ТИПОВОЙ ВИД

СИНОНИМИЯ

ОСНОВНЫЕ СИНОНИМЫ

1. Tomognathus Mayr, G. 1861: 56. Типовой вид: Myrmica sublaevis Nylander, по монотипии. Поскольку это название оказалось младшим гомонимом рода рыб Tomognathos Agassiz, 1850 (Pisces), то А.Форель предложил замещающее название Harpagoxenus Forel, 1893.
2. Harpagoxenus Forel, 1893: 167. Типовой вид: Myrmica sublaevis Nylander, по монотипии. Замещающее название для младшего гомонима Tomognathus Mayr, 1861.
   
   
   

СИСТЕМАТИ-ЧЕСКОЕ ПОЛОЖЕНИЕ

БИОЛОГИЯ

• Муравьи-рабовладельцы мелкого размера (3-5 мм), использующие в качестве рабов виды рода Leptothorax. Новая семья основывается молодой самкой, которая проникает в гнездо хозяина и убивает там местную матку.
• Рабами служат как правило виды Leptothorax acervorum и Leptothorax muscorum, реже Leptothorax tuberum.
• Гнезда в почве.
• Семьи немногочисленные, все работы выполняют только рабочие вида-раба. Рабочие самого вида-рабовладелца только совершают периодические набеги для захвата новых куколок рабов.

МОРФОЛОГИЯ

• Голова большая, почти прямоугольная с широкой затылочной выемкой, с широкими острыми жвалами без зубцов на жевательном крае.
• Лобные валики почти прямые и длинные. Наличник короткий, широкий.
• Глаза маленькие, слабоовальные, глазки отсутствуют.
• Усики 11-члениковые с 4-члениковой булавой (у самцов усики 12-члениковые).
• Формула щупиков: 5,3.
• Тело коренастое с коротким и мощным стебельком.
• Окраска рабочих желтовато-коричневая, брюшко темно-коричневое.
• Скульптура: голова и брюшко блестящие, грудь и стебелек слабоблестящие. Голова в продольных морщинках, остальное тело - в зернистой скульптуре.
• Тело покрыто длинными, густыми, серебристыми отстоящими волосками.
• Проподеум с 2 широкими зубцами.
• Постпетиолюс снизу с острым зубцом, направленным вниз и вперед. Петиолюс без цилиндрической части.
• Брюшко короткое с 2-3 видимыми сегментами.
• Передние крылья с 1 радиомедиальной и 1 дискоидальной ячейками.
• Встречаются как нормальные, так и эргатоидные самки.
• Размеры, как правило, мелкие, 3-5 мм.

РАСПРОСТРАНЕНИЕ

• Это очень маленький по числу видов род муравьев. На сегодня род Harpagoxenus включает 3 вида (список смотрите ниже), распространенных только в Палеарктике и Неарктике (Канада, США).
• В Неарктике (Мексика, юг США) известен 1 вид - Harpagoxenus canadensis Smith, 1939.
  

ФАУНА МИРА

ХРОМОСОМЫ

ЛИТЕРАТУРА ПО ФАУНЕ, ЭКОЛОГИИ И СИСТЕМАТИКЕ

Здесь собраны из базы Formis-2003 основные работы (род Harpagoxenus упоминается в 217 работах и в заголовках 57 из них), включая 5 диссертаций: A.Bourke (1987) и др.

1. Allies, A. B., A. F. G. Bourke, et Franks, N.R. (1986). "Propaganda substances in the cuckoo ant Leptothorax kutteri and the slave-maker Harpagoxenus sublaevis." J. Chem. Ecol. 12: 1285-1293.
   Впервые обнаружено наличие 'propaganda substances' у муравьев-инквилинов, у которых нет рабочей касты - у Европейского муравья L. kutteri Buschinger. Это вещество используются паразитической самкой в качестве оружия для защиты от рабочих вида-хозяина L. acervorum. Вещество это имеет необычный поведенческий эффект: оно заставляет рабочих вида-хозяина нападать друг на друга, и происходит аннулирование гнездового распознания соплеменников. Лабораторные эксперименты показали, что это вещество вырабатывается в Дюфуровой железе самок L. kutteri. Сходный эффект ранее был обнаружен у рабовладельцев H. sublaevis, которые используют секрецию Дюфуровой железы при рейдах и при основании новых колоний.
   
