Social Insect

Social immunity, which describes how individual behaviors of group members effectively reduce disease and parasite transmission at the colony level, is an emerging field in social insect biology. An understudied, but significant behavioral disease resistance mechanism in honey bees is their collection and use of plant resins. Honey bees harvest resins with antimicrobial properties from various plant species and bring them back to the colony where they are then mixed with varying amounts of wax and utilized as propolis. Propolis is an apicultural term for the resins when used by bees within a hive. While numerous studies have investigated the chemical components of propolis that could be used to treat human diseases, there is a lack of information on the importance of propolis in regards to bee health. This review serves to provide a compilation of recent research concerning the behavior of bees in relation to resins and propolis, focusing more on the bees themselves and the potential evolutionary benefits of resin collection. Future research goals are also established in order to create a new focus within the literature on the natural history of resin use among the social insects and role that propolis plays in disease resistance. Die „soziale Immunität“ als neues Forschungsfeld bei sozialen Insekten beschreibt, wie das individuelle Verhalten von Mitgliedern einer Gruppe wirkungsvoll die Verbreitung von Krankheiten und Parasiten auf der Ebene des Sozialstaates verhindern kann. Ein bisher zwar wenig untersuchtes aber wichtiges Verhaltensmerkmal zur Krankheitsabwehr bei Honigbienen ist die Verwendung von Pflanzenharzen. Honigbienen sammeln Harze mit antimikrobiellen Eigenschaften von verschiedenen Pflanzen, mischen diese dann im Bienenvolk mit unterschiedlichen Mengen von Wachs und benutzen dies als Propolis (Abb. 1–4). Propolis ist demnach der bienenkundliche Begriff für Harze, die im Bienenstock verwendet werden. Während es zahlreiche Untersuchungen zur Verwendung bestimmter Bestandteile des Propolis zur Krankheitsbekämpfung beim Menschen gibt, sind kaum Informationen über die Bedeutung von Propolis für die Bienengesundheit vorhanden. Dieses Review ist eine Zusammenstellung neuerer Forschungsergebnisse zum Verhalten der Bienen in Bezug auf Harze und Propolis mit dem Schwerpunkt auf die möglichen evolutiven Vorteile des Harzsammelns für die Honigbienen. Die Verwendung von Harzen durch Bienenvölker (Apis mellifera) ist weit verbreitet. Während es erhebliche Unterschiede zwischen einzelnen Völkern bzgl. der Menge an gesammelten Harzen und Propolis gibt, scheinen alle — und dabei insbesondere die wildlebenden — Bienenvölker das Propolis zur Auskleidung des gesamten Stockinneren zu benutzen. Es wird angenommen, dass Propolis dazu beiträgt, die Homöostase innerhalb des Bienenstockes aufrecht zu erhalten. Konkret könnte das Propolis dabei das mikrobielle Wachstum an den Beutenwänden reduzieren, unkontrollierten Luftzug ins Beuteninnere verhindern und zusätzlich mechanische Barrieren gegenüber Eindringlingen bilden. Einige Forschungsprojekte zeigen eindeutig, dass Propolis im Bienenstock direkt gegenüber Krankheitserregern (z.B. Amerikanische Faulbrut) und Parasiten (z.B. Kleiner Beutenkäfer, Varroa destructor) wirkt. Daneben scheint es aber auch eine subtilere Wirkung über die Unterstützung des individuellen Immunsystems zu geben. Die weiteren Forschungen sollten sich auf das bessere Verständnis der Verwendung von Harzen durch Honigbienen und andere soziale Insekten konzentrieren. Dafür gibt es eine Vielzahl an Forschungsfeldern, von den pharmazeutischen Möglichkeiten des Propolis für die menschliche Gesundheit über die Mechanismen der Sammelstrategie von Propolis auf den Ebenen der Einzelbienen und des Bienenvolkes bis hin zu einer möglichen Anwendung von Propolis als Bekämpfung von Bienenkrankheiten. Schließlich ermöglichen Informationen zur Verwendung von Harzen und deren Aufnahme in den Bienenstock spannende Forschungsansätze zum Einfluss der Umwelt auf Krankheitsresistenz und soziale Immunität.

