1. <strong>LABOULBENIALES</strong>1.1. DEFINITION OF <strong>LABOULBENIALES</strong>Laboulb<strong>en</strong>iales include over 2000 species in 141 g<strong>en</strong>era (SANTAMARIA, 1998; KIRKet al., 2001; WEIR & BLACKWELL, 2005) of obligate, ectoparasitic fungi, which liveassociated with arthropods, mostly true insects. Laboulb<strong>en</strong>iales have nomycelium; their thalli are small and of determinate growth, bearing antheridiaand perithecia on a receptacle with app<strong>en</strong>dages (TAVARES, 1985). Only sexualstages are known. Laboulb<strong>en</strong>iales are host-specific (THAXTER, 1896; SCHELOSKE,1969; TAVARES, 1985; MAJEWSKI, 1994; DE KESEL, 1996, 1997).The knowledge on Laboulb<strong>en</strong>iales is rather rec<strong>en</strong>t and has increased slowly.Large parts of their biodiversity are still unknown and many questions stillunresolved.1.2. HISTORICAL BACKGROUNDIn the 1840s, two Fr<strong>en</strong>ch <strong>en</strong>tomologists, Joseph Alexandre Laboulbéne andAuguste Rouget, did the earliest observations on Laboulb<strong>en</strong>iales resulting in afirst publication on the Laboulb<strong>en</strong>iales (ROUGET, 1850, ref. in TAVARES, 1985). Atfirst, the structures on Brachinus (Coleoptera, Carabidae) were thought to besupernumerary ant<strong>en</strong>nal segm<strong>en</strong>ts. ROUGET (1850, ref. in TAVARES, 1985)suggested that they were living organisms. The recognition as fungi came in1852 and the g<strong>en</strong>us Laboulb<strong>en</strong>ia was raised within the Pyr<strong>en</strong>omycetes – anext<strong>en</strong>sive group of Ascomycetes (ROBIN, 1853). Later on, specim<strong>en</strong>s onwingless dipteran parasites of bats were still mistak<strong>en</strong>ly described asArthrorhynchus, an acanthocephalan worm, by KOLENATI (1857).MAYR (1853) thought that the „hairlike structures‟ on Nebria (Coleoptera,Carabidae) were chitinous. Suggesting that they were outgrowths of theinsect‟s integum<strong>en</strong>t, he noticed differ<strong>en</strong>ces betwe<strong>en</strong> those structures onyounger and older beetles. On older individuals, there was always a variableswelling on the hair, whereas no such swelling was pres<strong>en</strong>t on the hair ofyounger beetles.PEYRITSCH (1871) made ext<strong>en</strong>sive observations on the structure anddevelopm<strong>en</strong>t of the housefly Laboulb<strong>en</strong>ia (L. muscae Peyr.). PEYRITSCH (1871)redescribed Arthrorhynchus as Laboulb<strong>en</strong>ia nycteribiae Peyr. He laterestablished the family Laboulb<strong>en</strong>iaceae in the Ascomycetes (PEYRITSCH, 1873)and laid a solid foundation for our understanding of the biology ofLaboulb<strong>en</strong>iales (PEYRITSCH, 1875), based on observations of laboratory coloniesof houseflies and studies of Laboulb<strong>en</strong>iales in the field.A systematic study of the Laboulb<strong>en</strong>iaceae was begun by Roland Thaxter in1890, wh<strong>en</strong> he published his first in a series of papers describing hundreds ofnew species. In 1896, THAXTER‟S first monographic volume on theLaboulb<strong>en</strong>iaceae appeared. It was particularly a summary upon all previouswork (TAVARES, 1985). The monograph of Thaxter – five volumes, published in1896, 1908, 1924, 1926 and 1931 – constituted the basis for later studies of thegroup. Unfortunately Thaxter died only one year after the publication of theP a g e | 9
fifth volume, therefore the sixth volume – which was meant to be a synthesis –was never prepared. More than 40 years study resulted in the description of103 g<strong>en</strong>era, approximately 1260 species and 13 varieties (BENJAMIN, 1971). Upto now, about two thirds of all the known species have be<strong>en</strong> described byThaxter (HULDÉN, 1983).THAXTER (1896) first separated the family of the Laboulb<strong>en</strong>iaceae into two„groups‟ (Table I below): the Endog<strong>en</strong>ae and the Exog<strong>en</strong>ae (based on thedevelopm<strong>en</strong>t of spermatia in the antheridia). The Endog<strong>en</strong>ae included twoorders: the Laboulb<strong>en</strong>ieae (with simple antheridia; 15 g<strong>en</strong>era) and thePeyritschielleae (with compound antheridia; 11 g<strong>en</strong>era). Those orders wouldcorrespond to tribes in modern classification systems. In the second volumeTHAXTER (1908) replaced the terms Endog<strong>en</strong>ae and Exog<strong>en</strong>ae by the subordinalnames Laboulb<strong>en</strong>iinae and Ceratomycetinae (Table I below); the majorsubdivisions