   
2. Alloway, T. M. (1979). "Raiding behaviour of two species of slave-making ants, Harpagoxenus americanus (Emery) and Leptothorax duloticus Wesson (Hymenoptera: Formicidae)." Anim. Behav. 27: 202-210.
   Сравнивается рабовладельческое поведение видов H. americanus и L. duloticus. In both species, raids follow 'scouting' by individuals or small groups of slave-makers. Single scouts return to their nest to lead a raiding party. Groups attack host-species nests at once, with or without recruiting additional raiders. Raiders usually guard the entrance to the raided nest, preventing workers from escaping with brood. L. duloticus raiders kill most adults in raided nests, but H. americanus raiders kill very few such individuals. H. americanus raiders always and L. duloticus usually transport the raided colony's brood back to their own nest. However, L. duloticus colonies sometimes emigrate into raided nests.
   
   
3. Alloway, T. M. (1982). How the slave-making ant Harpagoxenus americanus (Emery) affects the pupa-acceptance behavior of its slaves. The Biology of social insects : Proceedings of the Ninth Congress of the International Union for the Study of Social Insects. M. D. Breed, C. D. Michener and H. E. Evans. Boulder, Colorado, Westview Press: 261-265.
   
   
4. Alloway, T. M. and M. G. Del Rio Pesado (1983). "Behavior of the slave-making ant, Harpagoxenus americanus (Emery), and its host species under 'seminatural' laboratory conditions (Hymenoptera: Formicidae)." Psyche 90: 425-436.
   Колонии муравья-рабовладельца H. americanus (Emery), и 2 его видов-хозяев (L. ambiguus Emery и L. longispinosus Roger) исследовались в лабораторных гнездах. Some of the slave-maker and host-species colonies were polydomous. In some polydomous slave-maker colonies, the slaves carried all the H. americanus workers into one nest before the onset of raiding. When thus assembled, the slave-makers efficiently captured brood from nearby host-species colonies. In other polydomous colonies where the slave-makers remained in more than one nest, the parasites conducted unco-ordinated raids and incurred many casualties. Several kinds of slave aggression against the slave-maker are described. However, slaves 'peacefully' augmented the slave worker forces of some incipient H. americanus colonies.
   
   
5. Bourke, A. (1987). The social biology of the slave-making ant Harpagoxenus sublaevis, Ph.D. dissert., University of Bath, United Kingdom, 269 p.
   ДИССЕРТАЦИЯ посвящена социальной биологии муравья-рабовладельца Harpagoxenus sublaevis. H. sublaevis workers can increase their inclusive fitness by procuring Leptothorax slaves to rear their kin, or by producing male offspring parthenogenetically. I describe work exploring the consequences of worker reproduction for colony social structure, temporal division of labour, and productivity in H. sublaevis. I also use data from a field H. sublaevis population to test the genetic relatedness hypothesis of sex ratio determination, taking into account potential confounding factors such as intra-colony genetic relatedness and population mating structure. In addition, I review the occurrence and significance of worker reproduction throughout the advanced social Hymenoptera. I also deal with issues concerning social parasitism in ants, reviewing the origin of slave-making (I conclude it arose via the temporary parasitism route) and describing an investigation of the host-parasite relations between H. sublaevis and Leptothorax (H. sublaevis appears to be the permanent winner in an asymmetric interspecific 'arms race' with its slave species. My principal findings and conclusions are as follows. First, reproductive H. sublaevis workers, despite being full sisters (intra-colony relatedness is maximal) form competitive dominance orders in which rank correlates with ovarian development. Queens inhibit worker egg-laying and dominance activity, and most worker-derived males are produced by orphaned (queenless) workers. The social structure of H. sublaevis colonies therefore represents the current state of a kin-selected queen-worker conflict over male parentage. Second, the level of relative sex investment in the H. sublaevis study population confirms the genetic relatedness hypothesis of sex ratio. Third, throughout the advanced social Hymenoptera, as in H. sublaevis, worker reproduction and concomitant queen-worker conflict over male parentage have been and remain potent influences on colony organization and function. Hence this thesis highlights the importance of intra-group reproductive conflict in social evolution. [Dissert. Abstr. Int. C 49: 567] [Order # DEVCO-50058]
   
   
6. Bourke, A. F. G. (1987). Alternative reproductive strategies in workers of the slavemaking ant Harpagoxenus sublaevis. Chemistry and biology of social insects. J. Eder and H. Rembold. Munich, Peperny: 259.
   
