Arch Virol (2011) 156:1691–1705
DOI 10.1007/s00705-011-1080-2
BRIEF REVIEW
Pseudorabies virus in wild swine: a global perspective
T. Müller • E. C. Hahn • F. Tottewitz •
M. Kramer • B. G. Klupp • T. C. Mettenleiter •
C. Freuling
Received: 25 May 2011 / Accepted: 21 July 2011 / Published online: 12 August 2011
Ó Springer-Verlag 2011
Abstract Suid herpesvirus 1 (SuHV1, syn. Aujeszky’s
disease virus [ADV] or pseudorabies virus [PrV]), which
belongs to the family Herpesviridae, subfamily Alphaher-
pesvirinae, genus Varicellovirus is the causative agent of
Aujeszky’s disease (AD, pseudorabies), a notifiable dis-
ease, that causes substantial economic losses to the swine
industry in countries, where AD is present. Members of the
family Suidae (true pigs) are the only natural hosts for PrV,
although the virus can infect numerous other mammals
including ruminants, carnivores and rodents. Despite the
tremendous progress that has been made in controlling and
eliminating PrV in domestic pigs, there is mounting evi-
dence that PrV infections are more widespread in wild
swine across the world than originally thought. Unfortu-
nately, our understanding of the extent of PrV infections in
these wild populations and of the threat to domestic swine
is still fragmentary. This review aims at giving a global
perspective on PrV infections in wild swine by scrutinizing
the current state of knowledge concerning (i) the global
T. Müller (&)
M. Kramer
C. Freuling
Institute of Epidemiology, Friedrich-Loeffler-Institut (FLI),
Federal Research Institute for Animal Health, Seestrasse 55,
16868 Wusterhausen, Germany
e-mail: Thomas.Mueller@fli.bund.de
E. C. Hahn
Department of Veterinary Pathobiology, University of Illinois,
Urbana-Champaign 61802, IL, USA
F. Tottewitz
Institute for Forest Ecology and Forest Inventory, Johann
Heinrich von Thünen-Institut, 16225 Eberswalde, Germany
B. G. Klupp
T. C. Mettenleiter
Institute of Molecular Biology, Friedrich-Loeffler-Institut (FLI),
Federal Research Institute for Animal Health,
17493 Greifswald-Insel Riems, Germany
occurrence of PrV infections in free-living populations of
wild swine, e.g., wild boar and feral swine, (ii) the
molecular characterization of wild swine PrV, (iii) infec-
tion characteristics of PrV in populations of wild swine,
(iv) the risk of spillover infections to domestic pigs,
(v) potential risk-mitigating measures, focusing on further
research needs.
Introduction
Aujeszky’s disease (AD, pseudorabies) is a notifiable dis-
ease that causes substantial economic losses to the swine
industry in countries, where the disease is present. The
causative agent is Suid herpesvirus 1 (SuHV1, syn. Au-
jeszky’s disease virus [ADV] or pseudorabies virus [PrV]),
which belongs to the family Herpesviridae, subfamily
Alphaherpesvirinae, genus Varicellovirus [1]. Members of
the family Suidae (true pigs) are the only natural hosts for
PrV, although the virus can infect numerous other mam-
mals, including ruminants, carnivores and rodents [2, 3].
Since the early 1980s, the disease had spread nearly
globally. This development was in part fuelled by the
emergence of more virulent PrV strains and changes in
swine management, notably cohabitation and total con-
finement of large numbers of pigs and continuous farrow-
ing. Due to increased control efforts and the strict
implementation of national eradication programs, which
have included large-scale vaccination with gE-deleted
vaccines and the so-called DIVA (‘‘differentiating infected
from vaccinated animals’’) strategy, AD has virtually dis-
appeared from domestic pigs in several parts of Europe,
e.g., Austria, Cyprus, the Czech Republic, Denmark, Fin-
land, France (except a few departments), Germany, Hun-
gary, Luxembourg, the Netherlands, Sweden, Switzerland,
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Slovakia, Norway, and Great Britain (England, Scotland
and Wales). More recently, Canada, New Zealand and the
United States also became free of AD [4–6]. In countries
with an officially recognised AD-free status, vaccination
against PrV is prohibited. Today, PrV remains one of the
most important diseases of domestic pigs worldwide, par-
ticularly occurring in regions with dense pig populations,
i.e., some regions in Europe, Latin America and Asia.
Therefore, it remains included as an OIE notifiable disease
[7].
However, despite the achievements in eliminating PrV
in domestic pigs in many parts of the world, due to their
evolutionary relatedness to the domestic pig, the role of
wild swine as potential reservoirs has become increasingly
important. Wild boar (Sus scrofa, L. 1758) are considered
to be reservoirs for several viruses of livestock and
humans, including classical (CSFV) and African swine
fever viruses (ASFV), porcine circovirus type 2 (PCV2),
porcine reproductive and respiratory syndrome virus
(PRRSV), porcine parvovirus (PPV), hepatitis E virus
(HEV), swine influenza virus and Japanese encephalitis
virus (JEV) (for review see ref. 8). European wild boar and
feral swine seem to serve as a persistent reservoir for PrV
[8–12]. Unfortunately, however, although wild boar are
indigenous in many countries in the world, full apprecia-
tion of the extent of PrV infections in these wild popula-
tions and of the threat to domestic swine is still
fragmentary [13]. Since the first review on PrV infections
in wild pigs was published in 2000 [10], increased attention
has been paid to gaining more information on disease
epidemiology at the regional level. Therefore, this review
provides an update of the current knowledge of PrV
infections in populations of wild swine around the world
and attempts to elucidate their possible impact on contin-
uing PrV eradication programmes and the risks to the
PrV-free status.
Taxonomy and genetic relationship of wild swine
to domestic pigs
Sus scrofa (L. 1758) is the scientific name for wild and
domesticated breeds of the pig genus Sus, family Suidae,
artiodactyl suborder Suina. ‘‘Wild swine’’, as used
throughout this manuscript is an umbrella term for both
true wild boar and feral swine. The true wild boar repre-
sents genuine wildlife in Europe and Asia, comprising
nearly 23 subspecies in 4 subspecies groups, e.g., Western
breeds (scrofa group), Indian breeds (cristatus group),
Eastern breeds (leucomystax group) and Sundaic breeds
(vittatus group) [14], with a sharp genetic differentiation
between eastern Asian and European populations [15]. The
true wild boar is considered the ancestor of modern
123
T. Müller et al.
domestic pig hybrids, with several independent domesti-
cations across Eurasia [16]. Although, wild boar can
hybridize with domestic breeds, molecular studies indicate,
that this contributes only marginally to the diversity of wild
boar in Europe [15].
In contrast, the so-called feral swine or feral pig in the
USA originates from at least two ancestral pools of
(i) multiple introductions of European wild boar and (ii)
pigs descendent from domestic stock that have been
released during the last centuries. They easily interbreed,
producing descendants similar in appearance to wild boar,
making it difficult to distinguish them from true wild boar
[17]. However, the degree of crossbreeding in feral swine
may differ from region to region. Whether populations are
characterised as feral swine, i.e., domestic pigs run wild, or
introduced wild boar, is usually decided by where the
animals are encountered and what is known of their history.
