remove from one place to another with his hat on, nor is he to put
on his hat in coming in or removing, until he be set down in his
place. Scob., 6.
A question of order may be adjourned to give time to look into
precedents. 2 Hats., 118.
In Parliament, all decisions of the Speaker may be controlled by
the House. 3 Grey, 319.

SEC. XVIII.—ORDERS OF THE HOUSE.

Of right, the door of the House ought not to be shut, but to be kept
by porters, or Sergeants-at-Arms, assigned for that purpose. Mod.
ten. Parl., 23.
[By the rules of the Senate, on motion made and seconded to shut
the doors of the Senate on the discussion of any business which may,
in the opinion of a member, require secrecy, the President shall
direct the gallery to be cleared; and during the discussion of such
motion the doors shall remain shut. Rule 64.]
[No motion shall be deemed in order to admit any person or
persons whatsoever within the doors of the Senate chamber to
present any petition, memorial, or address, or to hear any such read.
Rule 19.]
The only case where a member has a right to insist on anything, is
where he calls for the execution of a subsisting order of the House.
Here, there having been already a resolution, any person has a right
to insist that the Speaker, or any other whose duty it is, shall carry it
into execution; and no debate or delay can be had on it. Thus any
member has a right to have the House or gallery cleared of strangers,
an order existing for that purpose; or to have the House told when
there is not a quorum present. 2 Hats., 87, 129. How far an order of
the House is binding, see Hakew., 392.
But where an order is made that any particular matter be taken up
on a particular day, there a question is to be put, when it is called for,
whether the House will now proceed to that matter? Where orders of
the day are on important or interesting matter, they ought not to be
proceeded on till an hour at which the House is usually full, [which
in Senate is at noon.]
Orders of the day may be discharged at any time, and a new one
made for a different day. 3 Grey, 48, 313.
When a session is drawing to a close, and the important bills are
all brought in, the House, in order to prevent interruption by further
unimportant bills, sometimes comes to a resolution that no new bill
be brought in, except it be sent from the other House. 3 Grey, 156.
All orders of the House determine with the session; and one taken
under such an order may, after the session is ended, be discharged
on a habeas corpus. Raym., 120; Jacob’s L. D. by Ruffhead;
Parliament, 1 Lev., 165, Pritchard’s case.
[Where the Constitution authorizes each House to determine the
rules of its proceedings, it must mean in those cases (legislative,
executive, or judiciary) submitted to them by the Constitution, or in
something relating to these, and necessary toward their execution.
But orders and resolutions are sometimes entered in the journals
having no relation to these, such as acceptances of invitations to
attend orations, take part in processions, &c. These must be
understood to be merely conventional among those who are willing
to participate in the ceremony, and are therefore, perhaps,
improperly placed among the records of the House.]

SEC. XIX.—PETITION.

A petition prays something. A remonstrance has no prayer. 1 Grey,

58.
 Petitions must be subscribed by the petitioners, Scob., 87; L. Parl,
c. 22; 9 Grey, 362, unless they are attending, 1 Grey, 401, or unable
to sign, and averred by a member, 3 Grey, 418. But a petition not
subscribed, but which the member presenting it affirmed to be all in
the handwriting of the petitioner, and his name written in the
beginning, was on the question (March 14, 1800) received by the
Senate. The averment of a member, or of somebody without doors,
that they know the handwriting of the petitioners, is necessary, if it
be questioned. 6 Grey, 36. It must be presented by a member—not
by the petitioners, and must be opened by him holding it in his hand.
10 Grey, 57.
[Before any petition or memorial addressed to the Senate shall be
received and read at the table, whether the same shall be introduced
by the President or a member, a brief statement of the contents of
the petition or memorial shall verbally be made by the introducer.
Rule 14.]
Regularly a motion for receiving it must be made and seconded,
and a question put, whether it shall be received? but a cry from the
House of “received,” or even its silence, dispenses with the formality
of this question. It is then to be read at the table and disposed of.

SEC. XX.—MOTIONS.

