Further notes on European taxa of the lichen genus Leptogium,with emphasis on the small species

Abstract:Leptogium imbricatum P.M. Jørg. sp. nov. and L. intermedium (Arnold) Arnold are established as the correct names for two fairly widespread, small species: L. imbricatum has an arctic-alpine distribution. Leptogium intermedium was previously mostly named L. minutissimum (Flörke) Fr., the type of which is L. subtile (Schrader) Torss., a distinct species usually growing on rotting wood or debris. Leptogium magnussonii Degel. & P.M. Jørg, mainly from maritime rock outcrops in Norway and Sweden, is described as new. The species complexes of L. plicatile (Ach.) Leighton and L. schraderi (Bernh.) Nyl. are discussed. The differences between L. byssinum (Hoffm.) Nyl. and L. biatorinum (Nyl.) Leighton are given, and L. aquale (Arnold) P.M. Jørg. comb. nov., a taxon related to the latter, is considered to be a distinct species growing in mountain brooks in Central Europe. Leptogium rivale Tuck. is recorded as new to Europe, from Romania. Leptogium bellopratense Hy is shown to be a synonym of L. brebissonii Mont. New records for L. coralloideum (Meyen & Flotow) Vainio, L. massiliense Nyl and L. rivulare (Ach.) Mont. are given. A preliminary key to the small European species (north of the Alps) is appended. The following names have been lecto- or neotypified: Collema byssinum Hoffm., Collema pulvinatum Hoffm., Leptogium bellopratense Hy., Leptogium crenulatum Watson, Leptogium massiliense Nyl., Leptogium minutissimum var. intermedium Arnold, Leptogium minutissimum f. plumbeum Zwackh ex Arnold, Leptogium pusillum var. aquale Arnold, Leptogium rivulare var. crenatulum Nyl., Leptogium scotinum var. crenatum Nyl., Lichen byssinus Hoffm., Lichen minutissimus Flörke, Lichen schraderi Bemh., Lichen subtilis Schrader and Parmelia scotina var. lophaea Ach.     

石耳科(子囊菌亚门)的间断分布和替代现象

本文报道了石耳科的两个亚洲的,也是美洲以外的新记录种,即角石耳与深色石耳。进一步证实了单果石耳在南大西洋的,即南美洲和南部非洲的间断分布。对于上述种类以及本科其它一些种类的间断分布和替代现象进行了分析讨论。     

新疆博格达山岩面生地衣群落结构特征

根据对新疆博格达山岩面生地衣群落20个样点(20m×20m)调查的数据,以各地衣种的盖度为指标结合双向指示种分析方法(TWINSPAN)和除趋势对应分析法(DCA)对博格达山岩面生地衣群落进行数量分类并分析了群落结构特征及其多样性和相似性。采用典范对应分析法(CCA)对各群落的物种分布格局与环境因子的关系进行研究。结果表明,TWINSPAN分析和DCA排序将分布在博格达山的37种岩面生地衣分为以下5个群丛。群丛1:斑纹网衣(Lecidea tessellate Florke)+粉芽盾衣(Peltula euploca(Ach.)Poelt)+杜瑞氏黄梅(Xanthoparmelia durietzii Hale)群丛,有25个种,总覆盖度为30.145%,多样性为4.025;群丛2:袋衣(Hypogymnia physodes(L.)Nyl.)+白边平茶渍(Aspicilia sublaqueata(H.Magn.)J.C.Wei)+砖孢胶衣(Collema subconveniens Nyl.)群丛,有17个种,地衣总盖度为15.885%,多样性为3.196;群丛3:聚茶渍(Lecanora accumulate H.Magn.)+丽石黄衣(Xanthoria elegans(Link)Th.Fr.)+亚洲平茶渍(Aspicilia asiatica(H.Magn.)Yoshim.)群丛,有30个种,地衣总盖度为37.87%,多样性为4.357;群丛4:中华石果衣(Endocarpon sinense H.Magn.)+伴藓大孢蜈蚣衣(Physconia muscigena(Ach.)Poelt.)+垫脐鳞衣(Rhizoplaca melanophthalma(DC.)LeuckertPoelt)群丛,有24个种,地衣总盖度为30.458%,多样性为3.912;群丛5:石胶衣(Collema flaccidum(Ach.)Ach.)+短绒皮果衣(Dermatocarpon vellereum Zschacke)+绿黑地图衣(Rhizocarpon viridiatrum(Wulfen)Korber.)群丛,有18个种,地衣总盖度为19.331%,多样性为3.515。CCA排序结果反映,该地区岩面生地衣的分布与海拔高度、光照强度、岩石pH和人为干扰有关,其中影响最大的因素是海拔高度,其次为光照强度和干扰。坡向和岩石大小对地衣种类分布的影响不显著。     