   
7. Bourke, A. F. G. (1988). "Dominance orders, worker reproduction, and queen-worker conflict in the slave-making ant Harpagoxenus sublaevis." Behav. Ecol. Sociobiol. 23: 323-333.
   Исследованы колонии моногинного муравья-рабовладельца Harpagoxenus sublaevis, a subset of workers formed a linear dominance order in which dominance was correlated with ovarian development, frequency of trophallaxis, length of time spent in the nest, but not body size. Identical dominance orders occurred in queenless colonies. Experiments in which the top-ranking workers were removed from queenless colonies demonstrated that worker dominance behaviour inhibits egg-laying in subordinates. A similar removal experiment showed queens restrict dominance behaviour and egg-laying in workers, probably pheromonally. Observations of slave raids indicated ovary-developed workers spent significantly less time scouting for slaves, and tended to participate less in slave raids, than workers without ovarian development. These findings suggest that potentially fertile H. sublaevis workers aggressively compete for egg-laying rights, consume extra food for egg development, and safeguard their reproductive futures by avoiding risks outside the nest. Hence worker reproduction in this species strongly influences the colony's social structure, nutrient flow, and division of labour, even though all workers in a colony are full sisters. I hypothesize that worker reproduction was formerly even more prevalent in H. sublaevis, with workers following the strategy of raising sisters and producing sons predicted by kinship theory. Its continued existence despite queen opposition conceivably results from selection on orphaned workers to reproduce, and the inability of slave-maker workers to raise female-biased broods. The social organization of H. sublaevis therefore highlights the importance both of worker reproduction and of the concomitant queen-worker conflict over male parentage in Hymenopteran social evolution.
   
   
8. Bourke, A. F. G., T. M. van der Have, et al. (1988). "Sex ratio determination and worker reproduction in the slave-making ant Harpagoxenus sublaevis." Behav. Ecol. Sociobiol. 23: 233-245.
   In a population of the monogynous slave-making ant Harpagoxenus sublaevis in S.E. Sweden, the mean proportion of dry weight investment in queens was 0.54. This result differed significantly from 0.75 but not from 0.5, matching the prediction from the genetic relatedness hypothesis of sex ratio applied to slave-makers, given (as confirmed by this study) single mating of queens, population-wide mate competition, and relatively low levels of worker male production. Sex investment appeared unaffected by resource availability. In the same 47 colony population sample, fertile slave-maker workers were found in every queenless colony (ca. 30% of all colonies), and in 58% of queenright colonies. Fertile workers occurred at a significantly higher frequency in the queenless colonies (19.2%) than in the queenright ones (9.8%), confirming that queenless conditions promote worker fertility. Fertile and sterile workers were similar in size. Electrophoretic allozyme analysis of ants from 49 colonies showed that: 1) queens mated singly; 2) female nestmates were full sisters (their regression coefficient of relatedness (+/- SE) was 0.735 +/- 0.044); 3) inbreeding did not occur; 4) queen and worker siblings were not genetically differentiated. Worker male production in queenright colonies was neither confirmed nor ruled out by the genetic data. However, production data indicated that queenless workers produced between 4.4 and 21.6% of all males. Overall colony productivity was largely determined by slave number, itself positively correlated with the number of slave-maker workers. There was an abrupt switch from all worker to all sexual production as colony size rose, as predicted by life history models. In queenright colonies, fertile slave-makers did not discernibly reduce colony productivity. Such workers occurred in queenright colonies with most slaves, suggesting they exploited energetic surpluses. Worker reproduction in H. sublaevis therefore appears to have greater influence at the level of individual behaviour than at colony or population level.
   