When a motion has been made, it is not to be put to the question

or debated until it is seconded. Scob., 21.
[The Senate says: No motion shall be debated until the same shall
be seconded. Rule 42.]
It is then, and not till then, in possession of the House, and cannot
be withdrawn but by leave of the House. It is to be put into writing, if
the House or Speaker require it, and must be read to the House by
the Speaker as often as any member desires it for his information. 2
Hats., 82.
[The rule of the Senate is, when a motion shall be made and
seconded, it shall be reduced to writing, if desired by the President or
any member, delivered in at the table, and read by the President,
before the same shall be debated. Rule 42.]
 It might be asked whether a motion for adjournment or for the
orders of the day can be made by one member while another is
speaking? It cannot. When two members offer to speak, he who rose
first is to be heard, and it is a breach of order in another to interrupt
him, unless by calling him to order if he departs from it. And the
question of order being decided, he is still to be heard through. A call
for adjournment, or for the order of the day, or for the question, by
gentlemen from their seats, is not a motion. No motion can be made
without rising and addressing the Chair. Such calls are themselves
breaches of order, which, though the member who has risen may
respect, as an expression of impatience of the House against further
debate, yet, if he chooses, he has a right to go on.

SEC. XXI.—RESOLUTIONS.

When the House commands, it is by an “order.” But fact,

principles, and their own opinions and purposes, are expressed in
the form of resolutions.
[A resolution for an allowance of money to the clerks being moved,
it was objected to as not in order, and so ruled by the Chair; but on
appeal to the Senate, (i. e., a call for their sense by the President, on
account of doubt in his mind, according to Rule 6,) the decision was
overruled. Jour. Senate, June 1, 1796. I presume the doubt was,
whether an allowance of money could be made otherwise than by
bill.]

SEC. XXII.—BILLS.

[Every bill shall receive three readings previous to its being

passed; and the President shall give notice at each whether it be first,
second, or third; which readings shall be on three different days,
unless the Senate unanimously direct otherwise. Rule 23.]

SEC. XXIII.—BILLS, LEAVE TO BRING IN.

 [One day’s notice, at least, shall be given of an intended motion for
leave to bring in a bill. Rule 22.]
When a member desires to bring in a bill on any subject, he states
to the House in general terms the causes for doing it, and concludes
by moving for leave to bring in a bill, entitled, &c. Leave being given,
on the question, a committee is appointed to prepare and bring in
the bill. The mover and seconder are always appointed of this
committee, and one or more in addition. Hakew., 132; Scob., 40.
It is to be presented fairly written, without any erasure or
interlineation, or the Speaker may refuse it. Scob., 41; 1 Grey, 82, 84.

SEC. XXIV.—BILLS, FIRST READING.

When a bill is first presented, the Clerk reads it at the table, and
hands it to the Speaker, who, rising, states to the House the title of
the bill; that this is the first time of reading it; and the question will
be, whether it shall be read a second time? then sitting down to give
an opening for objections. If none be made, he rises again, and puts
the question, whether it shall be read a second time? Hakew, 137,
141. A bill cannot be amended on the first reading, 6 Grey, 286; nor
is it usual for it to be opposed then, but it may be done, and rejected.
D’Ewes, 335, col. 1; 3 Hats., 198.

SEC. XXV.—BILLS, SECOND READING.

The second reading must regularly be on another day. Hakew.,

143. It is done by the Clerk at the table, who then hands it to the
Speaker. The Speaker, rising, states to the House the title of the bill;
that this is the second time of reading it; and that the question will
be, whether it shall be committed, or engrossed and read a third
time? But if the bill came from the other House, as it always comes
engrossed, he states that the question will be read a third time? and
before he has so reported the state of the bill, no one is to speak to it.
Hakew., 143–146.
[In the Senate of the United States, the President reports the title
of the bill; that this is the second time of reading it; that it is now to
be considered as in a Committee of the Whole; and the question will
be, whether it shall be read a third time? or that it may be referred to
a special committee?]

SEC. XXVI.—BILLS, COMMITMENT.

