REAL-PhD

Neumanné Virág, Ildikó

Az integráció hatása az EU tagországok külkereskedelmére – vizsgálatok gravitációs modellel = Impacts of the integration on trade of EU members - The impact of the integration on the trade of EU members-approach with gravity model

Somfay, Örs

Az I. világháború magyar vonatkozású köztéri, valamint közösségi hősi emlékei és ezek adatbázisa

Albertné Zombori, Katalin

Forradalom és megtorlás a Bács-Kiskun megyében lezajlott tanár- és diákperek tükrében (1956-58)

Nemes, Gábor

Az Apostoli Szentszék és a Jagelló-kori Magyarország kapcsolata - különösen VII. Kelemen pápa idején (1523–1526)

Koller, Miklós

Spatial–Temporal Event Recognition and Metastable Oscillations in CNN Wave Computer

Virág, András

Hálózatok a cserkészetben - cserkészhálózatok avagy a magyarországi cserkészet titkos története, különös tekintettel az 1939-től 1990-ig terjedő időszakra

Csáki, András

Andrés Segovia művészete 21.századi szemmel

Solymosi Tari, Emőke

Lajtha László színpadi művei: Egy ismeretlen műcsoport a szerzői életút kontextusában

Mátrai, Norbert

Genetikai és madárgyűrűzési vizsgálatok hazai költő és afrikai telelő nádirigó (Acrocephalus arundinaceus) populációkon

Vas, Zoltán

Szent István University 
Postgraduate School of Veterinary 
Science 
 
 
Host-parasite relationship of birds (Aves) 
and lice (Phthiraptera) – evolution, 
ecology and faunistics 
 
PhD theses 
 
Zoltán Vas 
 
 
2013 
2 
 
Supervisors and consultants: 
 
Prof. Dr. Jenő Reiczigel 
Szent István University, Faculty of Veterinary Science, 
Department of Biomathematics and Informatics 
supervisor 
 
Dr. Lajos Rózsa 
MTA-ELTE-MTM Ecology Research Group 
supervisor 
 
Dr. Andrea Harnos  
Szent István University, Faculty of Veterinary Science, 
Department of Biomathematics and Informatics 
consultant 
 
Dr. Péter László Pap  
Babes-Bolyai University, Faculty of Biology and Geology; 
University of Debrecen, Behavioural Ecology Research Group 
consultant 
 
 
 
 
3 
 
Introduction  
The host-parasite relationship is one of the most complex and 
intimate associations in nature. Given the astonishing diversity 
of parasitic animals, as well as that of their hosts, a huge 
variety of interactions evolved between the two partners. 
These interactions can be studied from several different points 
of view, each contributing to the general understanding of 
host-parasite relationships. At an ecological level, this 
relationship is often characterized as an arms race – 
emphasizing the conflicting adaptations and counter-
adaptations of the partners. Nevertheless, often the present 
partners’ ancestors have already been associated through 
evolutionary ages, raising the possibility to study how host and 
parasite lineages have affected each other’s adaptation and 
diversification. Such studies are able to draw reliable inference 
only if based on a sound classical zoological background: 
faunistics, taxonomy, and systematics. 
In this PhD thesis I present a multidisciplinary approach 
investigating the relationship of parasitic lice (Insecta: 
Phthiraptera) and their avian and mammalian hosts 
(Vertebrata: Aves, Mammalia). I apply both modern statistical 
methodologies of evolutionary comparative analysis and 
classical zoological methodologies such as sampling in the 
field for faunistical purposes. The two main chapters of the 
thesis indicate the two fields I was most concerned in the 
recent years: (i) the linkage between host and parasite 
diversity at a macroevolutionary level and (ii) various aspects 
of louse faunistics. 
Louse richness is remarkably variable among host taxa. The 
features of the bird-louse relationship offer a unique system to 
study the inference of macroevolutionary and macroecological 
4 
 
factors shaping two closely associated lineages. In this 
chapter I focus on the diversity component of host-parasite 
relationships, using birds (and sometimes mammals) and their 
lice as a model system. Parasites and other pathogens have a 
quite specific role in the formation and maintenance of 
biodiversity; they not only represent themselves a 
considerably large proportion of biodiversity but  they have a 
unique importance in facilitating the diversity of their hosts as 
well, hence contributing to the global biodiversity as a whole at 
a great extent. 
Local faunistical surveys are essential to discover the patterns 
in the geographical distribution of lice, the local variability in 
louse-host associations and in parasite assemblages. 
Although faunistics is often considered to be a low-prestige 
field of biology nowadays, I argue that it is – and will remain to 
be – vital to provide basic data about the studied organisms 
and thus to allow further evolutionary and ecological 
investigations. 
 
