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Diversity of Fungi from the Mound Nests of Formica Ulkei and Adjacent Non-Nest Soils

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Canadian Journal of Microbiology Diversity of fungi from the mound nests of Formica ulkei and adjacent non-nest soils Journal: Canadian Journal of Microbiology Manuscript ID cjm-2015-0628.R2 Manuscript Type: Article Date Submitted by the Author: 25-Feb-2016 Complete List of Authors: Duff, Lyndon B.; Brandon University, Biology Urichuk, Theresa M.; Brandon University, Biology Hodgins, Lisa N.; Brandon University, Biology Young, JocelynDraft R.; Brandon University, Biology Untereiner, Wendy; Brandon University, Biology Keyword: Aspergillus, fungal biodiversity, xerotolerant, mound-building ant https://mc06.manuscriptcentral.com/cjm-pubs Page 1 of 29 Canadian Journal of Microbiology Duff et al.; Fungi from nests of Formica ulkei 1 1 Diversity of fungi from the mound nests of Formica ulkei and adjacent non-nest soils 2 3 Lyndon B. Duff, Theresa M. Urichuk, Lisa N. Hodgins, Jocelyn R. Young, and Wendy A. 4 Untereiner 1 5 Department of Biology, Brandon University, 270 18 th Street, Brandon, Manitoba, R7A 6A9, 6 Canada 7 8 L.B. Duff ([email protected]) 9 T.M. Urichuk ([email protected]) 10 L.N. Hodgins ([email protected]) 11 J.R. Young ([email protected])Draft 12 W.A. Untereiner ([email protected]) 13 14 15 16 17 18 19 20 21 22 1 Corresponding author https://mc06.manuscriptcentral.com/cjm-pubs Canadian Journal of Microbiology Page 2 of 29 Duff et al.; Fungi from nests of Formica ulkei 2 23 Abstract 24 Culture-based methods were employed to recover 3929 isolates of fungi from soils collected 25 in May and July 2014 from mound nests of Formica ulkei and adjacent non-nest sites. The 26 abundance, diversity, and richness of species from nest mounds exceeded those of non- 27 mound soils, particularly in July. Communities of fungi from mounds were more similar to 28 those from mounds than non-mounds; this was also the case for non-mound soils with the 29 exception of one non-mound site in July. Species of Aspergillus , Paecilomyces and 30 Penicillium were dominant in nest soils and represented up to 81.8% of the taxa recovered. 31 Members of the genus Aspergillus accounted for the majority of Trichocomaceae from nests 32 and were represented almost exclusively by Aspergillus navahoensis and A. pseudodeflectus . 33 Dominant fungi from non-mound sites Draftincluded Cladosporium cladosporioides , Geomyces 34 pannorum and species of Acremonium , Fusarium , Penicillium and Phoma. Although mound 35 nests were warmer than adjacent soils, the dominance of xerotolerant Aspergillus in soils 36 from mounds and the isolation of the majority of Trichocomaceae at 25˚C and 35˚C suggests 37 that both temperature and water availability may be determinants of fungal community 38 structure in nests of F. ulkei . 39 40 41 Key words: Aspergillus , fungal biodiversity, mound-building ant, xerotolerant 42 43 44 45 46 47 https://mc06.manuscriptcentral.com/cjm-pubs Page 3 of 29 Canadian Journal of Microbiology Duff et al.; Fungi from nests of Formica ulkei 3 48 Introduction 49 The mound-building ant Formica ulkei Emery (Hymenoptera: Formicidae) ranges from Alberta 50 to Nova Scotia (Canada) and southward to Illinois, Indiana and Iowa (USA) (Holmquist 1928; 51 Sherba 1958; Glasier et al. 2013). This species builds conspicuous nests in meadows and 52 pastures along the margins of forests and sparsely wooded areas (Holmquist 1928; Dreyer 53 and Park 1932; Sherba 1958). Nests are composed of excavated soil and covered by a layer 54 of thatch (i.e., small pieces of grass and other plant material) (Sherba 1958, 1959, 1962). 