If on motion and question it be decided that the bill shall be

committed, it may then be moved to be referred to Committee of the
Whole House, or to a special committee. If the latter, the Speaker
proceeds to name the committee. Any member also may name a
single person, and the Clerk is to write him down as of the
committee. But the House have a controlling power over the names
and number, if a question be moved against any one; and may in any
case put in and put out whom they please.
Those who take exceptions to some particulars in the bill are to be
of the committee, but none who speak directly against the body of
the bill; for he that would totally destroy will not amend it, Hakew.,
146; Town., col. 208; D’Ewes, 634, col. 2; Scob., 47, or, as is said, 5
Grey, 145, the child is not to be put to a nurse that cares not for it, 6
Grey, 373. It is therefore a constant rule “that no man is to be
employed in any matter who has declared himself against it.” And
when any member who is against the bill hears himself named of its
committee, he ought to ask to be excused. Thus, March 7, 1606, Mr.
Hadley was, on the question being put, excused from being of a
committee, declaring himself to be against the matter itself. Scob.,
46.
[No bill shall be committed or amended until it shall have been
twice read; after which it may be referred to a committee. Rule 24.]
In the appointment of the standing committees, the Senate will
proceed, by ballot, severally to appoint the chairman of each
committee, and then, by one ballot, the other members necessary to
complete the same; and a majority of the whole number of votes
given shall be necessary to the choice of a chairman of a standing
committee. All other committees shall be appointed by ballot, and a
plurality of votes shall make a choice. When any subject or matter
shall have been referred to a committee, any other subject or matter
of a similar nature, may, on motion, be referred to such committee.
 The Clerk may deliver the bill to any member of the committee,
Town., col. 138; but it is usual to deliver it to him who is first named.
In some cases the House has ordered a committee to withdraw
immediately into the committee chamber, and act on and bring back
the bill, sitting the House, Scob., 48. A committee meet when and
where they please, if the House has not ordered time and place for
them, 6 Grey, 370; but they can only act when together, and not by
separate consultation and consent—nothing being the report of the
committee but what has been agreed to in committee actually
assembled.
A majority of the committee constitutes a quorum for business.
Elsynge’s Method of Passing Bills, 11.
Any member of the House may be present at any select committee,
but cannot vote, and must give place to all of the committee, and sit
below them. Elsynge, 12; Scob., 49.
The committee have full power over the bill or other paper
committed to them, except that they cannot change the title or
subject. 8 Grey, 228.
The paper before a committee, whether select or of the whole, may
be a bill, resolutions, draught of an address, &c., and it may either
originate with them or be referred to them. In every case the whole
paper is read first by the Clerk, and then by the chairman, by
paragraphs, Scob., 49, pausing at the end of each paragraph, and
putting questions for amending, if proposed. In case of resolutions
on distinct subjects, originating with themselves, a question is put on
each separately, as amended or unamended, and no final question on
the whole, 3 Hats., 276; but if they relate to the same subject, a
question is put on the whole. If it be a bill, draught of an address, or
other paper originating with them, they proceed by paragraphs,
putting questions for amending, either by insertion or striking out, if
proposed; but no question on agreeing to the paragraphs separately;
this is reserved to the close, when a question is put on the whole, for
agreeing to it as amended or unamended. But if it be a paper referred
to them, they proceed to put questions of amendment, if proposed,
but no final question on the whole; because all parts of the paper,
having been adopted by the House, stand, of course, unless altered or
struck out by a vote. Even if they are opposed to the whole paper, and
think it cannot be made good by amendments, they cannot reject it,
but must report it back to the House without amendments, and there
make their opposition.
The natural order in considering and amending any paper is, to
begin at the beginning, and proceed through it by paragraphs; and
this order is so strictly adhered to in Parliament, that when a latter
part has been amended, you cannot recur back and make any
alteration in a former part. 2 Hats., 90. In numerous assemblies this
restraint is doubtless important. [But in the Senate of the United
States, though in the main we consider and amend the paragraphs in
their natural order, yet recurrences are indulged; and they seem, on
the whole, in that small body, to produce advantages overweighing
their inconveniences.]
To this natural order of beginning at the beginning, there is a
single exception found in parliamentary usage. When a bill is taken
up in committee, or on its second reading, they postpone the
preamble till the other parts of the bill are gone through. The reason
is, that on consideration of the body of the bill such alterations may
therein be made as may also occasion the alteration of the preamble.
Scob., 50; 7 Grey, 431.
On this head the following case occurred in the Senate, March 6,
1800: A resolution which had no preamble having been already
amended by the House so that a few words only of the original
remained in it, a motion was made to prefix a preamble, which
having an aspect very different from the resolution, the mover
intimated that he should afterwards propose a correspondent
amendment in the body of the resolution. It was objected that a
preamble could not be taken up till the body of the resolution is done
with; but the preamble was received, because we are in fact through
the body of the resolution; we have amended that as far as
amendments have been offered, and, indeed, till little of the original
is left. It is the proper time, therefore, to consider a preamble; and
whether the one offered be consistent with the resolution is for the
House to determine. The mover, indeed, has intimated that he shall
offer a subsequent proposition for the body of the resolution; but the
House is not in possession of it; it remains in his breast, and may be
withheld. The rules of the House can only operate on what is before
them. [The practice of the Senate, too, allows recurrences backward
and forward for the purposes of amendment, not permitting