Aims  
Here I briefly list the main questions and aims of my research. 
Chapter 1 – Louse diversity and its macroevolutionary 
correlates 
I investigated: 
 whether there is a co-variation between avian 
cognitive/innovative capabilities and the taxonomic 
richness of their lice; 
5 
 
 whether louse diversity is reduced on brood parasitic 
bird clades due to the lack of vertical transmission 
routes (i.e. from parents to chicks) for lice; 
 whether more diverse host groups harbor more 
diverse parasites by testing Eichler’s assumption (the 
so-called Eichler’s rule) for the first time. 
Chapter 2 – Louse faunistics and conservation biology 
I aimed to: 
 prepare the Hungarian louse checklist by critically 
reviewing the literature; 
 find new louse species for the Hungarian fauna by 
identifying the ectoparasite samples collected in the 
last few years by myself and my colleagues; 
 test whether bird ringing procedure reduces avian 
louse burdens, thus causing a bias in louse 
infestation measures of birds sampled in bird ringing 
centers; 
 prepare a global list of critically co-endangered and 
co-extinct louse species to draw attention to the 
conservational status and value of parasitic animals.  
 
6 
 
Results and discussion 
Chapter 1 – Louse diversity and its macroevolutionary 
correlates 
The study entitled “Clever birds are lousy: Co-variation 
between avian innovation and the taxonomic richness of their 
amblyceran lice” unveils a formerly unexpected positive 
correlation between the host cognitive/innovative capabilities 
and louse richness across avian families. Using data of 108 
avian families (controlling for phylogenetic effects, host 
species richness within a family, body size, and research 
effort) I found that host cognitive/innovative capabilities and 
relative brain size co-vary positively with Amblyceran genera 
richness, but not with Ischnocerans. I propose several 
alternative and mutually non-exclusive hypotheses to explain 
this phenomenon. 
The study entitled “Avian brood parasitism and ectoparasite 
richness – scale-dependent diversity interactions in a three-
level host-parasite system” explores the effect of hosts’ brood 
parasitic life-style and the complex metapopulation structure of 
foster-generalist cuckoo species on their louse richness. 
Brood parasitic birds, together with their parasites and their 
foster birds constitute a complex three-level evolutionary 
system. Brood parasitic birds harbor host-specific louse 
species despite the complete lack of the vertical route of louse 
transmission, as their nestlings never get into direct physical 
contact with their genetic parents. I showed that host clades’ 
past switches to brood parasitism reduced both Amblyceran 
and (to a lesser extent) Ischnoceran genera richness, most 
probably because several louse lineages were unable to adapt 
to the lack of vertical transmission. On the other hand, I also 
7 
 
showed that the supposedly more complex and dynamic 
subpopulation structure of foster-generalist (i.e. utilizes several 
to many foster species) cuckoo species facilitates Ischnoceran 
species richness; hence for the first time diversity interactions 
across a three-level host-parasite system were recognized. 
The study entitled “Evolutionary co-variation of host and 
parasite diversity – the first test of Eichler’s rule using parasitic 
lice (Insecta: Phthiraptera)” re-visits and tests with modern 
methodologies the more or less forgotten “Eichler’s rule”. 
Eichler’s original assumption that more diverse host groups 
harbor more diverse parasite faunae was strongly supported 
by my study. I found positive co-variation between the species 
richness of 175 avian and 147 mammalian families and the 
genera richness of their Amblyceran, Ischnoceran, and 
Anopluran lice. Host diversity proved to be the strongest 
predictor of parasite diversity found so far. The potential 
macroevolutionary and macroecological mechanisms beyond 
this phenomenon were also discussed. 
Chapter 2 – Louse faunistics and conservation biology 
I compiled a checklist of avian and mammalian lice occurring 
in Hungary that includes 279 louse species (and subspecies) 
which have been recorded since 1945. Their hosts represent 
156 bird and 30 mammal species. Additionally, I also listed 
further 550 louse species whose occurrence may seem to be 
likely (as judged from their geographic and host distribution) 
but have not been detected yet. 
I identified ectoparasite samples collected by myself and my 
colleagues in recent years. This study increases the number of 
louse species and subspecies known from Hungary to 299, 
8 
 
infesting 170 bird species (14 new avian host species as 
compared to the checklist mentioned above). 
Ectoparasite sampling of wild birds is usually carried out 
parallel with bird ringing activities all over the World. Bird 
ringing stations work with the appropriate licenses issued by 
the relevant conservation authorities and offer excellent 
conditions for louse sampling as a huge number of birds 
belonging to several different species are caught there in 
standardized ways. However, the handling of host individuals 
during the ringing procedure may lead to ectoparasite loss. To 
test this idea Barn Swallows (Hirundo rustica) were assigned 
in the breeding season into two experimental groups – the 
birds were subjected to either a standard (recording biometry, 
fat and other condition scores, feather hole counts), or a 
reduced ringing procedure (only feather hole counts). I used 
feather holes (presumed traces of louse chewing) as a 
measure of louse loads. The birds were recaptured and holes 
were recounted after a month. Significantly more new holes 
appeared in the reduced ringing procedure group indicating 
that ringing procedures effectively reduces louse loads. 
The global conservational status of parasitic lice was 
assessed by combining the known conservational status of the 
avian and mammalian hosts with the published information on 
the host-specificity of their lice. The list contains 6 co-extinct, 
and 40 (possibly 41) critically co-endangered louse species, 
and further 2 (but possibly 4) cases of conservation-induced 
extinction. The purpose of this parasite “red lists” is to draw 
attention to the endangerment of dependent taxa – such as 
symbionts, parasites and parasitoids – and their biological 
value. 
9 
 