Global distribution of wild swine
The wild boar is the most widely distributed species of the
Old World Suidae and has one of the widest geographic
distributions of all terrestrial mammals. This range has
been greatly expanded by human activity. The species now
occurs in pure wild or barely modified feral form on all
continents except Antarctica, and on many oceanic islands
[18]. In Europe, wild boar regained territories in the latter
half of the 20th century after they had almost disappeared
from many parts by the end of the 17th century and during
the two world wars [19]. Their current range extends from
western Europe and the Mediterranean basin, from north-
ern Africa (Atlas Mountains) to southern Scandinavia and
the far eastern parts of the Eurasian continent, including the
islands of Japan, and through India and Southeast Asia as
far as Indonesia [14], and there is evidence that this con-
tinental expansion is still continuing [19].
Since the first introduction of pigs to the Americas by
the Polynesians and the early Spanish explorers in the 16th
century, multiple introductions of European wild boar and
releases of domestic swine have occurred in the USA. In
the recent past, due to translocation of animals for hunting
purposes and regular movement of the population in search
of new habitats, crossbreeds of wild swine populations
have expanded into many of the central states of the US
and even Canada, with expansions now including semi-arid
areas [20]. Currently, wild swine are distributed across 39
of the 50 United States [21]. The National Feral Swine
Reporting System monitors the distribution of feral swine
in the US with monthly updates (http://128.192.20.53/
nfsms/). European wild boar has also been introduced by
humans, mainly for hunting, to South America, New Gui-
nea, New Zealand, Australia and other islands. Elsewhere,
Pseudorabies virus in wild swine
populations also have become established after escape of
wild boar from captivity [22]. Wild swine have adapted
easily to these different environments and established
considerable populations in many areas of these continents,
where they have interbred with free-roaming domestic
pigs. Similarly, feral swine are abundant in certain regions
of Europe, e.g., Corsica, Sardinia, Croatia, Rumania and
Hungary [10, 23].
Population densities
The population densities of wild boar in western Eurasia
show high variation [12, 24], with up to 90 individuals per
sqkm in parts of Spain [25]. In fact, the wild boar is the
second most abundant ungulate in Europe [23]. On a bio-
geographical scale, there is a west-to-east decline in den-
sities, primarily affected by colder climate and diminishing
food resources [24]. Since the 1950s, populations of wild
ungulates, and particularly wild boar, exploded, which
increased their range and, where they were already estab-
lished, their density [23]. Based on official hunting statis-
tics, Germany and France are believed to have among the
highest population densities of wild boar in Europe, fol-
lowed by Spain, Poland and the Czech Republic (Table 1).
Although no hunting statistics are available for Italy, the
wild boar population is believed to amount to 600,000
individuals [23]. Several reasons for the rapid population
increase have been postulated, e.g., radical changes in
agricultural land use, climate change, lack of predator
species, increasing mast seeding years of oak (Quercus
ssp.) and beech (Fagus ssp.) trees, reintroductions due to
human activities, uncontrolled growth, and low hunting
pressure (reviewed in ref. 26).
A similar trend can be observed in the USA, where feral
pigs have increased in numbers and expanded their range
[27]. Over the last 10 years, the feral pig population has
quadrupled and is distributed across 39 states [21], mainly
resulting from intentional or accidental introductions or the
dispersal of established feral populations [28]. Unfortu-
nately, there are no current density data available regarding
feral swine in the United States. In general, the increasing
range and density also provide ample opportunities for
direct human and domestic swine contacts with possible
transmission of pathogens, including PrV [8].
Occurrence of PrV infection in wild swine populations
Serological surveys on PrV in wild swine are largely lim-
ited to the current disease situation, driven by public
interest and funding [10]. Despite the tremendous progress
made toward controlling and eliminating AD in domestic
1693
pigs, there is mounting evidence that PrV infections are
more widespread in wild swine across the world than
originally thought. First indications of the occurrence of
PrV in wild swine in the USA and certain regions of
Europe were provided as early as the mid 1980s [10]. In the
USA, where serological field studies on PrV have been
conducted for three decades, PrV is endemic in feral swine
populations in the southern and central states (Table 2).
Recently, PrV has also been found in isolated groups of
feral swine in Nebraska and Michigan [29]. The source
of these infections could be traced to illegal introduction of
feral swine from the southeast. While the overall sero-
prevalence was determined to be about 19% nationwide
[30], periodic seroprevalence can vary over a wide range,
between 0.5% and 61% at the regional level, depending on
the sample size (Table 1).
In Europe, since the year 2000, PrV infections in wild
boar populations have received considerable attention.
Serosurveys revealed the occurrence of PrV infections in
wild boar in at least 14 European countries, including
Belgium, Croatia, the Czech Republic, the former Yugo-
slavia, France, Germany, Italy, the Netherlands, Poland,
Romania, Russia, Slovenia, Spain, and Switzerland, with
the average seroprevalence for the investigated time peri-
ods ranging between 0.3% and 66% at the national or
regional level. Only in Lithuania and Sweden did PrV
infections seem to be absent in wild boar (Table 3). PrV
infections in wild boar were also reported in three prov-
inces (Atlas Mountains) of Tunisia for the time period
1993-1995, with an overall seroprevalence of 63% [31].
Also, 42.8% of tested wild boar sera (N = 28) from Iran
were positive [32].
In contrast, serological surveys conducted in Japan and
New Zealand between 1993 and 1995 revealed no evidence
for infections of wild swine with PrV [33, 34]. The exact
prevalence of PrV infection in populations of wild swine in
other parts of the world is not known.
In general, the reported seroprevalence in populations of
wild boar and feral swine has to be interpreted with care,
since these findings may have been biased by sampling, the
serological assays used or the investigation periods. Sero-
prevalence over continuous time periods, for example, had
only been studied in Germany, France, Italy, the Czech
Republic and the US state of Florida. While in Europe,
samples sizes ranged between a few dozen and several tens
of thousands, the largest sample size in the US amounted to
about 1660 (Tables 2 and 3). In general, the larger the
sample size, the more accurate the estimated seropreva-
lence (Tables 2 and 3). With more than 100,000 wild boar
sera tested over a 24-year period, East Germany is the most
intensively studied area ever. Here, seroprevalences can
exceed 40%, and a ‘‘steady state’’ may have been reached
[35]. PrV infections in wild boar in this area appeared to be
123
1694 T. Müller et al.

