Добавлено 16-10-2005 

Крупноглазые: филогения и эволюция

Philip S. Ward & Douglas A. Downie* E-mail: psward@ucdavis.edu Department of Entomology, University of California, Davis, CA 95616, U.S.A. *Present address: Department of Zoology and Entomology, Rhodes University, Grahamstown 6140, South Africa. The ant subfamily Pseudomyrmecinae (Hymenoptera: Formicidae): phylogeny and evolution of big-eyed arboreal ants

"Systematic Entomology", 2005, Volume 30, Issue 2 , July 2005, Pages 310-335 (April 2005)

      Подсемейство Pseudomyrmecinae включает три рода крупноглазых древесных муравьев, широко распространенных в тропических и субтропических областях: Pseudomyrmex (200 видов в Америке), Myrcidris (2 вида в Южной Америке) и Tetraponera (100 видов, Палеотропика). The phylogenetic relationships among these ants were investigated using DNA sequence data (5.2 kb from 18S rDNA, 28S rDNA, wingless, abdominal-A, and long-wavelength rhodopsin genes) and 144 morphological characters, both separately and in combination. Data were gathered from a representative set of 49 pseudomyrmecine species, plus 18 species from various outgroups. There was substantial agreement among the results obtained from different datasets, and from different methods of phylogenetic inference (parsimony, Bayesian inference).
      Поддержана монофилия следующих групп (100% bootstrap support and 1.00 posterior probability in the molecular dataset): Pseudomyrmecinae, Pseudomyrmex, и Pseudomyrmex + Myrcidris. Статус рода Tetraponera менее ясен: the DNA sequence data indicate that the genus is paraphyletic, but morphological features and a unique insertion in the 28S gene support the monophyly of this taxon.
      7 из 9 видовых групп рода Pseudomyrmex, выделенных ранее только на основании морфологических признаков, are strongly upheld, but monophyly is rejected for the P. pallens group and the P. viduus group. In the latter case, molecular evidence indicates the existence of two independent clades, associated with the ant-plants Triplaris and Tachigali, respectively, whose convergent morphological features had caused them to be placed erroneously in the same species group. Полученные результаты подтверждает более раннее утверждение, что облигатные ассоциации с растениями, несущими специализированные образования domatia возникли по крайней мере двенадцать раз в этом древесном подсемействе.
      Молекулярные и морфологические данные поддерживают гипотезу об объединении в качестве сестринских групп двух близких подсемейств: Pseudomyrmecinae и Myrmeciinae (84% parsimony bootstrap, combined dataset), которые имеют общий корень в Меловом периоде в южном полушарии. Pseudomyrmecinae вероятно возникли впервые в Палеотропике и уже затем распространились на других материках: в Африке и Южной Америке, где произошло мощное видообразование. Сходные отношения можно обнаружить в самой современной системе муравьев Болтона (Bolton, 2003), в которой в группу подсемейств "The myrmeciomorph subfamilies" объединяются Myrmeciinae (+Prionomyrmecini=Nothomyrmeciinae) и Pseudomyrmecinae.



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      PSEUDOMYRMECINAE

      Псевдомирмецины это весьма характерные древесные обитатели тропических лесов. Они обладают привлекательной яркой окраской и сильным жалом, быстро передвигаются. Большинство из 300 видов обитают в ветвях древесных растений, обычно в полостях ранее сделанных личинками жуков и бабочек. Более чем 40 видов этих муравьев являются облигатными обитателями специализированных муравьиных растений - мирмекофитов. Муравьи обитают в специальных полостях (myrmecodomatia) в живых тканях таких мирмекофитов, обеспечивая им некоторую степень защиты от фитофагов (Janzen, 1966, 1972; Benson, 1985; Ward, 1991; Davidson & McKey, 1993). Подобные мутуалистические ассоциации возникали в ходе эволюции как минимум 12 раз в подсемействе Pseudomyrmecinae (Ward, 1991).

      В составе подсемейства Pseudomyrmecinae выделяют 3 рода: Myrcidris Ward (2 вида), Pseudomyrmex Lund (около 200 видов) и Tetraponera F. Smith (около 100 видов). Кроме того остаются неописанными десятки новых видов. Прежние морфологичесике исследования говорили о монофилии трех родов и анцестральное положение рода Myrcidris по отношению к сестринской группе (Pseudomyrmex + Tetraponera) (Ward, 1990, 1991).

      Род Myrcidris известен только из нескольких мест в Бразилии и Гайаны (Guyana) и в этой работе был исследован вид Myrcidris epicharis.

      Род Pseudomyrmex широко распространен в Новом Свете с наибольшим разнообразием в Южной Америке. В составе рода Pseudomyrmex выделяют 9 групп видов (Ward, 1989, 1993, 1999) (далее в скобках указаны исследованные виды):
1, gracilis group (Pseudomyrmex godmani, P. gracilis, P. nigropilosus);
2, sericeus group (P. ita, P. sericeus, P. cordiae);
3, tenuis group (P. boopis, P. tenuis, P. denticollis, P. termitarius);
4, pallens group (P. apache, P. elongatulus, P. pallens, P. phyllophilus);
5, oculatus group (P. cubaensis, P. oculatus);
6, subtilissimus group (P. spiculus, P. tenuissimus);
7, pallidus group (P. pazosi, P. simplex, P. holmgreni, P. pallidus);
8, ferrugineus group (P. flavicornis, P. nigrocinctus);
9, viduus group (P. concolor, P. tachigaliae, P. dendroicus, P. viduus, P. kuenckeli). Все 5 видов из последней группы (P. viduus group) живут в мирмекофитных растениях рода Triplaris (Polygonaceae) (P. dendroicus и P. viduus) и рода Tachigali (Fabaceae) (P. concolor и P. tachigaliae). Однако 5-й вид P. kuenckeli связан с растениями не так тесно (not an ant-plant specialist; Ward, 1999).
И ещё 4 вида как incertae sedis (P. oki, P. filiformis, P. haytianus, P. subater).