from the former group Endog<strong>en</strong>ae were substituted by families,Laboulb<strong>en</strong>iaceae and Peyritschiellaceae. Thaxter did not recognize aseparate family within the Ceratomycetinae. The name Ceratomycetaceae –now universally accepted – first has be<strong>en</strong> introduced by MAIRE in 1916.Volume I Family Laboulb<strong>en</strong>iaceaeTable I: Classification in Thaxter‟s monographs I and II„Group‟ Endog<strong>en</strong>ae Exog<strong>en</strong>aeOrder Laboulb<strong>en</strong>ieae PeyritschielleaeVolume II Order Laboulb<strong>en</strong>ialesSuborder Laboulb<strong>en</strong>iinae CeratomycetinaeFamily Laboulb<strong>en</strong>iaceae PeyritschiellaceaeP a g e | 10After Thaxter‟s death in 1932, various authors contributed to the knowledge bypublishing some short papers and/or describing new species and g<strong>en</strong>era:Spegazzini, Picard, Maire, Chatton, Cépède, Giard and Istvánffi. The majorityof the numerous publications that have appeared were regional studies,collection reports and rather short (TAVARES, 1985; DE KESEL, 1997). However,wh<strong>en</strong> considered together, they have added ext<strong>en</strong>sive data on thedistribution of these fungi.Some researchers suggested a relationship betwe<strong>en</strong> red algae andLaboulb<strong>en</strong>iales and thought that Dikaryomycota derived from red algae, withthe Laboulb<strong>en</strong>iales as an intermediate step KARSTEN, 1869; SACHS, 1874). Sexualreproductive structures, including a highly differ<strong>en</strong>tiated trichogyne and theabs<strong>en</strong>ce of mycelium, are unusual ascomycete characteristics andsuperficially resemble floridean algal features (BLACKWELL, 1994). CÉPÈDE (1914)believed the Laboulb<strong>en</strong>iales as a whole occupied a special place among thefungi; he placed them in the Phycascomycetes, a name he proposedbecause of the superficial similarities both to the Ascomycetes and to theRhodophyta (red algae). Until g<strong>en</strong>eticists unquestionably proved thatAscomycetes derived from Chytridiomycetes-like ancestors, this “Red Algal (orFloridean) Hypothesis” persisted (JAMES et al., 2006). The analysis of JAMES et al.(2006) suggests that several clades of the Chytridiomycota (true fungi, stillproducing zoospores) form a paraphyletic assemblage at the base of the
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Table XXIV: Family Cerambycidae fro
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Table XXVIII: Family Dermestidae fr
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Adults prey on other flower-visitin
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Table XXXV: Family Scarabaeidae fro
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Creophilus Samouellemaxillosus (Lin
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4.3. LABOULBENIALES OF CARRION BEET
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The knowledge about the preferences
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Table XLII: Comparison of carrion b
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5. CONCLUSION AND SUGGESTIONSNo spe
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1. INTRODUCTION1.1. NATURAL LANDSCA
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In the 40s, Middelhoek studied and
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2. MATERIALS & METHODS2.1. HOSTS2.1
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Screening of the insects was done w
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3. RESULTS3.1. PARASITE-HOST LISTA
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Figure XV: Detail of thallus of Lab
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isodiametric or slightly elongate,
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occurs in the lower receptacle (cel
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Discussion:Two species of the genus
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anches (15-)34-38(-40) µm long, te
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5. CONCLUSION AND SUGGESTIONSIn The
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P a g e | 100ADRIAENS, T. & GYSELS,
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transmission, habitat preference an
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STALPERS, J.A., VILGALYS, R., AIME,
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MELIS, C., TEURLINGS, I. LINNELL, J
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SCHILTHUIZEN, M. & VALLENDUUK, H. (