   
9. Buschinger, A. (1966). "Untersuchungen an Harpagoxenus sublaevis Nyl. (Hym. Formicidae), II: Haltung und Brutaufzucht." Insect. Soc. 13: 311-322.
   
   
10. Buschinger, A. (1966). "Untersuchungen an Harpagoxenus sublaevis Nyl. (Hym. Formicidae). I. Freilandbeobachtungen zu Verbreitung und Lebensweise." Insect. Soc. 13: 5-16.
    
    
11. Buschinger, A. (1968). "Locksterzein begattungsbereiter ergatoider Weibchen von Harpagoxenus sublaevis Nyl. (Hymenoptera, Formicidae)." Experientia 24(3): 297.
    [=Attractive behavior of ergatoid females of Harpagoxenus sublaevis Nyl. (Hymenoptera, Formicidae) prepared for copulation]
    
    
12. Buschinger, A. (1968). "Untersuchungen an Harpagoxenus sublaevis Nyl. (Hymenoptera, Formicidae), III: Kopula, Koloniegrьndung, Raubzьge." Insect. Soc. 15: 89-104.
    
    
13. Buschinger, A. (1971). "Weitere Untersuchungen zum Begattungsverhalten sozialparasitischer Ameisen (Harpagoxenus sublaevis Nyl. und Doronomyrmex pacis Kutter, Hym., Formicidae)." Zool. Anz. 187: 184-198.
    = Further studies of the mating behavior of the social parasitic ants (Harpagoxenus sublaevis Nyl. and Doronomyrmex pacis Kutter, Hymenoptera, Formicidae).
    
    
14. Buschinger, A. (1972). "Begattungsverhalten sozialparasitischer Ameisen aus dem Tribus Leptothoracini (Harpagoxenus sublaevis, Doronomyrmex pacis, Leptothorax kutteri)." Insect. Soc. 19: 416.
    = Copulatory behavior of socially parasitic ants from the Tribe Leptothoracini (Harpagoxenus sublaevis, Doronomyrmex pacis, Leptothorax kutteri)
    
    
15. Buschinger, A. (1972). "Giftdrusensekret als Sexualpheromon bei der ameise Harpagoxenus sublaevis." Naturwissenschaften 59: 313-314.
    = Poison gland secretion as pheromone of Harpagoxenus sublaevis.
    
    
16. Buschinger, A. (1973). Zum Sexualverhalten der dulotischen Ameise Harpagoxenus sublaevis (Nyl.) (Hym., Formicidae), Proc. 7 Congr. IUSSI, Lond., p. 50-56.
    
    
17. Buschinger, A. (1974). "Experimente und Beobachtungen zur Grundung und Entwicklung neuer sozietaten der sklavenhaltenden Ameise Harpagoxenus sublaevis (Nyl.)." Insect. Soc. 21: 381-406.
    = Experiments and observations on foundation and development of new colonies of the slavemaker ant, Harpagoxenus sublaevis (Nyl.)
    
    
18. Buschinger, A. (1975). "Eine genetische Komponente im Polymorphismus der dulotischen Ameise Harpagoxenus sublaevis." Naturwissenschaften 62: 239-240.
    = A genetic component in the polymorphism of the dulotic ant -Harpagoxenus sublaevis
    
    
19. Buschinger, A., G. Frenz, et al. (1975). "Untersuchungen zur Geschlechtstierproduktion der dulotischen Ameise Harpagoxenus sublaevis (Nyl.) (Hym., Formicidae)." Insect. Soc. 22: 169-181.
    = Investigations on sexual production in the dulotic ant, Harpagoxenus sublaevis (Nyl.) (Hymenoptera, Formicidae) Buschinger, A. and U. Winter (1975). "Der Polymorphismus der sklavenhaltenden Ameise Harpagoxenus sublaevis (Nyl.)." Insect. Soc. 22: 333-362.
    = Polymorphism of the slavemaker ant, Harpagoxenus sublaevis (Nyl.)
    