      Род Tetraponera ограничен Палеотропикой, с ископаемыми представителями из Балтийского янтаря Европы (Eocene/Oligocene) (Wheeler, 1915; Dlussky, 1997; Ward, 1990; Bolton, 1995). В Афротропике встречается около 25 видов Tetraponera, на Мадагаскаре - около 40, и в Азии - около 35. Были исследованы 15 видов (Tetraponera allaborans, T. clypeata, T. penzigi, T. tessmanni, T. ambigua, T. ophthalmica, T. morondaviensis, T. aethiops, T. rufonigra, T. pilosa, T. natalensis, T. caffra, T. nigra, T. punctulata, T. grandidieri). Таксономическая ревизия Индо-Австралийских видов Tetraponera показала наличие 4 групп видов, но это предварительные данные. Вероятной родиной этого рода является Африка, откуда он несколькими путями проник в Азию (Ward, 2001). Некоторые старые родовые таксоны совпадают с новыми группами видов: Pachysima Emery (типовой вид T. aethiops) с группой rufonigra group; Tetraponera (типовой вид T. nigra) с группой T. nigra group (в этом исследовании представлены T. nigra и T. punctulata); и Sima Roger (типовой вид T. allaborans).

      ИЗВНЕ. Из внешних групп были исследованы 15 муравьев из других подсемейств: Paraponerinae (Paraponera clavata), Ectatomminae (Ectatomma opaciventre, Typhlomyrmex rogenhoferi), и другие (Cerapachys larvatus, Proceratium stictum, Amblyopone pallipes, Hypoponera opacior, Leptomyrmex erythrocephalus, Formica moki, Myrmica tahoensis), Myrmeciinae (Myrmecia pilosula, Myrmecia fulvipes, Myrmecia pyriformis, Myrmecia picta, Nothomyrmecia macrops) и 3 вида жалящих перепончатокрылых (оса-Bradynobaenidae вида Chyphotes mellipes и оса-веспиды Mischocyttarus flavitarsis, пчела Apis mellifera).

      Но проблема эволюционных взаимоотношений этих трех родов до сих пор оставалась нерешенной окончательно. Что и потребовало привлечь к данным морфологии, зоогеографии и биологии ещё и молекулярные данные и исследование ДНК. Из 144 исследованных в этой новой работе признаков у рабочих взяли 22 признака, у самок - 3, а у самцов - 61 признак. Ещё 46 признаков были сходны у самок и рабочих, 2 признака - у самцов и самок, и 10 признаков были одинаковы у всех трех каст.

      ГЕНЫ. Молекулярные данные включали сегменты 5 ядерных генов: гены РНК малой (18S) и большой (28S) субъединиц рибосомы, кодирующий белок ген abdominal-A (abd-A), и гены wingless (wg) и long-wavelength rhodopsin (LW Rh). Процедура Polymerase chain reaction (PCR) для генов 18S и 28S была сходной с методикой Ward & Brady (2003).

      Всего при составлении кладограмм учтены 5335 признаков = 144 морфологических + 5191 признаков по 5 генам (1868 по гену 18S + 1835 по гену 28S + 412 по гену wg + 618 по гену abd-A + 458 по гену LW Rh). Однако вариабельными из них оказались только 1460 = 144 + 1316 (169+467+216+220+244), соответственно.

      СПЕЦИАЛИСТЫ. Из 49 исследованных видов псевдомирмецин 12 (Pseudomyrmex flavicornis, P. nigrocinctus, P. dendroicus, P. viduus, P. concolor, P. tachigaliae, P. tenuissimus, P. nigropilosus, P. cordiae, Myrcidris epicharis, Tetraponera aethiops, T. tessmanni, T. penzigi, T. punctulata) являются облигатными жильцами (специалистами) растений-мирмекофитов и ещё 2 вида (P. tenuissimus, T. punctulata) близки к некоторым из этих ant-plant specialists (например, к неисследованным здесь P. subtilissimus и T. tucurua, соответственно). Заселяемые псевдомирмецинами мирмекофиты представлены 12 семействами растений и 19 их родами (Ward, 1991; Davidson & McKey, 1993).

Здесь я подобрал в базе Formis-2003 основные работы (около 75) по муравьям Pseudomyrmecinae, который дал 199 ссылок, в том числе 28 в заголовках статей.

Поиск по другим словам дает такие результаты: Pseudomyrmex (который дал 506 ссылок, в том числе 55 в заголовках статей), Myrcidris (9 и 0) и Tetraponera (145 и 30), acacia-ants (который дал 6 ссылок, в том числе 28 в заголовках статей).

1. Beulig, M. L. and D. H. Janzen (1969). "Variation in behavior among obligate Acacia-ants from the same colony, (Pseudomyrmex nigrocincta)." J. Kansas Entomol. Soc. 42: 58-67.
   
   
2. Billen, J. and A. Buschinger (2000). "Morphology and ultrastructure of a specialized bacterial pouch in the digestive tract of Tetraponera ants (Formicidae, Pseudomyrmecinae)." Arthropod Struct. Dev. 29: 259-266.
   Исследован пищеварительный тракт древесных муравьев некоторых видов рода Tetraponera, который характеризуется заметным грушевидным дивертикулумом (diverticulum) при переходе между midgut и intestine, что пока не было найдено ни у одного другого вида муравьев. Поскольку этот орган заполнен массой бактерий, авторы предлагают определять его как бактериальный мешок или сумка (pouch lumen). Its distal wall is formed by a thin ectodermal epithelium, through which tens of tracheal branches penetrate into the bacterial mass that fills the pouch lumen. The proximal wall, in contrast, is formed by a cylindrical epithelium with a conspicuous microvillar differentiation of the apical cell membrane, but without a cuticular lining. The contact region between both epithelia occurs as a complex fold surrounding the pouch like a belt. The Malpighian tubules open into the pouch through the cylindrical epithelium adjacent to the belt fold. The functional significance of the bacterial pouch remains unknown, although the abundant presence of bacteria may indicate a symbiotic function. The conspicuous tracheolar supply illustrates the metabolic activity in the pouch, while the microvillar differentiation of the cylindrical epithelium may be interpreted in the uptake of metabolites from the pouch lumen.
   