乌鲁木齐南部山区森林生态系统树生地衣群落结构

为了探明乌鲁木齐南部山区树生地衣种类及其群落结构特征, 应用多元分析中的主成分分析及聚类分析方法, 对分布在乌鲁木齐南部山区森林生态系统中的树生地衣植物群落进行数量分类, 并对其群落结构的物种多样性、相似性和均匀度等群落参数进行了比较系统的研究。结果表明, 分布在乌鲁木齐南部山区的树生地衣共有39种, 隶属于5目13科26属。根据多元分析结果, 将该地区的树生地衣划分为3种类型: (1) 小茶渍(Lecanora hageni (Ach.) Ach.) + 蜈蚣衣(Physcia stellaris (L.) Nyl.) + 柳茶渍(L. saligna (Schrad.) Zahlbr.)群落; (2) 斑面蜈蚣衣(Physcia aipolia (Humb.) Furm.) + 对开蜈蚣衣(Ph. dimidiata (Arn.) Nyl.) + 喇叭石蕊(Cladonia pyxidata (L.) Hoffm.)群落; (3) 拟石黄衣(Xanthoria fallax (Hepp) Arnold.) + 丽石黄衣(X. elegans (Link.) Th. Fr.)群落。群落3的多样性指数最大, 为1.509, 其次为群落2, 其多样性指数为1.109, 群落1的多样性指数最低, 为1.088。同时, 研究发现海拔高度和树种是影响乌鲁木齐南部山区森林生态系统树生地衣结构的两个重要因素。     

Selectivity of photobiont choice in a defined lichen community: inferences from cultural and molecular studies

Axenic cultures of lichen photobionts isolated from bark-inhabiting lichen thalli of the Physcietum adscendentis Ochsner were identified by light microscopy and sequence comparisons of internal transcribed spacer rDNAs to investigate principles of lichenization within a defined lichen sociological unit. The photobiont identity of eight lichen species is reported for the first time (photobiont species in square brackets): Lecania cyrtella (Ach.) Th. Fr. [ Trebouxia arboricola Puym.], Lecania naegelii (Hepp) Diederich & v. d. Boom [ Dictyochloropsis symbiontica Tscherm.-Woess], Candelaria concolor (Dicks) B. Stein [ Trebouxia jamesii (Hildreth & Ahmadjian) Gärtner], Candelariella cf. reflexa (Nyl.) Lettau [ T. jamesii ], Lecanora spec. [ T. arboricola ], Phaeophyscia orbicularis (Neck.) Moberg [ T. impressa Ahmadjian], Physcia adscendens (Fr.) H. Olivier [ T. impressa ] and Lecidella elaeochroma (Ach.) M. Choisy [ T. arboricola ] and could be confirmed for another two species, Physcia stellaris (L.) Nyl. [ Trebouxia impressa ] and Xanthoria parietina (L.) Th. Fr. [ Trebouxia arboricola ]. The observation that pioneer lichens without vegetative propagules, growing on smooth bark, had Trebouxia arboricola as photobiont can be explained by the assumption of a free-living population of Trebouxia arboricola . Species of photobionts from Xanthoria parietina were morphologically and genetically different from those of Physcia adscendens and Phaeophyscia orbicularis , respectively; a finding that does not support the previous assumption that Xanthoria parietina takes over its algal partner from a Physcia species, at least at the sites investigated.     

An in vitro method for establishing mycorrhizae on coniferous tree seedlings

Summary A method for in vitro synthesis of mycorrhizae on coniferous tree seedlings is described. Tree seedlings (Larix decidua Mill., Picea abies (L.) Karst, and Pinus sylvestris L.) and fungi Amanita muscaria (L. ex Fr.) Hooker, Piloderma croceum Erikss. et Hjorst., Pisolithus tinctorius (Pers.) Coker et Couch, and Suillus grevillei (Klotzsch) Singer were maintained under sterile conditions in petri dishes. Typical ectomycorrhizae were established within 2–3 weeks after inoculation and within 2 months after germination of seedlings. Eventually a high percentage of mycorrhizal root tips was obtained.     