Bibliographies are lists which convey idenfication information of different publications like books,
papers and articles on specific subjects in a certain order. They are prepared with the intention to provide
those who conduct studies in a certain field with the related source material in its entirety. They make it
possible to quickly review previous work on the subject being studied. They also provide an opportunity to
make more realistic, effective and less time consuming plans in reaching the target.
The first comprehensive bibliography on entomology and agricultural zoology in Turkey was
prepared by Prof. Dr. Feyzi Önder, Prof. Dr. Yusuf Karsavuran, Dr. E. Pervin Önder, Dr. Aynur Önuçar and
Dr. Ertaç Tutkun within the study framework of Turkish Entomology Association. It was printed in 1982 by the
General Directorate of Plant Protection and Quarantine under the name of “Bibliography of Entomology
and Agricultural Zoology of Turkey (Fauna, Pest Control, Pests of Plants, Methods) (1595-1978)”. The
second bibliography in this field was prepared by Prof. Dr. Feyzi Önder, Prof. Dr. Yusuf Karsavuran, Dr. E.
Pervin Önder, Dr. Ertaç Tutkun and Prof. Dr. Serdar Tezcan and was published by TURDOK, Tübitak-
Turkish Scientific and Technical Documentation Center in 1986 under the name of “Bibliography of
Entomology and Agricultural Zoology of Turkey (1979-1983)”. Bibliographical information of 4416
publications are available in the first of these bibliographies and there are 1361 of them in the second
bibliography. Bibliographical information of a total of 5777 publications are classified by topics and can be
found in these two volumes.
In the 30 years since 1983 there are numerous new research institutes and universities opened in
Turkey. As a consequence, new researchers were bred and have conducted and finished projects in the
fields of entomology and agricultural zoology. Additionally, technical developments in printing techniques
resulted with many new journals and periodicals. And, also, the number and frequency of scientific meetings
increased. All these have played a role in the considerably increased number of publications. Advancements
in the areas of communication and information technologies had marked influence on, especially, the second
half of the last 30-year period. They effected the production speed of information as well as the speed of
reaching this information. While some of the produced information was instantaneously attainable, some was
harder to be reached due to the huge amount of information output.
In the preparation phase of the book called “650 Entomologists, Acarologists and Nematologists from
Turkey with Their Brief Life Histories” written by Prof. Dr. Serdar Tezcan and Dr. Füsun Tezcan which was
published by the Entomological Society of Turkey, scientists who conducted studies in these topics were
contacted. Along with their biographical information, information concerning their publications as of
December 31, 2011 was also tried to be gathered.
Within the scope of the studies which lasted approximately seven months, authors were requested to
send in their work from their publication lists which were in line with the concept of this publication. Those
which were present in previous bibliographies were sorted out and the remaining which belonged to the 28-
year period between 1984-2011 were compiled. They were standardized to the best that it could and have
been listed alphabetically by the author last names. Those publications which belonged to a period before
1984 but not found in previous bibliographies were also included in this compilation. It should be noted that
X
this study which features 10823 publications does not contain all literature published between 1984-2011.
Some works of our colleagues could not be included in this study because they did not tend to share their
publication lists despite they were accessed. Furthermore, the publications which belonged to foreign
researchers could only be included if they were co-produced with a local colleague. Unfortunately, their other
studies could not be included. It should also be noted that, differing from previous ones, the entries in this
book are not classified according to their topics.
The contents of this study is subject to become out of date quickly due to the rapid output rate of
information today. For this reason, this book has been printed in a limited number and has been distributed
to libraries and documentation centers. In order to facilitate and spread the utilization of the book, an e-book
format of it is posted in the electronic library in the website of the Entomological Society of Turkey
(http://www.entomoloji.org.tr/). This will make it available free of charge for those local and foreign
people who wish to make use of it. The e-book can be scanned or bibliographical information can be
extracted by writing either the author of the publication, title or name of the journal after clicking on the
related file.
The pdf files of the two volumes of Bibliography of Entomology and Agricultural Zoology of
Turkey, within the scope of a graduation thesis written in 2013 in the Department of Plant Protection of Ege
University, Faculty of Agriculture, has been posted in the abovementioned website with the intention to
facilitate the scanning of publications prior to 1984. We would like to point out that this will enable the user to
make use of the old publications by using the same method.
It is technologically easy to access information today; however, it may be harder to find it in a short
period of time due to the increased speed of output of information. We are wishing that this bibliographical
reference we have prepared will further contribute to the production of new knowledge today. We are
thinking that the target audience who would make use of this reference book is comprised of researchers in
the fields of biology, agriculture, forestry, veterinary, medicine, environmental sciences and standing on the
top of the list are our young colleagues who will study in the fields of entomology, acarology and nematology.
We would like to extend our gratitutes to the President of the Entomological Society of Turkey, Prof.
Dr. Enver Durmuşoğlu and to the members of the Board of Directors for their support by assuming the
printing of this study and posting its e-book form in their website which will make it possible to reach the
target audience. We would like to thank to all our colleagues, dear families and colleagues of our deceased
colleagues who have made this book possible by sending in information. We also would like to extend our
cordial thanks to Mr. Olcay Sütüven and his valued colleagues for their dedicated and meticulous work in
materializing the printing of this book.
March 2013, Bornova
Serdar TEZCAN-Füsun TEZCAN-Nilay GÜLPERÇİN

Honey bee (Apis mellifera L.) workers are essentially sterile females that are used to study how complex social behavior develops. Workers perform nest tasks, like nursing larvae, prior to field tasks, like foraging. Despite worker sterility, this behavioral progression correlates with ovary size: workers with larger ovaries (many ovary filaments) start foraging at younger ages on average. It is untested, however, whether the correlation confers a causal relationship between ovary size and behavioral development. Here, we successfully grafted supernumerary ovaries into worker bees to produce an artificial increase in the amount of ovary tissue. We next measured fat body mRNA levels for the yolk precursor gene vitellogenin, which influences honey bee behavioral development and can correlate with ovary size. Vitellogenin was equally expressed in surgical controls and bees with supernumerary ovaries, leading us to predict that these groups would be characterized by equal behavior. Cont...