    
20. Buschinger, A. (1978). "Genetisch bedingte Entstehung geflugelter Weibchen bei der sklavenhaltenden Ameise Harpagoxenus sublaevis (Nyl.) (Hym., Form.)." Insect. Soc. 25: 163-172.
    = Genetically induced origin of alate females in the slavemaking ant, Harpagoxenus sublaevis (Nyl.)
    
    
21. Buschinger, A. (1983). "Sexual behavior and slave raiding of the dulotic ant, Harpagoxenus sublaevis (Nyl.) under field conditions (Hym., Formicidae)." Insect. Soc. 30: 235-240.
    A colony of Harpagoxenus sublaevis was artificially established in a near-natural habitat. The sexual calling behavior of young ergatomorphic queens was observed for the first time under field conditions on July 7 and 8 1981, between 20:15 and 21:30 Central European Time. Simultaneously, on July 7, the Harpagoxenus colony conducted a slave raid on a neighboring nest of Leptothorax acervorum ((Fabricius). Previously the sexual behavior was known only from laboratory studies, these observations now show that it is performed in an identical manner under natural conditions.
    
    
22. Buschinger, A. and T. M. Alloway (1978). "Caste polymorphism in Harpagoxenus canadensis M. R. Smith (Hym., Formicidae)." Insect. Soc. 25: 339-350.
    
    
23. Buschinger, A. and T. M. Alloway (1978 (1977)). "Population structure and polymorphism in the slave-making ant Harpagoxenus americanus (Emery) (Hymenoptera: Formicidae)." Psyche 84: 233-242.
    During the summer of 1977, we collected numerous colonies of the slave-making ant Harpagoxenus americanus in southern Ontario and northern Ohio. The number of dealate Harpagoxenus queens, intermorphs, and workers were recorded; and the production of young Harpagoxenus workers and sexuals were observed in 41 colonies. Some of the Harpagoxenus queens, all the intermorphs, and most of the Harpagoxenus workers were dissected to determine the structure and function of their reproductive organs. The queens, intermorphs, and most of the workers have 6 ovarioles. However, since only alate and dealate full queens have a spermatheca, they are the only individuals that can be inseminated. Thus, no true ergatoid queens exist in H. americanus. Nevertheless, many workers have functional ovaries and lay eggs, sometimes even in queenright colonies. We found 3 incipient colonies, 13 queenright colonies, and 23 'branch colonies' that lacked a dealate queen but contained one or more fully or partially fertile worker. Younger 'branch colonies' produce males, queens and workers, supposedly from larvae of their 'mother colony,' older 'branch colonies' only yield males. The significance of this population structure is discussed.
    
    
24. Buschinger, A. and T. M. Alloway (1979). "Sexual behaviour in the slave-making ant, Harpagoxenus canadensis M. R. Smith, and sexual pheromone experiments with H. canadensis, H. americanus (Emery), and H. sublaevis (Nylander) (Hymenoptera; Formicidae)." Z. Tierpsychol. 49: 113-119.
    
    
25. Buschinger, A. and U. Winter (1977). "Rekrutierung von Nestgenossen mittels Tandemlaufen bei Sklaveraubzugen der dulotischen Ameise Harpagoxenus sublaevis (Nyl.)." Insect. Soc. 24: 183-190.
    = Recruitment of nestmates by tandem-running during slave-raids of the dulotic ant, Harpagoxenus sublaevis (Nyl.)
    
    
26. Buschinger, A. and U. Winter (1978). "Echte Arbeiterinnen, fertile Arbeiterinnen und sterile Wirtsweibchen in Volkern der dulotischen Ameise Harpagoxenus sublaevis (Nyl.) Hym., Form.)." Insect. Soc. 25: 63-78.
    = True workers, fertile workers, and sterile host females in colonies of the dulotic ant, Harpogoxenus sublaevis (Nyl.) (Hymenoptera, Formicidae).
    
