Four species of Ardisia (Myrsinaceae, Magnoliopsida) with rosette or low-lying leaves in China (including Hong Kong) have been found to be the hosts for 12 species of epiphyllous liverworts which belong to 4 families and 9 genera. However, no obvious species-specific hostepiphyte relationship could be recognized

But, Paul P.P.H.

Wang, Mei-zhi

Wu, Pan-cheng

English

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27TROPICAL BRYOLOGY 19 (2000)Epiphyllous liverworts on Ardisia speciesTropical Bryology 19: 27-30, 2000Epiphyllous liverworts on rosette leaves of Ardisia species(Myrsinaceae) in China1Paul P.P.H. But,  1,2 Pan-cheng Wu,  and  1,2 Mei-zhi Wang1Department of Biology, Chinese University of Hong Kong, Shatin, N.T., Hong Kong;2Institute of Botany, Academia Sinica, Beijing 100093, China.Abstract: Four species of Ardisia (Myrsinaceae, Magnoliopsida) with rosette or low-lying leavesin China (including Hong Kong) have been found to be the hosts for 12 species of epiphyllousliverworts which belong to 4 families and 9 genera.  However, no obvious species-specific host-epiphyte relationship could be recognized.IntroductionEpiphyllous liverworts manifest an interestingrelationship with broad-leaved vascular plants.Found in tropical and subtropical regions wherethe habitats have a high relative humidity in theair and shaded areas with only diffuse light, theseliverworts grow on the upper leaf surface of theirhosts. The biodiversity and host-epiphyterelationship has attracted much attention,particularly in China (But & Gao, 1991; Chen &Wu, 1964; Gao & But, 1988; Li & Wu, 1988;Lin et al. 1982; Liu et al., 1988; Luo, 1990; Wuet al., 1983, 1987a, b; Wu & Guo, 1986; Wu &Lin, 1978, 1988, 1994; Wu & Luo, 1978; Zhu1995).  However, little scientific studies havebeen made on the microhabitats and thebiological relationship between epiphyllousliverworts and their vascular hosts.  But & Gao(1991) have observed that the host leaves areoften of two or more years old, flat, broad, ortogether they form a broad surface (e.g.Selaginella), smooth or hairy.  Such leavesinclude those on stoloniferous and creepingspecies and thus lying close to the ground andalso those growing on trees up to 2 m above theground.  Wu et al. (1987a, b) have compiled somepreliminary observation on the diffuse lightcondition of the general environment ofepiphyllous liverworts.  Berrie & Eze (1975) havedemonstrated that in Radula flaccida, a generallyterrestrial liverwort but occasionally epiphytic onleaves, the water and P32 nutrient could movefrom the host leaves to the liverwort.  Thus, theyconcluded that epiphyllous R. flaccida is a semi-parasite rather than a simple epiphyte.  The lack28TROPICAL BRYOLOGY 19 (2000)But, Wu & Wangof more detailed studies on the epiphyte-hostrelationship may possibly be caused by thedifficulty in maintaining a convenient system forproper investigation.  Recently, But (1997)reported that epiphyllous liverworts were foundon the leaves of rosette-leaved angiosperms andsuggested that the hosts and the epiphytestogether could be transplanted to green housesor laboratories for controlled experimentalstudies.Following this lead, we have further checked onvarious rosette-leaved angiosperms in the fieldand herbaria.  In this report, we report on theliverworts found on host plants belonging to theangiosperm family Myrsinaceae.Myrsinaceae is a family of dicotyledon plantsdistributed mainly in tropical regions (Hutchinson1959).  Majority of the genera and species in thisfamily are trees or shrubs, but Ardisiaprimulaefolia Garn. et Champ. is sub-herbaceousand has a stunted stem with only rosette leaveslying flat on the ground.  The leaves are obovateor oblong-obovate, 6–17 x 3–10 cm, somewhatsucculent, densely gland-dotted and soft-hairy.It is widely distributed in southern China,including such provinces as Yunnan, Guangxi,Fujian, Hainan and Guangdong as well as HongKong, growing on the ground in hillsidewoodlands at altitudes between 500–1400 m.Very frequently, epiphyllous liverworts are foundgrowing on the leaves of this plant, especially inwoodland areas close to running water or fed bymorning fogs.  Another three species, Ardisiamamillata Hance, A. pubivenula Walker, and A.villosa Roxb., are not really rosette-like in leafformation.  They are, however, short shrubby orsub-herbaceous plants with low-lying stemsbearing leaves that are close to the ground (Chen,1979).  Their leaves are also gland-dotted andstiff, or soft and hairy.  Liverworts are foundgrowing on their leaves.  