   
3. Blum, M. S. and P. S. Callahan (1963). "The venom and poison glands of Pseudomyrmex pallidus (F. Smith)." Psyche 70: 69-74.
   
   
4. Blum, M. S. and H. R. Hermann (1978). Venoms and venom apparatuses of the Formicidae: Myrmeciinae, Ponerinae, Doylinae, Pseudomyrmecinae, Myrmicinae, and Formicinae. Handbuch der Experimentellen Pharmakologie. 48: 801-869.
   
   
5. Buschinger, A., R. W. Klein, et Maschwitz, U. (1994). "Colony structure of a bamboo-dwelling Tetraponera sp. (Hymenoptera: Formicidae: Pseudomyrmecinae) from Malaysia." Insect. Soc. 41: 29-41.
   
   
6. Cokendolpher, J. C. and O. F. Francke (1990). The ants (Hymenoptera, Formicidae) of western Texas. Part II. Subfamilies Ecitoninae, Ponerinae, Pseudomyrmecinae, Dolichoderinae, and Formicinae, Special Publications, The Museum, Texas Tech University number 30, p. 1-76.
   Вторая часть описания муравьев Техаса. В Части I из этой серии другие авторы - Moody and Francke (1982) - рассмотрели фауну подсемейства Myrmicinae, которое представленов Западном Техасе 17 родами и 89 видами муравьев. Теперь исследованы 24 рода и 83 вида Западного Техаса из других подсемейств: Ecitoninae, Ponerinae, Pseudomyrmecinae (Pseudomyrmex apache, Pseudomyrmex pallidus), Dolichoderinae, и Formicinae. Two additional species (Formica neoclara and Hypoponera punctatissima) are probably also present, based on their known distributions elsewhere in Texas and surrounding states. Although several species are wide ranging, most are restricted to one or two vegetative regions. Thirteen wide ranging species are present in all of the western Texas regions: Brachymyrmex depilis, Camponotus discolor, C. festinatus, Conomyrma bicolor, C. flava, C. insana, Forelius foetidus, F. pruinosus, Formica gnava, Labidus coecus, Myrmecocystus placodops, Odontomachus clarus, and Paratrechina vividula. Eight other species are relatively common and were found in four of the five regions sampled: Camponotus sansabeanus, C. vicinus, Hypoponera opacior, Myrmecocystus depilis, M. mendax, M. mimicus, Neivamyrmex nigrescens, and Paratrechina terricola. More species were encountered in the Trans-Pecos region than in any other region in western Texas. Furthermore, more species are restricted to the Trans-Pecos area than all others taken together from the other regions in western Texas. Of the 58 species recorded from the Trans-Pecos, 21 were restricted to that region: Acanthomyops arizonicus, Amblyopone pallipes, Camponotus abdominalis transvectus, C. cuauhtemoc, C. ocreatus, C. ulcerosus, Cerapachys davisi, Colobopsis pylartes, Formica nitidiventris, F. puberula, Hypoponera opaciceps, Lasius sitiens, Liometopum luctuosum, Myrmecocystus melliger, Neivamyrmex fallax, N. macropterus, N. minor, N. pauxillus, Paratrechina austroccidua, P. bruesii, and P. pallidus. The wide variety of habitats- from lowland deserts to highland coniferous forests certainly- accounts for much of this diversity in ants. Approximately the same numbers of ant species occur in the High Plains, Rolling Plains, and portion of the Edwards Plateau in western Texas; 34, 38, and 47, respectively. Those species restricted to the High Plains of western Texas include: Formica bradleyi, F. gynocrates, F. montana, F. sp. (near integroides), and Myrmecocystus romainei. Species recorded only from the Rolling Plains are: Acanthomyops interjectus, Formica pallidefulva, F. subsericea, and Myrmecocystus navajo. The two species recorded only from the Edwards Plateau are Neivamyrmex melsheimeri and Proceratium compitale. Only 23 species are recorded from that portion of the Rio Grande Plains which extends into western Texas. Of those species, only three, Neivamyrmex fuscipennis, N. texanus, and Pachycondyla villosa do not occur elsewhere in western Texas.
   
   
7. Dejean, A., J. Orivel, Corbara, B., Olmsted, I., Lachaud, J.P. (2001). "Nest site selection by two polistine wasps: the influence of Acacia- Pseudomyrmex associations against predation by army ants (Hymenoptera)." Sociobiology 37: 135-146.
   
   
8. Dey, S. (1998). "A scanning electron microscopic study on the body cuticle and associated sensilla of an ant (worker) Tetraponera rufonigra." Proc. Nat. Acad. Sci. India Sect. B Biol. Sci. 68: 37-44.
   
   
9. Diniz, J. L. M. (1997). "O aparelho de ferrгo de formigas. Parte I. Pseudomyrmex gracilis (Fabricius) (Pseudomyrmecinae) e Myrmecia nigriscapa Roger (Myrmeciinae) (Hymenoptera, Formicidae)." Pap. Avulsos Zool. 40: 175-188.
   Исследован жалящий аппарат муравьев Pseudomyrmex gracilis и Myrmecia nigriscapa Roger.
   
   
10. Dlussky, G. M. and A. G. Radchenko (1990). The ants of Vietnam. S/f Pseudomyrmecinae. S/f Myrmicinae. Tribes Calyptomyrmecini, Meranoplini and Cataulacini. [in Russian]. News of faunistics and systematics. I. A. Akimov, I. G. Emelianov, M. D. Zerova and e. al. Kiev. 184 p., Naukova Dumka: 119-125.
    