    
27. Clausen, R. (1939). "Harpagoxenus sublaevis Nyl. in der Schweiz." Mitt. Schweiz. Entomol. Ges. 17: 493-500.
    
    
28. Creighton, W. S. (1927). "The slave-raids of Harpagoxenus americanus." Psyche 34: 11-29.
    
    
29. Creighton, W. S. (1929). "Further notes on the habits of Harpagoxenus americanus." Psyche 36: 48-50.
    
    
30. Czechowski, W. and W. Czechowska (1999). "New sites in Poland and notes on the biology of socially parasitic ants Formicoxenus nitidulus (NYL.) and Harpagoxenus sublaevis (NYL.) (Hymenoptera, Formicidae)." Fragm. Faun. (Warsaw) 42: 1-6.
    
    
31. Del Rio Pesado, M. G. (1983). Polydomy in the slave-making ant, Harpagoxenus americanus (Emery) (Hymenoptera:Formicidae), M.Sc. Thesis, University of Toronto.
    
    
32. Del Rio Pesado, M. G. and T. M. Alloway (1983). "Polydomy in the slave-making ant, Harpagoxenus americanus (Emery) (Hymenoptera:Formicidae)." Psyche 90: 151-162.
    Field maps were made while collecting nests of the slave-making ant, Harpagoxenus americanus, and two of its host species, Leptothorax ambiguus and L. longispinosus. The ants were then transferred to artificial nests arranged to reconstruct the natural spatial relationships among nests. Ants from adjacent slave-maker nests often exchanged nestmates and brood for a period of time before moving into a single nest; and ants which had been living in a single nest in the laboratory sometimes moved into two nests. However, in other instances, ants from adjacent nests fought. These observations were interpreted as indicating that colonies of H. americanus sometimes occupy more than one nest (facultative polydomy). Nest population data were also presented and discussed.
    
    
33. Friedrich, B. (1981). Untersuchgen zum Wirtwahlverhalten von Jungkoniginnen der sklavenhaltenden Ameise Harpagoxenus sublaevis (Nyl.), Thesis, Darmstadt, 84 p.
    
    
34. Gregg, R. E. (1945). "The worker caste of Harpagoxenus canadensis Smith (Formicidae)." Can. Entomol. 77: 74-76.
    
    
35. Haeseler, V. (1990). "Zum Vorkommen von Harpagoxenus sublaevis (Nylander) in einem aufgeforsteten Hochmoorrest bei Oldenburg (Hymenoptera: Formicidae)." Braunschw. Naturkd. Schr. 3: 757-762.
    = On the occurrence of Harpagoxenus sublaevis (Nylander) in an afforested part of a former bog near near Oldenburg (Hymenoptera: Formicidae).
    
    
36. Heinze, J., D. Ortius, Kaib, M., Holldobler, B. (1994). "Interspecific aggression in colonies of the slave-making ant Harpagoxenus sublaevis." Behav. Ecol. Sociobiol. 35: 75-83.
    
    
37. Heinze, J., R. J. Stuart, Alloway, T.M., Buschinger, A. (1992). "Host specificity in the slave-making ant Harpagoxenus canadensis M. R. Smith." Can. J. Zool. 70: 167-170.
    
    
38. Kaib, M. and J. Heinze (1994). Cuticular hydrocarbons and aggression in colonies of the slave-making ant Harpagoxenus. Les Insectes Sociaux. 12th Congress of the International Union for the Study of Social Insects, Paris, Sorbonne, 21-27 August 1994. A. Lenoir, G. Arnold and M. Lepage. Paris, Universitй Paris Nord: 143.
    
    
39. Klemm, W. (1952/53). "Eine seltene Ameise, Harpagoxenus sublaevis (Nylander), im lande Salzburg." Mitt. naturw. Arbeitgem. Haus d. Natur. Salzburg, Zool. Arbeitgr. 3/4: 1-3.
    