55 The mound nests of F. ulkei are thermoregulatory in function and are constructed to 56 achieve and maintain higher temperatures than adjacent undisturbed soils during the months 57 when the ants are most active (Sherba 1962). Nests are built in exposed sites and are 58 oriented to maximize their exposure toDraft solar radiation (Sherba 1958); they gain heat from 59 solar radiation in the early spring and maintain temperatures that are higher and more stable 60 than those of surrounding soils because of the insulating properties of thatch (Sherba 1962; 61 Frouz and Jilková 2008). This layer of organic material prevents the overheating of mounds 62 during the warmest parts of the year in other ant species that construct thatched nests 63 (Bollazzi and Rocces 2010; Kadochová and Frouz 2014) and it may serve the same function 64 in F. ulkei . 65 Although it is recognized that mound-building ants are capable of dramatically modifying 66 their environments and altering the chemical and physical properties of soils (Beattie and 67 Culver 1977; Frouz and Jilková 2008; Jilková et al. 2011), few studies have explored the 68 impact of microclimatic conditions on the composition of the communities of fungi in these 69 soils (Ba et al. 2000; Zettler et al. 2002; Rodrigues et al. 2014). Given the availability of a 70 large group of nests of F. ulkei in south-eastern Manitoba, we undertook a study to 1) confirm 71 the temperature characteristics of the mound nests of this species reported in previous 72 studies, and 2) test the hypothesis that the community of culturable fungi from soils from https://mc06.manuscriptcentral.com/cjm-pubs Canadian Journal of Microbiology Page 4 of 29 Duff et al.; Fungi from nests of Formica ulkei 4 73 nests differs from adjacent, non-nest soils. We were also interested in comparing the species 74 richness and diversity of the communities of culturable fungi of separate mound nests of F. 75 ulkei . 76 77 Materials and Methods 78 Collection of soils and temperature data 79 Thermocron iButton data loggers (DS1921G, Maxim Integrated Products, San Jose, USA) 80 that had been pre-set to measure temperature every two hours were coated in Performix 81 Plasti Dip (Plasti Dip International, Blaine, USA) to prevent moisture damage (Roznik and 82 Alford 2012). Data loggers were buried 5 cm deep in soil on the top, south side and north side 83 of three mound nests of Formica ulkei Draftlocated on the un-forested edge of a cattle pasture that 84 had not been grazed in approximately 10 years, south of White Mud Falls, Manitoba (UTM 85 coordinates of mound 1 = 14U 0707355 5588945; mound 2 = 14U 0707363 5588913; mound 86 3 = 14U 0707367 5588908). One data logger was buried at a depth of 5 cm at one location 1 87 m south of each mound. Another data logger was also secured at a height of 2 m to the north 88 (i.e., the shaded) side of a tree located in the middle of the study area to collect air 89 temperatures. Data loggers recorded temperatures from 9 May to 18 September 2014. 90 Nests were sampled on 11 May and 14 July 2014 by collecting the uppermost 3 cm of 91 soil beneath the thatch from the top and south sides of each mound. Each site on all mounds 92 was sampled using a new plastic spoon. Soils to a depth of 3 cm were collected from 93 adjacent non-mound soil 1 m south of nests using a soil core sampler that was sterilized in 94 100% ethanol and rinsed in sterile distilled water between samples. Samples were placed into 95 separate, unused plastic freezer bags, sealed and transported in an ice cooler to the 96 laboratory. Each sample was emptied into a clean aluminum pan, air-dried at room https://mc06.manuscriptcentral.com/cjm-pubs Page 5 of 29 Canadian Journal of Microbiology Duff et al.; Fungi from nests of Formica ulkei 5 97 temperature (18-21˚C), subjected to sieving using a 2 mm mesh to remove plant debris, and 98 stored in a new freezer bag. 99 100 Isolation and identification of fungi 101 Individual soil samples were used within 3 days following collection to prepare ten-fold serial 102 dilutions in sterile distilled water ranging from 10 -1 to 10 -7. Each dilution was plated in triplicate 103 on Dextrose-Peptone-Yeast Extract agar (DPYA) (Papavizas and Davey 