    
11. Duelli, P. (1977). "Das soziale Trageverhalten bei neotropischen Ameisen der Gattung Pseudomyrmex (Hym., Formicidae): eine Verhaltensnorm als Hinweis fьr Phylogenie und Taxonomie?" Insect. Soc. 24: 359-365.
    Рассмотрено все многообразие способов переноски собратьев по гнезду у муравьев. The carrying posture observed in three neotropical species belonging to the subfamily Pseudomyrmecinae corresponds exactly with the carrying posture of some myrmicinae ants (Fig. 1). The Myrmicinae belong to the Poneroid complex, whereas Pseudomyrmex is supposed to join the Myrmecioid complex (Fig. 2). All other members of the myrmecioid complex studied to present show basically differing carrying postures (Formicinae, Dolichoderinae) or only non-stereotyped carrying behaviour (Myrmeciinae). Two hypotheses are discussed: 1. The observed carrying behaviour has evolved independently in both Pseudomyrmecinae and Myrmicinae. This behavioral analogy comes along with the proposed anatomical analogy concerning the double-noded abdominal pedicel as well as the occurrence of stridulatory organs. 2. The carrying posture of Pseudomyrmex and of some Myrmicinae evolved from the same behavioral origin. Consequently, the phylogenetic relation and thus the taxonomic position of Pseudomyrmex ought to be changed considerably with respect to the Poneroid complex. It would be interesting to know the carrying behaviour of some pseudomyrmecine species from the old world tropics.
    
    
12. Emery, C. (1915). "Sima oder Tetraponera?" Zool. Anz. 45: 265-266.
    
    
13. Errard, C. (1984). "Evolution, en fonction de l'age, des relations sociales dans les colonies mixtes heterospecifiques chez les fourmis des genres Camponotus et Pseudomyrmex." Insect. Soc. 31: 185-198.
    Эволюция и функция смешанных колоний из муравьев разных родов Camponotus и Pseudomyrmex
    
    
14. Fraga, N. d. J., J. H. C. Delabie, et al. (1999). "Diversidade de Pseudomyrmex spp (Formicidae: Pseudomyrmecinae) em manguezais no sudeste da Bahia." Naturalia 24: 103-105.
    
    
15. Hink, W. F., P. W. Pappas, et al. (1994). "Partial biochemical characterization of venom from the ant, Pseudomyrmex triplarinus." Toxicon 32(7): 763-772.
    Биохимическое исследование ядоа муравьев Pseudomyrmex triplarinus. Яд содержит 12 протеинов с молекулярной массой более > 100,000-4200, которые составляют 41.5% сухого веса. In comparison with published data on ant, wasp, and bee venoms, whole venom has intense phospholipase activity and intermediate hemolytic activity. Four major proteins were isolated and purified by low pressure chromatography. The most abundant protein had a mol. wt of 4200 and weak hemolytic activity. The second most common protein was 20,400 and had phospholipase A2 activity. The other two major proteins had mol. wts of 24,500 and 14,100 and both exhibited phospholipase and direct hemolytic activities. There are eight minor proteins (> 100,00-40,000), each present at about 1% or less of the total protein. Assayed as a mixture, they had hyaluronidase activity. Seventeen free amino acids were detected with aspartic acid, glutamic acid, and proline together making up 72% of the total mass of amino acids. Glycerol was present at a concentration of 3.1% of the dry weight and the venom was devoid of lipids.
    
    
16. Jaffe, K., M. E. Lopez, et al. (1986). "On the communication systems of the ants Pseudomyrmex termitarius and P. triplarinus." Insect. Soc. 33: 105-117.
    
    
17. Janzen, D. H. (1972). "Protection of Barteria (Passifloraceae) by Pachysima ants (Pseudomyrmecinae) in a Nigerian rain forest." Ecology 53 (5): 885-892.
    Защита муравьями Pachysima деревьев Barteria в Нигерийских джунглях. In a Nigerian rain forest, occupation of Barteria fistulosa saplings by Pachysima aethiops ants results in the plants having more leaves, more branches, more leaves per branch, and less damage to the shoot tips than do unoccupied Barteria. At present this difference is the result of insect damage to unprotected Barteria, but several traits of the ant colony suggest that in the recent past, large browsing mammals may also have been a threat to Barteria. These traits include a few large workers per colony, a deeply painful sting, and the habit of frequently dropping from the Barteria crown. Pachysima workers attack plants adjacent to their Barteria and clean the large Barteria leaves of depris and epiphyllae. *[Pachysima aethiops (=Tetraponera, Pseudomyrmecinae)]
    
    
18. Janzen, D. H. (1965). The interaction of the Bull's-horn acacia (Acacia cornigera L.) with one of its ant inhabitants (Pseudomyrmex fulvescens Emery) in eastern Mexico, Ph.D. dissert., University of California, Berkeley, 527 p. [Dissert. Abstr. 26: 1228-9] [Order # 6508187]
    
    
19. Janzen, D. H. (1967). "Interaction of the bull's-horn acacia (Acacia cornigera L.) with an ant inhabitant (Pseudomyrmex ferruginea F. Smith) in eastern Mexico." Univ. Kans. Sci. Bull. 47: 315-558.
    
    
20. Janzen, D. H. (1973). "Evolution of polygynous obligate acacia-ants in western Mexico." J. Anim. Ecol. 42: 727-750.
    
    
21. Janzen, D. H. (1975). "Pseudomyrmex nigropilosa: a parasite of a mutualism." Science 188: 936-937.
    
    
22. Janzen, D. H. (1983). Pseudomyrmex ferruginea (hormiga del cornizuelo, acacia-ant). Costa Rican natural history. D. H. Janzen. Chicago. xi + 816 p., University of Chicago Press: 762-764.
    