62. G. Follmann: Chilenische Wanderflechten

• 1 . Es wird erstmalig über das Vorkommen von Wanderflechten in Südamerika berichtet
• 2 . Als solche kommen die Laub- und Strauchflechten Parmelia vagans (Nyl.) Nyl., Roccella cervicornis Follm. spec. nov. und Tornabenia ephebaea (Ach.) Kur. in der nordchilenischen Atacamawüste vor
• 3 . Geographie, Ökologie, Soziologie und Systematik der chilenischen Ärolichenen werden erörtert
• 4 . Die Diagnose der neuen endemischen Roccella cervicornis Follm. spec. nov. wird vorgelegt

     

Predicting lichen hydration using biophysical models

Two models for predicting the hydration status of lichens were developed as a first step towards a mechanistic lichen productivity model. A biophysical model included the water potential of the air, derived from measurements of air temperature, relative humidity and species-specific rate constants for desiccation and rehydration. A reduced physical model, included only environmental parameters, assuming instantaneous equilibration between the lichen and the air. These models were developed using field and laboratory data for three green algal lichens: the foliose epiphytic Platismatia glauca (L.) W. Culb., the fruticose epiphytic Alectoria sarmentosa (Ach.) Ach. and the fruticose, terricolous and mat-forming Cladina rangiferina (L.) Weber ex Wigg. The models were compared and validated for the same three species using data from a habitat with a different microclimate. Both models predicted the length and timing of lichen hydration periods, with those for A. sarmentosa and P. glauca being highly accurate—nearly 100% of the total wet time was predicted by both the biophysical and physical models. These models also predicted an accurate timing of the total realized wet time for A. sarmentosa and P. glauca when the lichens were wet. The model accuracy was lower for C. rangiferina compared to the epiphytes, both for the total realized wet time and for the accuracy of the timing for the hydration period. These results demonstrate that the stochastic and continually varying hydration status of lichens can be simulated from biophysical data. Further development of these models to also include water-related activity, light and temperature conditions during the hydration events will then be a potent tool to assess potential lichen productivity in landscapes and habitats of various microclimatic conditions.     

新疆梅衣属地衣生态地理特征分析

根据多年的实地调查资料和相关研究资料,对新疆梅衣属（Parmelia Ach．）地衣的种类及分布区、区系特征和垂直分布特征进行了初步研究。结果表明,分布在新疆的梅衣属地衣共有6种,分别为破裂梅衣（Parmelia erumpens Kurok．）、稀生梅衣（P．meiophora Nyl．）、脐梅衣[P．omphalodes（L．）Ach．]、石梅衣[P．saxatilis（L．）Ach．]、槽梅衣（P．sulcata Tayl．）和亚广开梅衣（P．fertilis Muell．）,主要分布在新疆的天山和阿勒泰山。根据对环境的适应特征和选择性,将新疆梅衣属地衣的地理分布区类型划分为：1）环低北极及北方种,包括石梅衣和脐梅衣2种;2）环北方种,包括槽梅衣和稀生梅衣2种;3）温带亚洲种,包括亚广开梅衣和稀生梅衣2种。研究结果还显示,分布在阿勒泰山和天山的梅衣属种类的垂直分布有明显差异。     

Screening the antioxidant and antimicrobial properties of the lichens Parmelia saxatilis, Platismatia glauca, Ramalina pollinaria, Ramalina polymorpha and Umbilicaria nylanderiana.