Summary 
Here I briefly list the main results: 
Chapter 1 – Louse diversity and its macroevolutionary 
correlates 
 a positive co-variation was detected between avian 
cognitive/innovative capabilities and the generic 
richness of Amblyceran lice; 
 avian hosts’ switching to brood parasitic way of life 
was shown to reduce the genera richness and 
taxonomic distinctness index of both Amblyceran and 
(to a lesser extent) Ischnoceran lice; 
 a positive evolutionary co-variation was found between 
the diversity measures (species richness, genera 
richness, and taxonomic distinctness) of cuckoos’ 
Ischnoceran lice and the number of their foster 
species; to my best knowledge, this represents the 
first diversity interaction ever found in a three-level 
host-parasite system; 
 Eichler’s assumption that more diverse host groups 
harbor more diverse parasites was revisited, tested, 
and evaluated by modern statistical methodologies 
covering the global louse fauna; this co-variation 
represents the strongest predictor of louse diversity 
recognized so far. 
 
10 
 
Chapter 2 – Louse faunistics and conservation biology 
 the Hungarian louse checklist was compiled by 
critically reviewing the literature; 
 20 louse species were found as new to the Hungarian 
fauna by identifying the ectoparasite samples 
collected in the last few years; 
 bird ringing procedure was shown to reduce avian 
louse burdens; 
 a global list of critically co-endangered and co-extinct 
louse species was provided, with particular emphasis 
on louse species which became extinct due to 
conservation efforts focusing on the host. 
11 
 
List of publications  
(in peer-reviewed journals or as book chapters between 
2009-2013, related to the present thesis) 
Vas Z., Fuisz T.I., Fehérvári P., Reiczigel J., Rózsa L. 2013. 
Avian brood parasitism and ectoparasite richness – 
scale-dependent diversity interactions in a three-level 
host-parasite system. Evolution 67(4): 959-968. IF: 
4.864 
Rózsa L., Vas Z. 2013. A list of co-extinct and critically co-
endangered species of parasitic lice (Phthiraptera) 
with remarkable cases of conservation-induced 
extinction. Oryx, in press. IF: 1.624 
Rózsa L., Vas Z. 2013. Host correlates of diversification in 
avian lice. In: Morand S., Krasnov B.R., Littlewood T. 
(eds.) Parasite diversity and diversification: 
evolutionary ecology meets phylogenetics. Cambridge 
Univerity Press, in press. 
Vas Z., Csorba G., Rózsa L. 2012. Evolutionary co-variation of 
host and parasite diversity – the first test of Eichler’s 
rule using parasitic lice (Insecta: Phthiraptera). 
Parasitology Research 111: 393-401. IF: 2.852; 
independent citations: 2. 
Vas Z., Privigyei Cs., Prohászka V.J., Csörgő T., Rózsa L. 
2012. New species and host association records for 
the Hungarian avian louse fauna (Insecta: 
Phthiraptera). Ornis Hungarica 20(1): 44-49. 
12 
 
Vas Z., Rékási J., Rózsa L. 2012. A checklist of lice of 
Hungary (Insecta: Phthiraptera). Annales historico-
naturales Musei nationalis hungarici 104: 5-109. 
Vas Z., Lefebvre L., Johnson K.P., Reiczigel J., Rózsa L. 
2011. Clever birds are lousy: Co-variation between 
avian innovation and the taxonomic richness of their 
amblyceran lice. International Journal for Parasitology 
41: 1295-1300. IF: 3.393; independent citations: 4. 
Vas Z., Fuisz T.I. 2010. Ringing procedure can reduce the 
burden of feather lice in Barn Swallows Hirundo 
rustica. Acta Ornithologica 45(2): 203-207. IF: 0.889; 
independent citations: 2. 
Vas Z., Fuisz T.I. 2010. Rendhagyó vedlésű füsti fecske 
(Hirundo rustica). Aquila 116-117: 132-133. 


Madarak (Aves) és tolltetveik (Phthiraptera) gazda-parazita kapcsolatának evolúciós, ökológiai és faunisztikai vizsgálata

Not a member yet