Table 1 Records and approximation of hunting bag, and population estimates for wild boar in European countries during past decades according
to ref. 25
Country Hunting bag
Lowest record Highest record Population
Year Number Year Number Year Estimates

Austria 1948 \1,000 2004 32,500

Belgium 2006 21,000
Croatia 2003 18,200
Czech Republic 1966 \5,000 2004 121,000
Denmark 2010 \25 Present
Estonia 1995 \2,500 2005 17,000
Finland 2010 \100
France 1973 40,000 2003 475,000
Germany 1984 180,000 2009 500,000
Great Britain 2004 230
Greece Present
Hungary 1960 3,900 2002 93,962
Ireland Not present
Italy 2005 600,000
Latvia 1923 \2,500 2005 47,000
Lithuania 1933 \2,500 2005 29,000
Macedonia 2003 525 2005 712
Netherlands 2010 2,300
Norway Not present
Poland 1975 40,000 2005 136,000
Portugal 1989 423 2000 8,000
Romania 1968 3,967 2005 10,714
Serbia 1995 1,356 1991/2000 3,783/3,000
Slovakia 1950 \2,000 2003 22,000
Slovenia 1975 1,078 2005 6,569
Spain 2010 150,000
Sweden 1990 \1,000 2005 18,000
Switzerland Present

dynamic, with the locations with the highest seropreva-
lence varying over time. A dynamic ‘‘emergence and dis-
solution of pockets of infection’’, i.e., a local cyclic
accumulation of infection with movement into non-infec-
ted areas followed by a decrease in seroprevalence, has
been suggested [36].
PrV seropositivity also has been reported in wild boar
held in pens from Serbia and Germany, although there
was no contact to free-roaming wild boar in the vicinity
[37, 38].
Isolation of PrV of wild swine origin
Because PrV enters a stage of latency in infected Suidae,
virus isolation from wild swine is often difficult. However,
some PrV isolates have been obtained directly from feral
swine and from European wild boar [39–44]. A recent
study characterized PrV isolated from wild boar from dif-
ferent European countries, including Germany (N = 13),
France (N = 1), Spain (N = 8), Slovakia (N = 3), Hun-
gary (N = 2) and Italy (N = 1) [45]. The viruses were
either isolated from brain [42], oral and nasal swabs [43],
tonsils [40, 43, 44] or genital secretions (prepuces, vaginae)
[41, 43, 46] of mostly seronegative animals.
Due to the severe clinical course of PrV infections in
carnivores, many isolates were obtained from hunting dogs,
that had been in contact with potentially infected wild
swine and eventually succumbed to the disease. In France,
from 1997 to 2004, a total of twenty hunting dogs died due
to infection with PrV of wild boar origin [47]. In Germany,
12 hunting dogs died and tested PrV-positive during the

123
Pseudorabies virus in wild swine 1695

Table 2 Serosurveys of PrV infections conducted in Europe between 1979 and 2008
Country Region Wild Time Sample Number Percent Method Reference
boar period size positive positive