    
23. Keeler, K. H. (1981). "Infidelity by acacia-ants." Biotropica 13: 79-80.
    
    
24. Kempf, W. W. (1958). "Estudos sobre Pseudomyrmex. II. (Hymenoptera: Formicidae)." Stud. Entomol. (n.s.)1: 433-462.
    
    
25. Kempf, W. W. (1960). "Estudo sobre Pseudomyrmex I. (Hymenoptera: Formicidae)." Rev. Brasil. Entomol. 9: 5-32.
    
    
26. Kempf, W. W. (1961). "Estudos sobre Pseudomyrmex. III. (Hymenoptera: Formicidae)." Stud. Entomol. 4: 369-408.
    
    
27. Kempf, W. W. (1967). "Estudos sobre Pseudomyrmex. IV (Hymenoptera: Formicidae)." Rev. Brasil. Entomol. 12: 1-12.
    
    
28. Kempf, W. W. and K. Lenko (1976). "Levantamento da formicifauna no litoral norte e ilhas adjacentes do Estado de Sгo Paulo, Brasil. I. Subfamilias Dorylinae, Ponerinae e Pseudomyrmecinae (Hym., Formicidae)." Stud. Entomol. 19: 45-66.
    = Survey of the ant fauna of the northern coast and islands adjacent to Sao Paulo State, Brazil: I. The subfamilies Dorylinae, Ponerinae, and Pseudomrymecinae (Hymenoptera: Formicidae). *[H. iheringi frequently misspelled H. jheringi. Acromyrmex, Atta, C. cingulatus, C. melanoticus are prey of army ants. P. angustus, Z. angustus, & Z. striativentris nested in same trees as Pseudomyrmex. Staphylinidae & E. parva found with L. praedator.]
    
    
29. Kistner, D. H. and R. W. Klein (1996). "A new species of Orphnebius (Coleoptera: Staphylinidae) associated with Tetraponera sp. (Hymenoptera: Formicidae) in Malaysia." Sociobiology 27: 79-91.
    
    
30. Klein, R. W. (1987). Colony structures of three species of Pseudomyrmex (Hymenoptera: Formicidae: Pseudomyrmecinae) in Florida. Chemistry and biology of social insects. J. Eder and H. Rembold. Mьnchen. xxxv + 757 p., Verlag J. Peperny: 107-108.
    
    
31. Klein, R. W. (1987). A workerless inquiline in Pseudomyrmex (Hymenoptera: Formicidae: Pseudomyrmecinae). Chemistry and biology of social insects. J. Eder and H. Rembold. Mьnchen. xxxv + 757 p., Verlag J. Peperny: 623-624.
    
    
32. Klein, R. W. (1990). Untersuchungen zum Nahrungserweb von Pseudomyrmex und Tetraponera (Formicidae: Pseudomyrmecinae), unter besonderer Berьcksichtigung des Spur- und Orientierungsverhaltens, Diplom-thesis, University of Frankfurt.
    
    
33. Klein, R. W. (1994). Orphnebius politus Cameron (Coleoptera: Staphylinidae), a specialized predator of the bamboo ant Tetraponera sp. (Hymenoptera: Formicidae: Pseudomyrmecinae) with subsocial behavior. 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: 200.
    meeting abstract *[Beetle attacks & paralyzes ant; feeds on it & raises its larvae on it. Males show sexual pheromone calling behavior, & may offer a dead ant as a nuptial gift.]
    
    
34. Klein, R. W. (1997). "A specialized ant predator with brood care: the SE-Asian staphylinid Orphnebius kleini Kistner (Coleoptera: Staphylinidae: Aleocharinae) preys on Tetraponera sp. bamboo-ants (Hymenoptera: Formicidae: Pseudomyrmecinae) and rears its larvae on them." Mitt. dtsch. Ges. Allg. Angew. Entomol. 11: 401-404.
    
    
35. Klein, R. W., D. Kovac, Buschinger, A., Schellerich, A., Maschwitz, U.(1994). Mealybug-transport by swarming queens, water bailing from flooded nest chambers, and other adaptive strategies of a southeast asian bamboo ant (Hymenoptera: Formicidae: Pseudomyrmecinae: Tetraponera). 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: 200.
    [meeting abstract] Bamboo with its hollow internodes is utilized as habitat by many generalist arboreal ants. Interestingly, a few species of ants have become obligate bamboo specialists, among them an as yet undescribed Tetraponera sp. near attenuata F. Smith (T. sp. PSW-80 in the collection of P. S. Ward, University of California at Davis). This ant inhabits the stem internodes of at least 3 spp. of large bamboo, including Gigantochloa scortechinii Gamble (up to 25m tall, 10 cm diameter), the most common host plant at our main investigation site, the Ulu Gombak Field Studies Center near Kuala Lumpur. The large monogynous colonies may occupy dozens of internodes in different culms. Only internodes with small access holes are inhabited that can be blocked by one to a few workers to ward off potential intruders. During heavy rainstorms, runoff-water may flow into the internodes through the entrance holes. T. sp. responds with an effective countermeasure: The workers ingest the liquid and regurgitate it to the outside until no standing water remains. T. sp. apparently derives much of its food requirements from the honeydew of its symbiontic pseudococcids, Kermicus wroughtoni Newstead, which are kept inside the internodes. The close symbiosis of T. sp. with Kermicus becomes particularly apparent during the mating flight: The young queens carry Kermicus crawlers along in their mandibles for colony foundation. Unlike many other Pseudomyrmecinae, T. sp. does not exhibit predatory behavior but instead scavenges decaying plant matter, feces, and dead arthropods. There are also extended periods with no outside foraging. In contrast to other pseudomyrmecine plant specialists that reach comparable colony sizes, T. sp. is unaggressive and does not defend its host. Several species of spiders and the specialized staphylinid beetle Orphnebius politus are important predators of T. sp.
    
    
36. Klein, R. W., D. Kovac, Maschwitz, U., Buschinger, A. (1994). "Tetraponera sp. nahe attenuata F. Smith, eine sudostasiatische bambusameise (Hymen.: Formicidae: Pseudomyrmecinae), mit ungewohnlichen anpassungen an ihren lebensraum." Mitt. Dtsch. Ges. Allg. Angew. Entomol. 9: 337-341.
    
    
37. Klein, R. W., U. Maschwitz, et al. (1993). "Flood control by ants- a southeat asian bamboo-dwelling tetraponera (Formicidae, Pseudomyrmecinae) bails water from its internode nests." Insect. Soc. 40: 115-118.
    
    
38. Mehrotra, A. K. (1988). "Activity and feeding habit of Tetraponera (=Sima) rufonigra (Jerdon) (Formicidae: Pseudomyrmecinae: Hymenoptera) in outer Siwalik Hills." Ann. Entomol. 6: 57-58.
    