    
40. Leighl, A. S. (1993). Factors affecting host specificity and habitat selection in the slave-making ant Harpagoxenus canadensis M.R. Smith and its host species, M.Sc. thesis, University of Toronto, Canada, 64 p.
    [Masters Abstr. Int. 32: 561] [Order # MAMM83378]
    
    
41. Ollett, D. G., E. D. Morgan, et al. (1987). "The contents of the Dufour gland of the ant Harpagoxenus sublaevis Nyl. (Hymenoptera: Formicidae)." Z. Naturforsch. Sec. C Biosci. 42: 141-146.
    The volatile components of the Dufour gland secretion of workers of the ant Harpagoxenus sublaevis have been analysed by gas chromatography and mass spectrometry. Over 20 components have been identified, consisting of linear and terpenoid hydrocarbons. Each gland contains approximately 6µg of hydrocarbons with n-heptadecene (40%) and n-†heptadecadiene (30%) as the major components. Two terpenoid hydrocarbons, (E)-Я-farnesene and a homofarnesene isomer were also identified.
    
    
42. Pisarski, B. (1963). "Nouvelle espиce du genre Harpagoxenus For. de la Mongolie (Hymenoptera, Formicidae)." Bull. Acad. Pol. Sci. Ser. Biol. 11: 39-41.
    
    
43. Schumann, R. and A. Buschinger (1991). "Selective acceptance of alien host species pupae by slaves of the dulotic ant, Harpagoxenus sublaevis (Hymenoptera, Formicidae, Myrmicinae)." Ethology 88: 154-162.
    
    
44. Smith, M. R. (1939). "The North American ants of the genus Harpagoxenus Forel, with the description of a new species (Hymenoptera: Formicidae)." Proc. Entomol. Soc. Washington 41: 165-172.
    
    
45. Stuart, R. J. (1982 (1981)). "Abdominal trophallaxis in the slave-making ant, Harpagoxenus americanus (Hymenoptera: Formicidae)." Psyche 88: 331-334.
    
    
46. Stuart, R. J. (1984). "Experiments on colony foundation in the slave-making ant Harpagoxenus canadensis M. R. Smith (Hymenoptera; Formicidae)." Can. J. Zool. 62: 1995-2001.
    Queens of obligatory socially parasitic ants are unable to found colonies independently and typically intiate new colonies by invading host-species nests and usurping the role of reproductive. In some cases, fission ('budding') of parasite colonies also occurs. This study is the first to investigate colony foundation in the obligatory slave-making ant Harpagoxenus canadensis M. R. Smith. Laboratory experiments examined whether colonies could be founded (i) independently by a parasite queen, (ii) jointly by a parasite and host queen, (iii) by budding, and (iv) by usurpation. Parallel experiments using host-species queens (Leptothorax sp.) were conducted for independent colony foundtation and joint colony foundation by two conspecific queens. The results indicate that parasitic queens found colonies by usurpation. Parasites invaded host nests using either an 'active' aggressive strategy or a relatively less aggressive 'passive' strategy, obtained host brood, and were occasionally adopted by a large portion of the host colonies' adults. Host-specific queens successfully founded colonies both independently and jointly.
    