The aim of this study is to investigate in vitro antimicrobial and antioxidant activities of the methanol extracts of Parmelia saxatilis (L) Ach., Platismatia glauca (L.) W.L. Club. & C.F. Culb., Ramalina pollinaria (Wesstr.) Ach., Ramalina polymorpha (Liljeblad) Ach. and Umbilicaria nylanderiana (Zahlbr.) H. Magn. Antioxidant activity was evaluated by two separate methods: scavenging of free radical DPPH and the inhibition of linoleic acid oxidation. Extracts of Parmelia saxatilis, Platismatia glauca., Ramalina pollinaria and Ramalina polymorpha did not exert any activity in both assays, whereas those of Umbilicaria nylanderiana provided 50% inhibition at 400.2 microg/ml concentration in the former and gave 53% inhibition at 2g/l concentration. Total phenolic constituents of extracts from lichen species tested (P. saxatilis, P. glauca, R. pollinaria, R. polymorpha and U. nylanderiana) were 1.0% (w/w), 1.1% (w/w), 1.0% (w/w), 0.8% (w/w) and (3.0% w/w), respectively (as gallic acid equivalent); implying that the observed activity could be related to the amount of polar phenolics. Extracts were also found to possess antimicrobial activity against some test bacteria and fungi and yeast.     

The lichen genus Diploschistes in India and Nepal

Abstract: A taxonomic revision of 15 taxa of Diploschistes from India and Nepal is presented. Two species, D. awasthii and D. nepalensis, are described as new. Diploschistes caesioplumbeus (Nyl.,) Vainio, D. diacapsis (Ach.) Lumbsch and D. euganeus (Massal.) Steiner are new records for the lichen flora of India.     

PHYLOGENETIC STUDIES OF THE GENUSARTHONIA

A phylogenetic analysis is presented, based on 57 morphological, anatomical, chemical and ecological characters, including 19 species from the generaArthonia,ArthotheliumandSyncesia. The aim of the study was to test the monophyly of some of the groups within the genusArthoniasuggested, for example, by Redinger (e.g. species with reddish ascomata, with grey–pruinose ascomata and with brown–black hypothecia). The results strongly support thatArthoniaandArthotheliumare paraphyletic genera. The best-supported node contains allArthoniaspecies together withA. crozalsianade Lesd.,A. ruanaA. Massal. (both hitherto placed inArthothelium) andSyncesia myrticola(Fée) Tehler. A well-supported clade is formed by a group of pioneer species, often non-lichenized or poorly lichenized, including the type species ofArthonia,A. radiata. The species with reddish and/or K+ reddish ascomata form one clade and the species with more or less brownish or blackish hypothecia form another clade withSyncesia myrticola, the sister group to theOpegraphaceaeandRoccellaceae. The results are discussed and compared with Redinger’s grouping. Relationships to other genera within Arthoniaceae are briefly discussed.Arthothelium scandinavicumTh. Fr.,Arthonia dispersa(Schrad.) Nyl.,A. punctiformisAch. andA. mediellaNyl. are lectotypified.     

黑粉菌属一新种及中国黑粉菌补遗VI

本文描述采自河北小五台山的黑粉菌一新种,即寄生在野青茅(Deyeuxia arundinacea(L.) Beauv)上的野青茅黑粉菌(Ustilago deyeuxiae L. Guo)。此种与网优黑粉菌(Ustilago scrobiculata Liro)近似,但野青茅黑粉菌网纹明显,网眼高。并报道三种黑粉菌中国新记录。1)酢浆草黑粉菌(Ustilago oxalidis Ellis&Tracy)寄生于醉浆草(Oxalis corniculata L.),此种是我国首次在酢浆草科(Oxalidaceae)植物上发现的黑粉菌。2)网状黑粉菌(Ustilago polygoni-alati Thirum. & Pavgi)寄生于尼泊尔萝(Polygonum nepalense Meisn.),作者对此种进行了订正研究。3)臭草条黑粉菌(Urocystis melicae (Lagerheim & Liro) Zundel)寄生于细叶臭草(Melica radula Fr.)。     

On the Lichen generaPlacocarpus andPlacidiopsis (Verrucariaceae)

Verrucaria schaereri (Fr.)Nyl. is accorded generic status asPlacocarpus Trev. emend.O. Breuss. The main differences between the two genera concern the thallus anatomy and the spores which are halonate inPlacocarpus. The following new combinations are proposed:Placocarpus schaereri (Fr.)O. Breuss andPlacidiopsis cinerascens (Nyl.)O. Breuss.     