Austria Lower Austria Free-living 2010–2011 162 98 60.5

Belgium Hainaut Free-living 2004–2005 47 15 ELISA [69]
Namur Free-living 2004–2005 403 22
Luxembourg Free-living 2004–2005 1422 17
Liège Free-living 2004–2005 983 15
Croatia Moslavacka gora Free-living 1999 44 24 54.55 ELISA [87]
Brodsko-posavska, Karlovacka, Free-living 2000–2003 745 30.4 [88]
Koprivnicko-krizevacka, Osjecko-baranjska,
Sisacko-moslavacka, Vukovarsko-srijemska,
Zagrebacka
Czech Entire country Free-living 1995–2005 338 101 29.88 ELISA [89]
Republic
Former Croatian part Free-living 1988 96 53 55.21 SNT [37]
Yugoslavia
France Central and northeastern France Free-living 1979 143 0 0 SNT [90]
Southeastern and central France Free-living 1979 260 0 0 SNT [91]
Corsica Free-living 1991–1998 492 159 32.32 ELISA [92]
Continental France Free-living 6025 199 3.30 ELISA
Continental France Farmed 5508 65 1.18 ELISA
Continental France Free-living 2000–2004 6801 0–54 ELISA [93]
Corsica Free-living 2000–2004 53 ELISA
Germany Mecklenburg-Western Pommerania Free-living 1989 5000 13 0.26 SNT [94]
Lower Saxony Free-living 1990/1991 722 12 1.66 SNT [95]
Saxony-Anhalt/Brandenburg Free-living 1990/1991 640 13 2.03 ELISA [96]
North Rhine Westphalia Free-living 1992–1996 719 51 7.09 ELISA [97]
Brandenburg Free-living 1991–1994 3143 281 8.94 ELISA [66]
Baden-Wurttemberg Farmed 1996 141 52 36.88 ELISA [38]
North Rhine Westphalia Free-living 1996–2000 929 92 9.90 ELISA [44]
Mecklenburg-Western Pommerania Free-living 1997/1998 99 13 13.13 ELISA [98]
Free-living 2000/2001 322 58 18.01 ELISA
Brandenburg Free-living 2004/2005 166 26 15.66 ELISA
Saxony-Anhalt Free-living 2001/2002 254 15 5.91 ELISA
Baden-Wurttemberg Free-living 2001/2002 159 11 6.92 ELISA
Rhineland Palatinate Free-living 2005 199 12 6.03 ELISA
Mecklenburg-Western Pommerania Free-living 1985–2008 15466 535 3.46 SNT/ELISA [35]
Saxony-Anhalt Free-living 1985–2008 7184 177 2.46 SNT/ELISA
Brandenburg Free-living 1985–2008 15190 927 6.10 SNT/ELISA
Berlin Free-living 2000–2006 5020 234 4.66 SNT/ELISA
Saxony Free-living 1985–2008 45674 4856 10.63 SNT/ELISA
Thuringia Free-living 1985–2008 13853 66 0.48 SNT/ELISA
Italy Tuscany Free-living 1983–1984 20 0 0 SNT [99]
Sardinia Free-living 1989–1991 2609 753 28.86 ELISA [100]
Sardinia Free-living 1994–1995 2379 986 41.45 ELISA [101]
Sardinia Free-living 1993–1997 4027 1120 27.81 ELISA [102]
Grosseto district Free-living 2002/2003 152 41 26.97 ELISA [68]
Campania Free-living 2006/2006 342 105 30.70 ELISA [103]
Liguria Free-living 1992–1993 430 40 9.30 ELISA [104]
Bologna Free-living 89 6 ELISA [105]
Rome Free-living 1997–1999 136 0 ELISA
Maremma (Tuscany) Free-living 1997–1999 265 47 ELISA

123
1696 T. Müller et al.

Table 2 continued
Country Region Wild Time Sample Number Percent Method Reference
boar period size positive positive

Lithuania Entire country Free-living 2004 24 0 0.00 ELISA [106]

Netherlands Veluwe region Free-living 1994 115 3 2.61 SNT [107]
Free-living 1994–1999 0 ELISA [108]
Wildbeheer Veluwe, Roerstreek,Hoge Veluwe, Free-living 1996–1999 368 0 0 ELISA [109]
‘t Loo Crown dominion
Wildbeheer Veluwe, Roerstreek Midden Limburg, Free-living 1996–1999 364 0 0 ELISA [110]
Hoge Veluwe, Kroondomeinen
Wildbeheer Veluwe, Roerstreek, Midden Limburg, Free-living 1996–2001 878 0 0 ELISA [111]
Hoge Veluwe, Kroondomeinen
Poland Olstyn Free-living 1994–1996 34 11.8 ELISA Szweda, W.,
cited in [112]
Entire country Free-living 1997–1998 1816 235 12.94 [113]
Free-living 1998–1999 1429 278 19.45
Free-living 1999–2000 4791 663 13.84
Romania Southern parts Free-living 2008 357 197 55.18 ELISA [114]
Russia Wladimirska, Moskowskaja Twerskaja, Free-living 48 19 39.58 ELISA [115]
Belgorodskaja, Chabarowski Kraj, Charkowskaja
Belgorodskaya, Vladimirskaya,Moskovskaya, Free-living 2002–2007 83 27 32.53 ELISA [116]
Tverskaya, Smolenskaya, Kirovskaya
Oblasts and Chabarovsky Krai
Slovenia Entire country Free-living 2003/2004 427 111 26.00 ELISA [117]
Entire country Free-living 2003/2004 178 55 30.90 ELISA [118]
Entire country Free-living 2007 698 115 16.48 ELISA [119]
Entire country Free-living 2008 749 175 23.36 ELISA
Entire country Free-living 2009 672 115 17.11 ELISA
Spain Sierra Morena Montes de Toledo Free-living 1999–2000 78 8 10.26 ELISA [120]
Central spain 16 9 56.25 ELISA [11]
Asturia; Ebro Bassin, Jaen Free-living 2000–2003 693 308 44.44 ELISA [54]
South-central Spain Free-living 2000–2003 66 28 42.42 ELISA [121]
Farmed 2000–2003 218 144 66.06 ELISA
South-central Spain Free-living 2004/2005 185 85 45.95 ELISA [122]
Catalonia Free-living 2004–2007 273 0.80 ELISA [123]
Sweden Southern Sweden Free-living 2006 231 0 0 ELISA [124]
Southern Sweden Free-living 2007 334 1 0.30 ELISA [125]
Southern Sweden Free-living 2008 542 0 0 ELISA [126]
Southern Sweden Free-living 2009 500 0 0 ELISA [127]
Switzerland North Alps Free-living 2001–2003 1450 35 2.41 ELISA [128]
South Alps Free-living 2001–2003 407 28 6.88
Entire country Free-living 2004–2005 ca. 1000 ? 3.50 ELISA [129]
Entire country Free-living 2004–2005 1060 30 2.83 ELISA [85]