1Chemical communication plays an important part in the lives of insects, and particularly in lives of those that live in groups or social organizations.2Chemicals which are used in communication in the general sense are called semiochemicals, and there are a number of subdivisions recognized under this title.3Pheromones are a category of semiochemicals which are used for communication between individuals of the same species.4Pheromones are in turn subdivided into primer and releaser pheromones. The former produce a relatively long-lasting physiological change in the receiver, and the latter stimulate the receiver to some immediate behavioural response.5Far more is known about releaser pheromones at present because they are easier to study.6Nine categories of releaser pheromone are recognized here, used by both social and non-social insects.7Sex pheromones are widely used to bring the sexes together for mating, and they have been extensively studied in Lepidoptera.8Invitation pheromones, encouraging the species to feed or oviposit at an explored site, are not extensively known.9Aggregation pheromones are designed to bring individuals together into groups which may be temporary in sub-social insects, or permanent in social insects.10Dispersal or spacing pheromones are used by other species to reduce intraspecific competition for scarce resources.11Alarm pheromones are a broad and sometimes unclearly defined group which communicate alarm or attack, chiefly in colonial species.12Trail pheromones, applied to a surface by an individual, to be followed by another, are confined to Hymenoptera, Isoptera and a few Lepidoptera as far as is known.13Territorial and home range pheromones may be widely distributed, but as yet few of them have been recognized.14Surface and funeral pheromones are even less well known. Surface pheromones may play a large part in species or colony recognition.15We can expect the number and complexity of pheromones to be much greater in social insects, a part of the subject which until now has received relatively less attention.16As our understanding of the subject grows we may expect other categories to be added to this list.Chemical communication plays an important part in the lives of insects, and particularly in lives of those that live in groups or social organizations.Chemicals which are used in communication in the general sense are called semiochemicals, and there are a number of subdivisions recognized under this title.Pheromones are a category of semiochemicals which are used for communication between individuals of the same species.Pheromones are in turn subdivided into primer and releaser pheromones. The former produce a relatively long-lasting physiological change in the receiver, and the latter stimulate the receiver to some immediate behavioural response.Far more is known about releaser pheromones at present because they are easier to study.Nine categories of releaser pheromone are recognized here, used by both social and non-social insects.Sex pheromones are widely used to bring the sexes together for mating, and they have been extensively studied in Lepidoptera.Invitation pheromones, encouraging the species to feed or oviposit at an explored site, are not extensively known.Aggregation pheromones are designed to bring individuals together into groups which may be temporary in sub-social insects, or permanent in social insects.Dispersal or spacing pheromones are used by other species to reduce intraspecific competition for scarce resources.Alarm pheromones are a broad and sometimes unclearly defined group which communicate alarm or attack, chiefly in colonial species.Trail pheromones, applied to a surface by an individual, to be followed by another, are confined to Hymenoptera, Isoptera and a few Lepidoptera as far as is known.Territorial and home range pheromones may be widely distributed, but as yet few of them have been recognized.Surface and funeral pheromones are even less well known. Surface pheromones may play a large part in species or colony recognition.We can expect the number and complexity of pheromones to be much greater in social insects, a part of the subject which until now has received relatively less attention.As our understanding of the subject grows we may expect other categories to be added to this list.

■ Abstract Reproductive-skew theory can be broadly divided into transactional models, in which reproduction is shared among group members in return for some fitness benefit, and tug-of-war models, in which reproductive sharing arises solely from an inability of each group member to fully control the others. For small-colony social insects in which complete reproductive control by a single individual is

A variety of social insects use visual cues for homing. In this study, we examine the possible factors affecting the learning and retention of nest-associated visual cues by the Australian desert ant Melophorus bagoti and the manner in which such cues are encoded by foraging ants. We placed four prominent cylindrical landmarks around a nest and trained foragers from that nest to a food source. Ants were tested with the landmark array in a distant testing field after (1) a known number of exposures to the landmarks (1, 3, 7 or 15 trials, spread over a period of 1 day, 2 days or ≥3 days) and (2) after a known period of delay (0, 24, 48, 96 or 192 h). The results show that a combination of an increase in training trials and an increase in number of training days affected the acquisition of landmark memory. Moreover, once the landmarks were learnt, they became a part of long-term memory and lasted throughout the ants’ foraging lifetime. To examine visual cue encoding behaviour, ants trained under similar conditions for 4 days were tested with (1) an identical landmark array, (2) landmarks of the same size used in training, but placed at twice the distance from each other, and (3) landmarks whose dimensions were doubled and placed at twice the distance from each other. In conditions (1) and (3), the ants searched extensively at the centre of the four landmarks, suggesting that, similar to the Saharan ant (genus Cataglyphis) and the honeybee, M. bagoti too uses a snapshot to match the view of the landmarks around the nest. But contrary to the snapshot model, in condition (2), the ants did not search extensively at the centre of the landmarks, but searched primarily 0.5 m from the landmark, the distance from each landmark to the nest during training. We discuss how various search models fare in accounting for these findings.