Problematik der Gattungsbegrenzung bei den Lichinaceen

Anhand von Typusarten und unter Berücksichtigung von weiteren Sippen werden die zur Gattungsbegrenzung von Lichinaceen mit einzelligen symbiotischen Algen geeigneten Merkmale diskutiert. Die wichtigsten Ergebnisse sind aus Tabelle 2 ersichtlich. Die neuen Arten Cryptothele negletta Henss., Gonohymenia inflata Henss. und Paulia aldabrensis Henss. werden beschrieben. Sechs neue Kombinationen werden gemacht: Cryptothele cylindrophora (Vain.) Henss., Gonohymenia cribellifera (Nyl.) Henss., Gonohymenia nummularia (Nyl.) Henss., Gonohymenia pulvinatum (Dahl) Henss., Pterygiopsis affinis (Mass.) Henss. und Synalissa mattogrossensis (Malme) Henss.     

Action inhibitrice de l'acide 3-indolyl-acétique sur la croissance de quelques Basidiomycètes mycorhizateurs

La croissance en milieu liquide de quelques champignons mycorhizateurs a étéétudiée en présence d'acide 3-indlolyl-acétique (AIA). Les champignons suivants ont été utilisés: Amanita muscaria (L. ex Fr.) Quél., A. rubscens (Pers. ex Fr.) Quél., Suillus granulatus (L. ex Fr.) O. Kuntze et S. variegatus (Sw. ex Fr.) O. Kuntze. Tous ont été inhibés, à des degrés divers, par l'AlA à la concentration do 1.75 mg/l. Soumis à diverses concentrations d'AIA, S. variegatus a montré une croissancc légèrement accrue au voisinage de 0.2 mg/l de cette substance. Cependant, pour des concentrations supérieures à celle-ci, l'inhibition de la croissancc a été proportion-nelle à la concentration en AIA; elle a été de 50 pour-cent en présence de 2.0 mg/l d'AIA.     

Lipid and Fatty Acid Composition of Tree-Growing and Terrestrial Lichens

A comparison of lipid and fatty acid composition was made of the tree-growing lichens Evernia prunastri (L.) Ach., Parmelia saxatilis (L.) Ach. and Hypogymnia physodes (L.) Ach. and the terrestrial species Cetraria islandica (L.) Ach. and Cladonia impexa Harm. In the terrestrial species the total lipid content varied strongly during spring, while the tree-growing species showed much less variation. Phospholipid and sterol content of all lichens was unusually low. Monoglycosyl diglyceride was absent from Parmelia saxatilis. Fatty acids common to higher plants as palmitic, stearic, oleic, linoleic and linolenic acid were invariably present in all lichen species. In addition large quantities of extra-long chain fatty acids like behenic acid, eicosadienoic acid and cyclic aliphatic lichen acids were present in the terrestrial species. The degree of (poly) unsaturation decreased in the order Evernia prunastri, Parmelia saxatilis. Fatty acids common to higher plants as palmitic, stearic, impexa, which decrease was compensated by an increase in extralong chain fatty acids and lichen acids. It is suggested that the lichen acids are of adaptive value for lichen species growing in the terrestrial habitats, which were characterized by extreme diurnal temperature variations. Just as the polyunsaturated fatty acids, lichen acids guarantee at lower temperatures a high flexibility of the membranes involved, at the same time as they are less susceptible to photo-oxidation at the high daytime temperatures.     

中国星文衣属地衣型真菌的分类研究

星文衣属(Sarcographa)隶属于真菌界(Fungi)、子囊菌门(Ascomycota)、茶渍纲(Lecanoromycetes)、厚顶盘目(Ostropales)、文字衣科(Graphidaceae),该属地衣体为壳状,子囊盘为放射分支状,多具子座,子囊含8孢子,子囊孢子褐色,横隔透镜型或亚砖壁型,主要分布于热带亚热带地区。该文通过形态学、解剖学、化学与分子生物学等方法进行该属分类学研究,共报道了中国该属9种,其中变黄星文衣[Sarcographa flavescens(Dal-Forno & Eliasaro)L. W. Chen & Z. F. Jia]为新组合(≡ Phaeographis flavescens Dal-Forno & Eliasaro); 曲线星文衣[Sarcographa labyrinthica (Ach.)Müll. Arg. ]为中国大陆新记录种。同时,该文对近似属拟星文衣属(Sarcographina)的1物种——异孢拟星文衣[Sarcographina heterospora(Nyl.)Z. F. Jia & Lücking]进行了描述,并提供了10个物种的特征提要及与其近似种的区别特征,并编制了分种检索表。该研究为中国地衣型真菌生物多样性研究提供了基础资料。     