A total of 12 species ofliverworts are identified on the leaves of thesefour species of Ardisia in China (List 1).Ardisia primulaefolia is the more popular hostplant for epiphyllous liverworts.  So far, sixepiphyllous liverworts, Cheilolejeunea intertexta,Cololejeunea goebelii, C. spinosa, Leptolejeuneaelliptica, Heteroscyphus argutus andLopholejeunea subfusca, are found on the leavesof A. primulaefolia.  On the other hand,Cololejeunea goebelii is the more widespreadliverwort, found on the leaves of not only Ardisiaprimulaefolia, but also of Ardisia mamillata andA. villosa.  No species-specific relationship,however, could be recognized.  It may be ofinterest to note that leaves of the four species ofArdisia are densely hairy, bearing around 100hairs per square centimeter.  Therefore, the leafystems of the 12 species of liverworts could onlyanchor on the hairless areas and spread theirbodies in the ‘forest’ of hairs.The four species of Ardisia are widely distributedin southern regions of China.  They are short andsmall plants.  Therefore, they can be easilytransplanted to green houses and laboratories forcontrolled experimentation on the biologicalrelationship between the hosts and epiphyllousliverworts.  It would be interesting if furtheranalysis on the presence of epiphyllous liverwortson these plants may suggest them to be indicatorsof specific ecological condition and degree ofbiodiversity.  Moreover, since it is quite oftenfor the angiosperm botanists to collect the wholeplant of rosette-leafed species as a herbariumspecimen, it would be convenient to simply checkthe preserved dried specimens in herbaria anddetermine the extent and diversity of theirepiphyllous liverworts that can suggest theecological conditions in the various collectionlocalities.List 1. A list of the epiphyllous liverworts on theleaves of MyrsinaceaeHerbertaceae1. Herberta adunca (Dicks.) S. GrayYunnan: Deqin Co., on leaves of Ardisiapubivenula, 2,700-3,200 m alt., (collectoranonymous 5576).Lepidoziaceae2. Bazzania tridens (Reinw., Blume et Nees)Trev.Hainan: Mt. Wu Zhi, on leaves of Ardisiamamillata (F.A. McClure & E. D. Merrill9333).Lophocoleaceae3.     Heteroscyphus argutus (Reinw., Blume etNees) SchiffnerGuangdong: Liang Co., Mt. Tianguang, onleaves of Ardisia primulaefolia, 800 m alt.(P-X. Tan 60201a).29TROPICAL BRYOLOGY 19 (2000)Epiphyllous liverworts on Ardisia speciesLejeuneaceae4.     Lopholejeunea subfusca (Nees) StephaniHong Kong: Mt. Parker, on leaves ofArdisia primulaefolia (S.-Y. Hu 9256).5.     Drepanolejeunea dactylophora (Nees)SpruceHainan: Baoting Co., on leaves of Ardisiapubivenula (K.-Z. Hou 73665b).6.     Drepanolejeunea thwaitesiana (Mitten)StephaniHainan: Baoting Co., on leaves of Ardisiapubivenula (K.-Z. Hou 73665a).7.     Cheilolejeunea intertexta (Lindenberg)StephaniHong Kong: Tai Mo Shan, on leaves ofArdisia primulaefolia (S.-Y. Hu 6121b).8.     Leptolejeunea elliptica (Lehmann et Linden-berg) SchiffnerFujian: Nanqing, Mt. Xinluo, on leaves ofArdisia primulaefolia, 510 m alt. (AmoyUniversity 618).9.     Lejeunea ulicina (Tayl.) GottscheYunnan: Mar-Li-Po, on leaves of Ardisiaprimulaefolia, ca. 1000 m alt. (P.-X. Tan60201).10.   Cololejeunea goebelii (Schiffner) SchiffnerHainan: Mt. Wuzhi, on leaves of Ardisiavillosa (F.A. McClure et E.D. Merrill8671).  Guangdong: Mt. Lofau, on leavesof Ardisia mamillata (C. Ford withoutnumber); Liang Co., near Wangjunding, onleaves of Ardisia primulaefolia (P.-X. Tan58551b).  Guangxi: Lang Co., on leaves ofArdisia primulaefolia, ca 400 m alt. (R.-C.Ching 5664).  Fujian: Liancheng Co., onleaves of Ardisia primulaefolia (R. Lin3824).  Hong Kong: Pokfulam, on leavesof Ardisia primulaefolia (H.-C. Tang 430);Mt. Gough, on leaves of Ardisiaprimulaefolia (H.-C. Tang 1486); FindleyRd., on leaves of Ardisia primulaefolia (H.-C. Tang 1188).11.   Cololejeunea spinosa (Horikawa) Pande etMisraHainan: Wengyuan Co., Mt. Bijia, onleaves of Ardisia primulaefolia (X.-X. Liu25270b). Yunnan: Mar-Li-Po, on leaves ofArdisia primulaefolia, 1,000 m alt. (C.-W.Wang 86842a).  Guangdong: Liang Co.,Mt. Tianguang, on leaves of Ardisiaprimulaefolia, 800 m alt. (P.-X. Tan60201b); Liang Co., near Wangjunding, onleaves of Ardisia primulaefolia, 700 m alt.(P.-X. Tan 58551a).12.   Cololejeunea reineckeana StephaniHainan: Wengyuan Co., Mt. Bijia, onleaves of Ardisia primulaefolia (X.-X. 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Epiphyllous liverworts on rosette leaves of Ardisia species (Myrsinaceae) in China

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