time from 2000 to 2010. Notably, there was an apparent
increase in recent years, with four dogs succumbing in
Germany in 2009 and 2010, of which two animals became
infected in Austria and one in Luxembourg [48]. In
neighboring Belgium, two hunting dogs also died after
infection with wild-boar transmitted PrV [49]. In Austria,
the first case of AD in a dog associated with wild boar was
reported in 2006 [50]. However, recently, another four
hunting dogs have been shown to have succumbed to PrV
infection in Austria after having had intensive contact with
wild boar [51].
Although wild carnivores have the highest chance of
contracting a PrV infection in areas where wild boar are
endemically infected while foraging and feeding on PrV-
infected wild swine cadavers, reports of wildlife being
infected with PrV through contact with wild swine are rare.
PrV has been detected in a panther (Puma concolor coryi)
from Florida [52], and red foxes (Vulpes vulpes) from
France and Germany. However, there is no direct evidence
for an association with wild swine. In Morocco, PrV was
isolated from African wild dogs (Lycaon pictus) held in
captivity that died of acute infection after being fed wild
boar meat [53]. It has been speculated that PrV infections
in wild boar may have an impact on conservation of
endangered wild carnivore populations in Spain, e.g., the
Iberian lynx (Lynx pardinus) and the Iberian wolf (Canis

123
Pseudorabies virus in wild swine 1697

Table 3 Serosurveys of PrV infections conducted in the Americas between 1979 and 2008
Country Region Feral swine Year Sample Number Percent Method Reference
size positive positive

USA California Free-living 1981–1983 130 4 3.08 SNT [130]

Southeast Free-living 1979–1983 423 22 [17]
Hawaii Free-living 116 46
Texas Free-living 1985 124 46 37.10 SNT [131]
Arizona Free-living 1986 192 1 0.52 SNT [132]
peccaries
Ossabaw Island Georgia Free-living 1981–1986 661 200 30.26 LAT/ [67]
MIDT
South Carolina Free-living 10.00 LAT/ Wood et al. (1992),
ELISA cited in [133]
Florida Free-living 1980–1989 1662 579 34.84 MIDT [13]
Great Smoky Mountains Free-living 1990 108 0 0 LAT/ [134]
ELISA
Oklahoma Free-living 1996 120 0 0 LAT/ [135]
ELISA
Kansas Free-living 1993/1994 20 0 0 LAT/ [20]
ELISA
South Carolina Free-living 1999 220 134 60.91 LAT/ [136]
ELISA
Texas, Alabama, South Carolina, Free-living 2001–2002 100 38 38.00 LAT [133]
Florida, Georgia,
Texas (East) Free-living 60 7 11.67 [137]
Texas (South) Free-living 42 16 38.10
Texas (East) Free-living 125 22 18.00 ELISA [75]
Texas (South) Free-living 244 87 35.66
Texas (South and Central) Free-living 2001–2007 409 145 35.45 LAT [138]
Great Smoky Mountains Free-living 497 16 3.22 ELISA [139]
South Carolina Free-living 2006–2007 20 10 50.00 LAT/ [140]
ELISA
North Carolina Free-living 120 0 0
North Carolina Free-living 2007–2009 482 0 0 ELISA [28]
Brazil Southeast Farmed 1998–2001 358 25.2–100 SNT/ [141]
ELISA

lupus signatus) [54], but a direct influence has not been
reported.
Molecular characterisation of PrV isolates of wild swine
origin
Isolation and characterisation of PrV from infected wild
swine can help to understand population diversity and may
facilitate tracing back the infection chain [10]. Classically,
the analysis of restriction fragment length polymorphisms
(RFLP) using BamHI restriction endonuclease was used to
characterize PrV isolates from domestic swine [55, 56].
The method is inexpensive and can detect variation across
large genomic regions [55]. Four major genome types
could be differentiated, with type I predominantly found in
the USA and Central Europe. Types II and III circulate in
Central and Northern Europe, respectively, whereas type
IV is restricted to Asia.
Using this technique, a comprehensive study of Euro-
pean PrV isolates of wild boar origin demonstrated that all
except one belonged to genotype I [45]. Based on the
observation that mainly type II strains were found in
domestic pigs in Central Europe, it has been suggested that
infections of wild boar by domestic pigs did not occur
recently [42, 57]. Within the major genome types, several
subtypes can be distinguished [55, 56]. Interestingly,
genetic profiles of East German wild boar PrV isolates

123
1698 T. Müller et al.