    
39. Mintzer, A. (1982). "Nestmate recognition and incompatibility between colonies of the acacia-ant Pseudomyrmex ferruginea [sic]." Behav. Ecol. Sociobiol. 10: 165-168.
    1. Compatibility between workers of 21 different colonies of the acacia-ant Pseudomyrmex ferruginea was examined. The colonies were reared from foundress queens in the greenhouse, on a clone of Acacia hindsii. Widespread incompatibility between colonies was encountered in these tests. Since diet and nesting environment are uniform, these results strongly suggest that the ants are producing recognition pheromones. 2. Worker brood from a stock colony was separated into groups of larvae and pupae, and each group was reared by a different foster reproductive female. The absence of antagonistic reactions between workers reared in the different groups indicates that the reproductive females are not the direct source of colony recognition pheromones.
    
    
40. Mintzer, A. and S. B. Vinson (1985). "Kinship and incompatibility between colonies of the acacia ant Pseudomyrmex ferruginea." Behav. Ecol. Sociobiol. 17: 75-78.
    The effect of kinship on incompatibility between colonies of the acacia-ant Pseudomrmex ferruginea was examined. Colonies were reared on a clone of Acacia hindsii in a standard environment. A slight but significant reduction in intercolony incompatibility was obtained with two inbred lineages, compared with the observed frequency of rejection for unrelated colonies. These results indicate that the relevant odor differences between colonies are probably not determined at a single gene locus. The 'gestalt' model for recognition pheromones does not apply to P. ferruginea, since nestmates are often treated differently upon introduction to a second colony. However, the resulting intermediate rejection frequencies (30-100%) fail to converge on the 50% level predicted by single locus models for recognition pheromones.
    
    
41. Mintzer, A. C. (1987). Experimental inbreeding, alate sex ratios and incompatibility between colonies of the Acacia ant Pseudomyrmex ferruginea. Chemistry and biology of social insects. J. Eder and H. Rembold. Munich, Peperny: 337-338.
    
    
42. Mintzer, A. C., H. J. Williams, et al. (1987). "Identity and variation of hexane soluble cuticular components produced by the acacia ant Pseudomyrmex ferruginea." Comp. Biochem. Physiol. B Comp. Biochem. 86: 27-30.
    
    
43. Morisita, M., M. Kubota, Onoyama, K., Ogata, K., Terayama, M., Kondoh, M., Imai, H.T. (1989). A guide for the identification of Japanese ants. I. Ponerinae, Cerapachyinae, Pseudomyrmecinae, Dorylinae, and Leptanillinae (Hymenoptera: Formicidae). [in Japanese]. Tokyo, Myrmecological Society of Japan.
    
    
44. Overal, W. L. (1982). Comparative study of social carrying in Pseudomyrmex and Ectatomma (Hymenoptera: Formicidae: Pseudomyrmecinae and Ponerinae). The Biology of social insects. Proceedings of the Ninth Congress of the International Union for the Study of Social Insects, Boulder, Colorado, August 1982. M. D. Breed, C. D. Michener and H. E. Evans. Boulder, CO. 420 p., Westview Press: 253.
    
    
45. Pan, J. and W. F. Hink (2000). "Isolation and characterization of myrmexins, six isoforms of venom proteins with anti-inflammatory activity from the tropical ant, Pseudomyrmex triplarinus." Toxicon 38: 1403-1413.
    
    
46. Pasteels, J. M., J. L. Boeve, et al. (1987). Chemical defense by contact poison in Tetraponera (Pseudomyrmecinae) and Crematogaster scutellaris (Myrmicinae). Chemistry and biology of social insects. J. Eder and H. Rembold. Mьnchen, Verlag J. Peperny: 423.
    
    
47. Perrault, G. H. (1988 ("1987")). "Notes sur des types de Pseudomyrmex dйcrits par F. Smith. I. (Hymenoptera, Formicidae)." Nouv. Rev. Entomol. (n.s.)4: 381-385.
    
    
48. Peters, J. M. (1997). "Microsatellite loci for Pseudomyrmex pallidus (Hymenoptera: Formicidae)." Mol. Ecol. 6: 887-888.
    
    
49. Radchenko, A. (1997). "Tetraponera pisarskii sp. nov. - the first native member of subfamily Pseudomyrmicinae from Palaearctic Asia." Ann. Zool., Warszawa 47: 479-480.
    
    
50. Reis, Y. T. and J. H. C. Delabie (1999). "Estudo de uma comunidade de Pseudomyrmecinae usando armadilhas de malaise numa reserva de Mata Atlвntica em Porto Seguro, Bahia." Naturalia 24: 119-121.
    
    
51. Seigler, D. S. and J. E. Ebinger (1995). "Taxonomic revision of the ant-acacias (Fabaceae: Mimosoideae, Acacia, series Gummiferae) of the New World." Ann. Mo. Bot. Gard. 82: 117-138.
    
    
52. Schultz, D. R. and P. I. Arnold (1977). "Characterization of factors in ant venom (Pseudomyrmex sp.) which cause inactivation of human complement proteins." Fed. Proc. 36: 1265.
    
    
53. Schultz, D. R. and P. I. Arnold (1977). "Venom of the ant Pseudomyrmex sp.: further characterization of two factors that affect human complement proteins." J. Immunol. 119: 1690-1699.
    
    
54. Schultz, D. R. and P. I. Arnold (1984). Immunochemical and clinical studies of venom from the ant Pseudomyrmex sp. Insect poisons, allergens, and other invertebrate venoms. A. T. Tu. N.Y., Marcel Dekker, Inc. 2: 243-290.
    
    
55. Smith, F. (1877). "Descriptions of new species of the genera Pseudomyrma and Tetraponera, belonging to the family Myrmicidae." Trans. Entomol. Soc. London 1877: 57-72. *[P. penetrator stat.rev by Ward, 1999; previously synonymized under P. concolor by Kempf 1967 and Ward 1989.] Taylor, B. (1976). Ants of the Nigerian forest zone (Hymenoptera: Formicidae). Part I Ponerinae, Cerapachyinae, Pseudomyrmecinae. An illustrated guide to worker caste, Cocoa Research Institute of Nigeria, Technical Bulletin No. 4: II, 41 p.
    