1959) lacking oxgall 104 and sodium propionate, and Dichloran Rose Bengal agar (DRBA) (King et al. 1979) 105 containing 25 mg Rose Bengal, 2 mg dichloran, and KH 2PO 4 rather than K 2HPO 4. Both media 106 were supplemented with 50 mg chlortetracycline hydrochloride and 50 mg streptomycin 107 sulphate. Duplicate sets of plates wereDraft incubated at 25˚C and 35˚C for 5 days. 108 All fungal colonies were transferred to Modified Leonian’s agar (MLA) (Malloch 1981), 109 incubated at room temperature and identified based on cultural and micro-morphological 110 characters. Isolates that could be discriminated as separate taxa within genera but not 111 identified to species were numbered. Sporulating fungi that could not be identified to the level 112 of genus were designated as “undetermined” whereas those taxa that did not sporulate on 113 MLA were labeled “sterile” (see supplemental Table S1). Non-filamentous fungi and 114 Zygomycota, which were isolated in very low numbers on both DRBA and DPYA, were 115 disregarded. Fungi recovered on DPYE were also excluded from analyses because of the 116 high levels of bacterial contamination, particularly in soils collected in July. 117 Dominant species of Aspergillus were characterized on Czapek Dox agar (CZ), Czapek 118 Yeast agar (CYA), Czapek Yeast agar with 20% sucrose (CY20S) and Malt Extract agar 119 following Klich (2002a) and on Creatine Sucrose agar (CREA) as described by Samson et al. 120 (2014). The thermotolerances of these taxa were determined by assessing their ability to 121 grow on CYA and MLA when incubated at 37˚C, 45˚C, and 50˚C. Cultures used for DNA https://mc06.manuscriptcentral.com/cjm-pubs Canadian Journal of Microbiology Page 6 of 29 Duff et al.; Fungi from nests of Formica ulkei 6 122 extraction were grown as described previously (Untereiner et al.
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  • 2009-Acclimation.Pdf

    2009-Acclimation.Pdf

    16 Acclimation Douglas W. Whitman 4120 Department of Biological Sciences, Illinois State University, Normal, IL, 61790 USA, Phone: (309) 438-5123. e-mail: [email protected] Abstract Acclimation refers to a physiological change in an individual stimulated by exposure to a different, often stressful, environment. As such it represents physiological phenotypic plasticity. This chapter reviews both early (1900 – 1960) and current research on arthropod acclimation, including: definitions, abiotic and biotic elicitors, types of acclimatory responses, tolerance and capacity acclimation, persistence and speed of response, confounding factors, including different experimental designs and metrics, graphic models, underlying physiological mechanisms, and possible adaptive value. Current acclimation research emphasizes molecular biology, environment- induced gene activation, passive vs. active responses, ecological and fitness consequence of acclimation, and its costs, adaptiveness, and evolution. Current studies attempt to integrate acclimation from genes-to-ecology, and relate acclimation to homeostatic physiology, phenotypic plasticity and stress studies. Understanding acclimation has numerous practical benefits. Everything old is new again Introduction The current literature on phenotypic plasticity often proclaims the novelty of this exciting research area. However, like many fields of science, phenotypic plasticity actually has a long and diverse history, some of which has been nearly forgotten. During the early and mid 20th Century, as geneticists and evolutionary biologists worked to develop the initial ideas about phenotypic plasticity (Baldwin 1896, 1902, Morgan 1896a,b, Osborn 1897, Woltereck 1909, Johannsen 1911, Nilsson-Ehle 1914, Dobzhansky 1937, Clausen et al. 1940, Goldschmidt 1940, Waddington 1942, Schmalhausen 1949, 676 Phenotypic Plasticity of Insects Bradshaw 1965), another group of scientists labored, largely beyond their view, on a sub-discipline of phenotypic plasticity: acclimation and acclimatization.