2874-ESP
FL82gC32
(a) FL81gC-1U
(b) 2873-ESP
2882-ESP
FL944855-gC1U 2885-ESP

H11725-1u 2886-ESP
2890-ESP
H351gC-1U
2960_ESP
HI354-gc1u 519-FRA
NCDog-gC1L 536-FRA
57 537-FRA
SC56AVH9909
550-GER_NRW
TXAndersonSOW-gC1L 551-GER_NRW
OK_18_PRV_F1_88 552-GER_NRW
553-GER_RP
OK_2_PRV_F1_72
554-GER_RP
OK_3_PRV_F1_81
56 555-GER_RP
OK_23_PRV_F1_93 556-GER_NRW
69
OK_21_PRV_F1_32 594-FRA
611-GER_RP
OK_8_PRV_F1_84
2872-ESP
OK_9_PRV_F1_96 561-ITA
67 OK_22_PRV_F1_92 558-SVK
54
54 559-SVK
TE-19F
549-SVK
OK_16_PRV_F1_24 560-SVK
TX11-1U- 613-GER_SN
612-GER_BRB
68 OK_17_PRV_F1_87 79
15-GER_BRB
CA35gC-1U
61 13-GER_BRB
67
AR9772LnL-PRV-F1-26 12-GER_BRB

66 TN669900gC21 11-GER_SA
10_GER_SA
MSferal031301-1U
92 641-GER
SC3-3-05gC1L 527-FRA
FL_6731B-FL-PRV-F1-42 576-HUN
61 563-HUN
AF158090
AF158090

0.005 0.005

Fig. 1 Phylogenetic trees of PrV of US feral swine (a, modified from GenBank AF158090) as an outgroup was used as implemented in
Hahn et al. [4]) and European wild boar (b, modified from Müller MEGA5. All positions containing gaps and missing data were
et al. [45]) using partial gC nucleotide sequences. The neighbor- eliminated. Bootstrap values (500 replications) greater than 50 are
joining method (Kimura 2-parameter) with the strain Ea (pig, China, indicated

differed considerably from known subtypes from domestic
animals [43]. Together with wild boar isolates from Hun-
gary, the East German isolates showed the same rare PrV
subtype, Iw (5-10/5-12 double fusion strains), confirming
previous associations with PrV isolates from domestic pig
and cats in Hungary. In contrast, the great majority of wild
boar PrV isolates from Europe showed BamHI restriction
patterns similar to the standard pattern, as exhibited by PrV
strain Kaplan, which has been identified as subtype Ip [45].
PrV strains isolated from wild boar and feral swine in
the USA were of subtypes Ip and Iip [42, 57, 58] and thus
distinct from European isolates and from strains isolated
from domestic swine. Interestingly, many isolates from
Florida and Georgia showed an additional fragment of
8-9 kb with a concomitant loss of fragments 14 and 5’ due
to a missing BamHI restriction site separating them. At
least one California isolate seemed to be closely related to
domestic pig isolates and may represent a transmission
from domestic to feral swine [59].
Nucleotide sequence variations are the basis for another
molecular approach for comparing PrV strains [60]. The
variable glycoprotein C (gC), a nonessential component of
the virion envelope involved in virus attachment to host
cells, was chosen for comparative studies. Sequence anal-
ysis of a region comprising parts of the gC gene allowed
differentiation of isolates exhibiting an identical RFLP
restriction pattern [45, 60], thus making it difficult to cor-
relate the two techniques [59].
However, sequence analysis was applied for the inves-
tigation of AD outbreaks in domestic pigs and for analysis
of PrV of wild swine in the USA [4, 59], the character-
ization of PrV from Brazil [61], and the characterization of
PrV associated with European wild boar [45]. PrV gC
sequences show a high degree of diversity (Fig. 1a and b).
At least eight different genetic types have been shown to
circulate in wild boar in Europe, with up to three circu-
lating in individual countries [45]. In the USA, PrV from
feral swine could be distinguished from virus circulating in
domestic pigs during the national epizootic [4]. In the
eastern states and the Hawaiian Islands, PrV gC sequences
showed a higher degree of identity than in Texas and
Oklahoma, where gC sequences were more variable
(Fig. 1a). Several PrV isolates from feral swine in the south
central states were closely related or identical to strains