    
47. Stuart, R. J. and T. M. Alloway (1983). "The slave-making ant, Harpagoxenus canadensis M.R. Smith, and its host-species, Leptothorax muscorum (Nylander): slave raiding and territoriality." Behaviour 85: 58-90.
    The territorial hypothesis suggests that slave raiding in ants evolved from the territorial behaviour of free living ancestors. In this study, the slave raiding behaviour of Harpagoxenus canadensis was observed and compared to the territorial behaviour of its host species, Leptothorax muscorum. Under laboratory conditions, colonies of both species attack and raid nearby L. muscorum nests, and display very similar raiding behaviours. At the conclusion of their raids, both species appropriate brood from raided nests. However, H.canadensis colonies rear the captured brood to produce slave workers, while L.muscorum colonies mutilate it and feed it to their larvae. In H. canadensis colonies, slaves join with the slave-makers in all aspects of raiding activity; and the differential treatment of captured brood was the only major difference we observed between the behaviour of enslaved and unenslaved L.muscorum workers. During interactions between colonies, workers of both species explore laboratory arenas ('scouting'), and attack alien members of either species that they encounter. Nonetheless H.canadensis workers are markedly superior to L.muscorum in their fighting abilities, and consequently their colonies are capable of conducting successful raids against much larger host-species colonies. Workers of both species recruit nestmates to the scene of fights by leading tandem runs ('alarm recruitment'); a behaviour which is typical of the territorial behaviour of various ant species. However, the discovery of such recruitment in H.canadensis is remarkable because all other known obligatory Leptothoraciae[!] slave-makers recruit nestmates during their slave raids only after they discover the target- colony's nest, and thereafter only lead recruitment to target nest itself. These more specialized forms of slave-raid recruitment occur in an aggressive intercolonial context virtually identical to that of alarm- recruitment and could easily have evolved from the alarm recruitment of free-living ancestors. Tandem-run recruitment also occurred during the transport of captured brood following raids by colonies of either species. However, the context of this recruitment was quite different from that performed during the early stages of raid. Since H.canadensis colonies rear captured brood, this recruitment may be analogous to that led during emigrations to new nest sites ('transport- recruitment'). However, for L.muscorum colonies, this recruitment might be properly considered as analogous to that led to food sources ('food- recruitment'). The results of this study demonstrate that all of the major elements which comprise the slave raids of H.canadensis and other Leptothoraciae[!] slave-makers occurring during the apparently opportunistic, territorial raids of L. muscorum, and implicate such territorial behaviour as the proximate ancestor to slave raiding in this group of ants. The relatively minor differences which do occur can be regarded as specialization for slave- making. Since H.canadensis retains alarm-recruitment as a means of slave-raid recruitment, its slave raids can be regarded as the most primitive in the genus Harpagoxenus; and relatively more primitive than those of any other known obligatory Leptothoracine slave-maker.
    
    
48. Sturtevant, A. H. (1927). "The social parasitism of the ant Harpagoxenus americanus." Psyche 34: 1-9.
    
    
49. Wesson, L. G. (1939). "Contributions to the natural history of Harpagoxenus americanus Emery (Hymenoptera: Formicidae)." Trans. Am. Entomol. Soc. 65: 97-122.
    
    
50. Winter, U. (1976). Untersuchgen zur Dulosis, insbesondere zum Raubzugverhalten der Ameise Harpagoxenus sublaevis (Nyl.), Diplomarbeit TH Darmstadt.
    
    
51. Winter, U. (1979). "Untersuchungen zum Raubzugverhalten der dulotischen Ameise Harpagoxenus sublaevis (Nyl.)." Insect. Soc. 26: 123-135. = Investigations on the raiding behavior of the dulotic ant Harpagoxenus sublaevis (Nyl.)
    
    
52. Winter, U. and A. Buschinger (1984). Genetically mediated queen polymorphism and caste determination in the slave-making ant, Harpagoxenus sublaevis. [Abstract], P. 516 in: XVII International Congress of Entomology. Hamburg, Federal Republic of Germany, August 20-26, 1984. Abstract Volume. Hamburg: 17th International Congress of Entomology, 960 p.
    
    
53. Winter, U. and A. Buschinger (1986). "Genetically mediated queen polymorphism and caste determination in the slave-making ant, Harpagoxenus sublaevis (Hymenoptera: Formicidae)." Entomol. Gen. 11: 125-137.
    In the European slavemaking ant Harpagoxenus sublaevis (Nylander 1849), there is a polymorphism for the form of the queens [females] controlled by a single locus with 2 alleles, E and e. Only homozygous ee specimens can be gynomorphic (winged) females, whereas the females derived from the other genotypes (EE and Ee) are wingless ergatomorphs. The females mate only once, the colonies are monogynous, and most natural colonies have an ergatomorphic female. Breeding experiments demonstrated that the E|e locus (or one or more others tightly linked to it) affects neither behavior nor reproductive rate, but does have a strong influence on caste determination beyond the gynomorph/ergatomorph dichotomy. This influence probably stems from 2 effects: (a) that the E allele slows the development of diploid larvae [L], and (b) that it increases the inhibitory effects that a female has on the likelihood of a L becoming a female. Thus, the potential of ee L to become female is high, whereas Ee and, especially, EE L are more likely to became workers. The polymorphism of the form of the female (gynomorph or egratomorph) therefore is now a side-effect of the polymorphism, with the main impact of selection falling on caste-biassing: the potential of a L to become a female of either kind.