    
56. Terron, G. (1968). "Composition des colonies de Tetraponera anthracina Santschi (Hym. Formicoidea, Promyrmicidae)." Ann. Fac. Sci. Cameroun 1: 89-100.
    
    
57. Terron, G. (1968 ("1967")). "Description des castes de Tetraponera anthracina Santschi (Hym., Formicidae, Promyrmicinae)." Insect. Soc. 14: 339 -348.
    
    
58. Terron, G. (1969). "Description de Tetraponera ledouxi espece nouvelle du Cameroun, parasite temporaire de Tetraponera anthracina Santschi (Hym. Formicidae, Promyrmicinae)." Bull. Inst. Fondam. Afr. Noire (Ser. A, Sci. Nat.) 31: 629-642.
    Описание нового вида Tetraponera ledouxi из Камеруны, который является временным социальным паразитом вида Tetraponera anthracina Santschi
    
    
59. Terron, G. (1970). Recherches morphologiques et biologiques sur Tetraponera anthracina Santschi et sur son parasite social temporaire Tetraponera ledouxi Terron (Hym. Formicidae, Promyrmicinae), Ph.D. diss., Universite Paul Sabatier, Toulouse. 313 p. [Cited by Dejean and Passera, 1974.].
    
    
60. Terron, G. (1970 ("1969")). "Mise en evidence du parasitisme temporaire de Tetraponera anthracina Santschi par Tetraponera ledouxi nov. spec. (Hym. Formicidae, Promyrmicinae)." Ann. Fac. Sci. Univ. Fed. Cameroun 3: 113-115.
    
    
61. Terron, G. (1971). "Description des castes de Tetraponera nasuta Bernard (Hym. Formicidae, Promyrmicinae)." Ann. Fac. Sci. Univ. Fed. Cameroun 6: 73-84.
    
    
62. Terron, G. (1977). "Evolution des colonies de Tetraponera anthracina Santschi (Formicidae Pseudomyrmecinae) avec reines." Bull. Biol. Fr. Belg. 111: 115-181.
    Исследовано развитие колоний Tetraponera anthracina, содержащих маток.
    
    
63. Van Borm, S., A. Buschinger, Boomsma, J.J., Billen, J. (2002). "Tetraponera ants have gut symbionts related to nitrogen-fixing root-nodule bacteria." Proc. R. Soc. Lond. B Biol. Sci. 269: 2023-2027.
    
    
64. Ward, P. S. (1985). "The Nearctic species of the genus Pseudomyrmex (Hymenoptera: Formicidae)." Quaest. Entomol. 21: 209-246.
    Ревизия 10 Неарктических видов древесных муравьев рода Pseudomyrmex из 3 групп: gracilis group (P. mexicanus Roger), elongatus group (P. cubaensis Forel. stat. nov.; P. elongatus Mayr), and pallidus group (P. apache Creighton; P. bruneus F. Smith; P. ejectus F. Smith; P. leptosus Ward, sp. nov. [type locality: John Pennekamp State Park, Munroe Co., Florida]; P. simplex F. Smith). P. peruvianus Wheeler, stat. nov., described as a variety of P. ejectus, is considered a distinct species. The following are proposed as new synonyms of P. simplex: P. delicatulus Forel = P. capperi Forel = P. panamensis Forel = P. vittatus Forel: The Nearctic Pseudomyrmex are characterized by marked geographical variation, coupled with the frequent occurrence of sympatric sibling species.
    
    
65. Ward, P. S. (1989). "Systematic studies on pseudomyrmecine ants: revision of the Pseudomyrmex oculatus and P. subtilissimus species groups, with taxonomic comments on other species." Quaest. Entomol. 25: 393-468.
    Описаны новые виды: P. alustratus Ward (from Peru), P. cretus Ward (Costa Rica), P. pisinnus Ward (Brazil), P. spiculus Ward (Costa Rica), и P. villosus Ward (Brazil). Таксон P. caeciliae (Forel) повышен до вида (stat. nov.) из расы P. eduardi race caeciliae. Новая синонимия... P. alliodorae (Wheeler) is a n.syn. of P. elongatus (Mayr), P. altinodus (Mann). P. tuberculatus (Enzmann), P. wessoni (Enzmann) are n.syn. of P. oculatus (F. Smith). P. chodati (Forel) & P. ogloblii (Santschi) are n.syn. of P. urbanus (F. Smith). P. culmicola (Forel) is a n.syn. of P. tenuissimus (Emery). The following are raised to species (stat. nov.) for P. pazosi (Santschi) from Pseudomyrma pazosi, P. rufiventris (Forel) from P. kurokii var. rufiventris, and P. tachigaliae (Forel) from P. latinoda tachigaliae. P. dendroicus (Forel) & P. osurus (Forel) are removed from synonymy (stat. reval.). A replacement name (P. laevifrons Ward. nom. nov. (P. laeviceps) is provided for P. laeviceps (F. Smith), 1877 (nec F. Smith, 1859).]
    
    
66. Ward, P. S. (1990). "The ant subfamily Pseudomyrmecinae (Hymenoptera: Formicidae): generic revision and relationship to other formicids." Syst. Entomol. 15: 449-489.
    Родовая ревизия всего подсемейства на родовом уровне. Из Бразилии описан новый вид муравьев Myrcidris epicharis n.sp. Новые синонимы: Ornatinoda Enzmann, Clavanoda Enzmann, Triangulinoda Enzmann, Apendunculata Enzmann, Latinoda Enzmann are n.syn. of Pseudomyrmex Lund. Родовые таксоны Pachysima Emery, Viticicola Wheeler, Sima Emery (nec Roger), Parasima Donisthorpe (все как n.syn.) сведены в синонимы к роду Tetraponera F. Smith.]
    