Oribatid mite (Acari, Oribatida) feeding on ectomycorrhizal fungi

The coexistence of a large number of soil animals without extensive niche differentiation is one of the great riddles in soil biology. The main aim of this study was to explore the importance of partitioning of food resources for the high diversity of micro-arthropods in soil. In addition, we investigated if ectomycorrhizal fungi are preferentially consumed compared to saprotrophic fungi. Until today, ectomycorrhizal fungi have never been tested as potential food resource for oribatid mites. We offered six ectomycorrhizal fungi [Amanita muscaria (L.) Hook., Boletus badius (Fr.) Fr., Cenococcum geophilum Fr., Laccaria laccata (Scop.) Fr., Paxillus involutus (Batsch) Fr. and Piloderma croceum J. Erikss. & Hjortstam], one ericoid mycorrhizal fungus [Hymenoscyphus ericae (D.J. Read) Korf & Kernan] and three saprotrophic fungi [Agrocybe gibberosa (Fr.) Fayod, Alternaria alternata (Fr.) Keissl. and Mortierella ramanniana (A. Møller) Linnem.] simultaneously to each of the mainly mycophagous oribatid mite species Carabodes femoralis (Nicolet), Nothrus silvestris Nicolet and Oribatula tibialis Nicolet. The ericoid mycorrhizal fungus H. ericae and the ectomycorrhizal fungus B. badius were preferentially consumed by each oribatid mite species. However, feeding preferences differed significantly between the three species, with O. tibialis being most selective. This study for the first time documented that oribatid mites feed on certain ectomycorrhizal fungi.     

The genera Dirina and Roccellina (Roccellaceae)

In the present monograph of the coastal lichens, generally referred to as Dirina (crustose), Lobodirina (placoid), and Roccellina (suffruticose), two genera comprising 38 taxa are recognized: Dirina Fr. (7 species, 2 subspecies, and 4 forms) and Roccellina Darbish. (23 species and 2 forms). One Chiodecton and one Enterographa species are transferred to Dirina. Lobodirina Follm. is included in Roccellina together with seven species formerly placed in Dirina, one in Schismatomma, and one in Dirinastrum. The relationships between the two genera and the relation to other genera in Roccellaceae are discussed. Eleven species, and four forms are new: Dirina approximata Zahlbr. ssp. hioramii (B. de Lesd.) Tehler f. sorediala Tehler, D. catalinariae Hasse f. sorediala Tehler, D. insulana (C. Tav.) Tehler f. sorediala Tehler, Roccellina badia Tehler, R. cerebriformis (Mont.) Tehler f. sorediala Tehler, R. chalybea Tehler, R. conformis Tehler, R. exspectata Tehler, R. flavida Tehler, R. inaequabilis Tehler, R. nigricans Tehler, R. nigrocincta Tehler, R. obscura Tehler, R. suffruticosa Tehler, R. terrestris Tehler. Sixteen new combinations are proposed: D. approximata Zahlbr. ssp. africana (Fee) Tehler, D. approximata Zahlbr. ssp. hioramii (B. de Lesd.) Tehler, Dirina cretacea (Zahlbr.) Tehler, D. insulana (C. Tav.) Tehler, D. massiliensis Durteu et Mont. f. sorediala (Mull. Arg.) Tehler, Roccellina accedens (Nyl.) Tehler, R. capensis (Nyl. ex Stiz.) Tehler, R. cerebriformis (Mont.) Tehler, R. chilena (Dodge) Tehler, R. cinerea (Mull. Arg.) Tehler, R. cinerea (Mull. Arg.) Tehler f. sorediosa (Müll. Arg.) Tehler, R. falklandica (Zahlbr.) Tehler, R. limitata (Nyl.) Tehler, R. lutosa (Zahlbr.) Tehler, R. mahuiana (Follm.) Tehler, R. niponica (Nyl.) Tehler. All Dirina and 18 Roccelina species have been collected and studied in the field. Gross morphology and chemistry arediscussed. Statistical measurements of spore size are made for all species and subspecies. A cladistic relationship of the species of both genera, and a reduced area cladogram for Roccellina are proposed. Dirina is mainly restricted to the Northern Hemisphere and Roccellina, except for four species, is found in the Southern Hemisphere. The species are mainly bound to mediterranean, arid or subtropical climates.