123
Pseudorabies virus in wild swine
from domestic pigs, suggesting recent transmission from
escaped or released domestic pigs [4]. When PrV sequen-
ces of wild boar and feral swine origin from Europe and
America were analyzed together, a great diversity was
again revealed. Surprisingly, one German isolate (Ger-614)
was identical to sequences from Oklahoma, while vice
versa, the Texas isolate 19F was very closely related to
strain Bartha and one isolate from Slovakia (data not
shown). As the region used for sequence analysis repre-
sents only a very small portion of the entire PrV genome
(700 bp versus approx.145 kbp), these data should be
interpreted with care regarding epidemiological links and
virulence, and for reliable molecular epidemiology, more
than one parameter should be included.
Infection characteristics of PrV in populations of wild
swine
Results obtained in experimental studies on the suscepti-
bility of wild swine for PrV seem inconsistent and may
have been impaired by the origin and virulence of PrV
strains or the doses used for inoculation. Whereas no
clinical signs were observed after inoculation of wild boar
with a domestic swine isolate [62], moderate to severe
disease was produced in feral swine and wild boar,
respectively, that were experimentally infected with other
domestic PrV strains [63, 64]. Compared to domestic pigs,
very young wild swine seem more resistant to the virus,
probably because of the earlier maturation of the important
CD8 T cells [65], but in an outbreak of clinical AD in wild
boar in Spain, higher mortality was observed in young
individuals [11].
However, several studies and field observations have
shown, that infection with PrV of wild swine origin seems
to result in low pathogenicity, indicating that in wild swine,
the virus is relatively attenuated. Inoculation of feral swine
in the US with an indigenous feral swine PrV isolate
resulted in subclinical, latent infection [63]. Also, a wild
boar PrV isolate from East Germany produced mild clinical
symptoms in experimentally infected wild boar and
domestic pigs, but only when higher inoculation doses (107
TCID50) were used [64]. Since there is a diversity of PrV in
wild swine [4, 45], the results obtained with one strain may
not necessarily mimic the outcome of infection with other
variants. Field observations in Europe and the Americas
suggest that PrV infection in free-living wild swine may
produce only mild respiratory symptoms and may go
unnoticed [10]. However, occasional clinical cases with
central nervous symptoms have been reported from an
outbreak of AD in wild boar in south-central Spain and
from two cases in an endemically infected area in East
Germany [11, 39].
1699
This supports the hypothesis that PrV strains of wild
swine origin may generally be less virulent than those of
domestic pigs and are highly adapted to co-existence with
their specific host population [39, 63, 64]. As a conse-
quence of low virulence, seroconversion can be delayed,
and some wild swine can harbour viral DNA without
detectable antiviral antibodies [63, 64].
Most studies show that seroprevalence in wild swine is
correlated with age, and significantly higher infection rates
have been found in adult wild boar than in juveniles and
piglets [13, 35, 44, 54, 66–69]. A significant difference in
prevalence rates between the sexes was found among wild
boar in Spain and Tunisia, where seroprevalence was
higher in females than in males [31, 54].
The seroprevalence in wild boar piglets decreases over
the course of a year, and this may reflect the natural half-
life of maternal antibodies (maAb), which can be detected
up to 27 weeks postpartum by ELISA in consecutive
generations of offspring born to infected sows [69, 70].
Virus isolation from various secretions and excretions
of wild swine has suggested that oral/nasal [43, 44, 62, 67]
and venereal [40, 41, 43, 71] shedding represent the two
major routes of transmission of PrV among free-roaming
pig populations. Based on experimental transmission
studies with virus from feral pigs in both captive feral and
domestic swine, venereal transmission has been suggested
to be the likely main route of transmission in free-living
feral swine populations [41]. Transmission may be trig-
gered by sexual maturity, perhaps in conjunction with
reactivation of the virus due to mating stress, with males
playing a key role in transmission and maintenance of PrV
infection. Considering the age dependence of seropreva-
lence, there is reason to believe that the virus may be
transmitted predominantly in younger animals, as there are
higher percentages of susceptible animals among piglets
and juveniles. Because wild swine piglets born to naturally
infected sows seem to be protected from infection by
virus-neutralising maAbs up to 16 weeks postpartum, it
has been speculated that juveniles get infected in autumn,
when contact rates and stress increases due to the start of
the mating season [70]. The apparent high rate of pro-
miscuity observed in feral swine in Texas may increase
transmission rates [72]. In addition to direct contact, ani-
mals might also get infected when sniffing at tree trunks
marked by male wild boar in heat with virus-containing
saliva.
Risk of spillover to domestic pigs
Based on the available data (Tables 1 and 2), there is
reason to believe that PrV infections in wild swine pop-
ulations will spread further in the future. In the light of the
123
1700
tremendous progress made in elimination of AD in
domestic pigs on one hand and the vast territories with
endemic PrV infections in wild swine populations in North
America and Europe on the other hand, the potential risk
of PrV transmission via infected wild swine has become a
growing matter of concern. Although, there is evidence
that PrV infections in domestic pigs and in wild swine
represent epidemiologically distinct infection cycles [35,
66], experimental studies suggest that there is no zero risk,
as infected wild swine can shed virus in sufficient amounts
to trigger infection [43, 63]. So far, however, no quanti-
tative studies on the risk of transmission have been
reported.
With the possibility of spillover of virus to non-porcine
species, including rodents, transmission through carcasses
of rats, raccoons, and other infected animals should also be
considered [73].
Two main routes of transmission, aerosol transmission
and direct contact, to domestic pigs could be relevant.
Given the attenuated nature of wild swine PrV and the
rather low concentration of individuals in sounders com-
pared to domestic pig holdings, aerosol transmission of
PrV over long distances, without having direct contact to
domestic pigs, is unlikely [35]. Thus, in countries with a
high level of biosecurity in pig-producing facilities, the
risk of spillover infection appears much higher for
‘‘backyard’’ or outdoor producers of swine [74]. In Texas,
it has been demonstrated that feral swine have direct
contact with domestic swine from such facilities [75].
Also, in the USA, anecdotal reports have suggested
infection following the commingling of wild and domestic
swine for refreshment of gene pools. Supplementary
feeding during the hunting season as practised in some
countries may concentrate wild swine and thus facilitate
direct transmission among conspecifics and possibly to
domestic pigs in outdoor holdings.
During the final phase of eradication, sometimes the
only supporting evidence of new outbreaks in domestic
pigs is deduced from the absence of other recognised
sources of infection, and infection is therefore blamed on
feral swine in the vicinity [10]. In fact, in Europe in 2004