    
67. Ward, P. S. (1991). Phylogenetic analysis of pseudomyrmecine ants associated with domatia-bearing plants. Ant-plant interactions. C. R. Huxley and D. F. Cutler. Oxford. xviii + 601 p., Oxford University Press: 335-352.
    Филогенетический анализ древесных муравьев, мутуалистически ассоциированных с растениями. Основные части работы: a) Phylogenetic relationships among the Pseudomyrmecinae. b) Nesting behaviour and plant associations of the pseudomyrmecine ants. c) Discussion: (Patterns of association) (Origins of pseudomyrmecine-plant relationships) (Characteristics of obligate plant-ants) d) Conclusions (in part): Phylogenetic analysis of the Pseudomyrmecinae indicates that obligate plant-ants, which comprise almost 20 % of the estimated 250-300 extant species in the subfamily, have arisen at least twelve times.
    
    
68. Ward, P. S. (1992). "Ants of the genus Pseudomyrmex (Hymenoptera: Formicidae) from Dominican amber, with a synopsis of the extant Antillean species." Psyche 99: 55-85.
    
    
69. Ward, P. S. (1993). "Systematic studies on Pseudomyrmex acacia-ants (Hymenoptera: Formicidae: Pseudomyrmecinae)." J. Hym. Res. 2: 117- 168.
    Синонимия акациевых муравьев. [Extensive errata sheet available from author OR Errata in J. Hym. Res. 3:309-310. (1994.10.15). P. alfari, P. bicinctus, P. nigrocincta bicincta, & P. peltatus are n.syn. of P. ferrugineus. P. atrox, P. spinicola atrox, P. gaigei, P. spinicola gaigei, P. infernalis, P. spinicola infernalis, P. scelerosus, & P. spinicola scelerosa are n.syn. of P. spinicola. P. thoracicus & P. thoracica are n.syn. of P. boopis. P. bicolor, P. canescens, P. dimidiata, P. dimidiatus, P. gracilis mexicana guayaquilensis, P. glabriventris, P. gracilis glabriventris, P. mexicana, P. mexicanus, P. pilosula, P. pilosulus, P. sericata, P. sericatus, P. gracilis velifera, P. veliferus, &, P. volatilis are n.syn. of P. gracilis. P. acaciarum & P. sericea acaciarum are n.syn. of P. ita (stat.nov.). P. gracilis major is a n.syn. of P. major (stat.nov.). Left unresolved is the relationship to P. gracilis to: P. alternans, P. gracilis argentinus, P. gracilis atrinodus, & P. santschii.]
    
    
70. Ward, P. S. (1996). "A new workerless social parasite in the ant genus Pseudomyrmex (Hymenoptera: Formicidae), with a discussion of the origin of social parasitism in ants." Syst. Entomol. 21: 253-263.
    Описан паразитический вид P. inquilinus sp. nov. из Аргентины
    
    
71. Ward, P. S. (1999). "Systematics, biogeography and host plant associations of the Pseudomyrmex viduus group (Hymenoptera: Formicidae), Triplaris- and Tachigali-inhabiting ants." Zool. J. Linn. Soc. 126: 451-540.
    Систематика группы видов Pseudomyrmex viduus group. Описаны новые виды: P. crudelis, P. deminutus, P. eculeus, P. ferox, P. hospitalis, P. insuavis, P. ultrix, & P. vitabilis are n.sp... Многочисленная новая синонимия: P. latinodus, P. endophytus syn. nov., & P. damnosus are syn. of P. concolor... P. emarginatus is a syn. of P. dendroicus... P. dichrous, P. bierigi, & P. crenulatus are syn. of P. kuenckeli... P. cholericus, P. crucians, & P. auripes are syn. of P. malignus... P. symbioticus is a n.syn. of P. mordax, n.comb., stat.rev... P. nigrescens n.syn., P. coronatus Wheeler syn. nov., & P. coronatus Enzmann are syn. of P. penetrator, stat.rev... P. bradleyi Wheeler & P. bradleyi Enzmann are syn. of P. tachigaliae... P. boxi Wheeler & P. boxi Enzmann are syn. of P. triplaridis... P. arborissanctae, P. cordobensis syn.nov., P. rurrenabaquensis syn.nov., & P. ecuadorianus are syn. of P. triplarinus... P. caroli, P. opacior syn.nov., P. sapii, P. ulei, P. baileyi, P. tigrinus, P. biolleyi, & P. trigonus are syn. of P. viduus... Pseudomyrmex fortis stat.nov... P. acaciarum Wheeler & P. acaciorum Enzmann are syn. of P. ita... P. heterogyna syn. nov., P. castus syn. nov., & P. humboldi syn. nov. are syn. of P. pictus... P. huberi is a syn. of P. rubiginosus... P. vinneni stat.nov... В определительные таблицы включены виды P. viduus group и все остальные виды рода Pseudomyrmex, обнаруженные на растениях Triplaris или Tachigali.
    
    
72. Ward, P. S. (2001). "Taxonomy, phylogeny and biogeography of the ant genus Tetraponera (Hymenoptera: Formicidae) in the Oriental and Australian regions." Invertebr. Taxon. 15: 589-665.
    
    
73. Wu, J. and C. Wang (1990). "A taxonomic study on the genus Tetraponera Smith in China (Hymenoptera: Formicidae). [In Chinese]." Sci. Silvae Sin. 26: 515-518.
    Tetraponera microcarpa described from China. Tetraponera nigra new record for China. Other species only in chinese key.
    
    
74. Wheeler, G. C. and J. Wheeler (1956). "The ant larvae of the subfamily Pseudomyrmecinae (Hymenoptera: Formicidae)." Ann. Entomol. Soc. Am. 49: 374-398.
    Первое подробное описание личинок древесных муравьев Pseudomyrmecinae
    
    
75. Wheeler, G. C. and J. Wheeler (1973). "Supplementary studies on ant larvae: Cerapachyinae, Pseudomyrmecinae and Myrmicinae." Psyche 80: 204-211.
    
    

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