and 2010, isolated cases of AD in outdoor farms in
southwestern France were suggested to be linked to PrV
infections in wild boar [76]. In the Czech Republic, one
AD case in a backyard holding facility was attributed to
contact with wild boar, since the farmer was a hunter, and
swill feeding was identified as the possible route of
infection [77]. Nevertheless, those reports of PrV trans-
mission are surprisingly rare, considering that wild boar
have been identified as the primary source of initial out-
breaks of classical swine fever in Germany [78]. Some
credit can be given to the improvements in biosecurity
made by commercial farmers.
123
T. Müller et al.
Risk-mitigating measures
According to the Terrestrial Code of the World Organisa-
tion for Animal Health (OIE), the recognition and main-
tenance of an AD-free status is not affected by the presence
of PrV infections in wild swine, given that measures have
been implemented to prevent transmission of PrV from
wild swine to domestic pigs [79]. Unfortunately, those
measures have not been defined.
Disease elimination in wild swine as one possible option
is hampered by their natural behaviour and population
dynamics. In view of the continuously expanding wild
swine populations, with or without human intervention
[23], a reduction of wild swine populations below a density
where transmission would cease is illusory. Another option
would be the oral vaccination of wild swine against PrV,
which has proved successful for classical swine fever
(CSF) [80]. Recently, experimental studies have shown that
wild boar develop an adequate immune response after oral
vaccination with attenuated PrV (Lange, Vahlenkamp and
Mettenleiter, unpublished). While suitable attenuated vac-
cine strains may be at hand [81], a manageable and cost-
efficient vaccine baiting strategy still needs to be devel-
oped. In a large scale study of feral swine vaccinated with a
gene-deleted vaccine, for example, no change in population
seroprevalence of wild-type viral antibody was found
(David Stallknecht, personal communication). Also, safety
issues of attenuated PrV strains for non-target species
would be a major constraint unless recombinant vaccines
expressing immunodominant PrV glycoproteins would be
available in the future. Thus, oral vaccination of wild boar
is not an option at present.
The attenuated nature of wild swine PrV may lead to
inapparent infection, a delayed immune response, and a
lack of immune reactivity. Thus, infection of domestic pigs
could be silent, requiring continuous surveillance [10, 63,
64]. Yearly serological monitoring of the domestic pig
population is required for the maintenance of an AD-free
status [79, 82], and this would also help to detect spillover
infections of wild swine origin. The implementation of a
similar periodic serological monitoring of wild swine
populations at a national level could be another way to gain
basic information on the presence or absence of PrV
infections in wild swine. The National Wildlife Disease
Program within USDA-APHIS routinely screens feral pigs
for antibodies to CSF, PrV, and Brucella suis [83]. Given
the latent character of PrV infections, monitoring at bi- or
even triennial intervals would be sufficient to obtain
information on the spread and dynamics of PrV infections
in populations of wild swine, as experience from long-term
serological surveys (24 years) in wild boar populations in
East Germany has shown [35]. An adequate sample size
should be defined by taking into consideration statistical
Pseudorabies virus in wild swine
(accepted confidence intervals), spatial (size of the area
under scrutiny) [84] and practical (feasibility of sample
collection) aspects and detection of other diseases [8]. If
possible, clustered sampling should be avoided. Also,
outdoor pig farms deserve close attention in serological
monitoring for the maintenance of an AD-free status.
Assuming that venereal and oral/nasal excretion/infec-
tion are the main routes of transmission, prevention of
direct contact between wild swine and domestic pigs seems
an appropriate risk-mitigating measure and should be given
utmost priority. Separating domestic pigs and wild swine
by double electric fencing should be required for pig
holdings and at the same time, strict sanitation measures
should be applied. In particular, outdoor pig farms should
be considered to have the highest risk of infection [85, 86].
This approach has been successful in areas with high PrV
seroprevalence in wild boar in Germany. During the past
2‘ decades, no spillover of PrV has been detected in
domestic pigs in those areas [35]. In Spain, the level of PrV
seroprevalence in domestic pig farms is not influenced by
the level of ADV seroprevalence in the coexisting wild
boar populations [86]. Free roaming of domestic pigs is a
traditional form of animal husbandry in some parts of
Europe, in particular in the Mediterranean basin and the
Balkans [23], and may be required for the production of
pork specialities, such as ham. Curtailing or abandoning
those traditions will be difficult to achieve. Under these
circumstances, PrV in wild swine may be considered a
constant threat, and continual vaccination of traditionally
outdoor-farmed domestic pigs with gE-deleted vaccines
and consideration of all of the consequences regarding
animal trade may be necessary to prevent disease in those
animals.
Given these limitations, the only recommended strategy
for the control of PrV in wild boar is adequate monitoring
for infection. This will allow the true risk to be estimated
and may help to stratify mitigating measures. In the future,
the new animal health law of the European Union implies
that member states should monitor the situation and take
measures to reduce and, as far as possible, to prevent the
spread of certain diseases in wild animals (Source EN,
SANCO/7221/2010, Working document: Draft proposal
for the EU Animal Health Law).
Conclusions
Despite successful elimination of PrV from domestic pigs
in several parts of the world, this virus seems to be cir-
culating globally in populations of non-domestic swine.
There has been a notable increase in PrV seroprevalence
for the past two decades that might have been favoured by
the dramatic increase of wild swine populations. The
1701
historic origin of PrV infections in domestic pigs as well
as in wild swine remains unknown. The genetic similarity
among wild swine and domestic pig isolates suggest that
transmission has taken place in both directions. Since PrV
infections in domestic pigs and wild swine appear to be
epidemiologically distinct, PrV in wild boar does not
impact on the OIE definition ‘‘freedom from disease’’ in
domestic pigs. However, infected populations of wild
swine represent a constant danger for reintroduction of
PrV into free domestic pig herds and regions, as observed
with CSF virus. Implementation of proper monitoring of
disease in wild swine populations should be mandatory,
particularly in areas where wild swine populations overlap
with domestic pig husbandry. Also, further studies are
needed that focus on factors that affect the risk of wild-
boar-associated PrV transmission, as well as character-
ization of isolates and successful measures to eliminate the
disease.
Acknowledgments The authors would like to acknowledge the
anonymous reviewers for their valuable comments and suggestions,
which helped to improve the manuscript. We also thank Viola
Damrau from the library at FLI for her continuous support. Gratefully
acknowledged is the financial support by the Federal Ministry of
Food, Agriculture and Consumer Protection.
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