J Hattori Bot. Lab. No. 100: 431- 441 (Aug. 2006)

WORLD DISTRIBUTION PATTERNS IN THE LOPHOZIACEAE / COMPLEX (HEPATICAE, BRYOPHYTA)

LARS SODERSTROM I AND ANA SENECA2

ABSTRACT. Global analysis ofbryophyte distribution ranges is generally absent from the literature. In order to understand general patterns of distribution we studied those for the members of the Lophoziaceae/Scapaniaceae complex at a worldwide scale. Number of species and number of re­ stricted species was investigated for the geographical units at three levels (continents, regions and areas). A rarity index was developed in order to identify areas where restricted species occurred and these areas were mapped. Differences in average distribution range and in the evenness of the distrib­ ution pattel11s were analyzed for various distribution groups. Himalaya, south-central China, Hawaii, Malesia, New Zealand, Tasmania. Venezuela, southern South America and some subantarctic Islands are hot spots for restricted species in the group. There are more species north of the tropics than in the tropics and south of the tropics and the northern species have much wider distribution and occur more evenly on available land areas than the rest.

K EY WORDS: Lophoziaceae, Scapaniaceae, global distribution, distribution patterns, rarity

INTRODUCTION There are a great number of publications describing distribution patterns, but the vast majority of them describe where a species (or a group of species) have its maximum of oc­ currence. Another topic much explored is the occurrence of endemic species (e.g. S6derstr6m 1996). However, more theoretic analyzes of the sizes of distribution ranges and the geographical distribution of distribution ranges are few and seems to be absent in bry­ ologicalliterature. As a part of a larger study on phylogeny and life history strategies in the liverwort family Lophoziaceae, distribution data was collected from literature for all taxa usually in­ cluded in the family. This family has variably been treated as a family of its own or as a subfamily of . At the time this project started, a couple of papers were published showing that Jungermanniaceae is not closely related to Lophoziaceae, and that Leiocolea and Jamesonielloideae, usually treated as parts of Lophoziaceae, do not belong there. Further the family Scapaniaceae (excl. Delavayellaceae and Blepharidophyllaceae) should be included in Lophoziaceae as a monophyletic group (Schill et aI. , 2004; Davis, 2004; Yatsentyuk et aI., 2004; De Roo et aI., unpubl.). But even after excluding these ele­ ments, the group is heterogeneous as e.g. Cephaloziaceae and Cephaloziellaceae seem to be nested within or at least sister to the rest. Species of Lophoziaceae and Scapaniaceae grows mainly on soil, epilithic or epixylic

1 Department of Biology, Norwegian University of Science and Technology, N-749 I , Trondheim, Norway. 2 Department of Botany and CIBIO, University of Porto, Rua Campo Alegre 1191 , P-4150-181 Porto, Portugal. 432 J. Hattori Bot. Lab. No. 100 2 0 0 6 and only very few are epiphytic. They are also mainly requiring moist or wet conditions and thus largely absent from drier regions, or regions with a prolonged dry period (Soderstrom et aI. , unpubl.). Within the complex, several species do occur over large areas (circumboreal) and some species (e.g. excisa and hatcheri) are also bipolar. But there are also species with very restricted distribution, e.g. tenue occurring in a few localities in Ontario, Canada. We are interested in distribution patterns within the Lophoziaceae/Scapaniaceae com­ plex. The questions we put forward in this paper are: 1) which species have the most re­ stricted distribution, 2) which areas are most species rich and which areas have most re­ stricted species, 3) is there a correlation between species richness and number of restricted species.

METHODS The Lophoziaceae/Scapaniaceae complex, as we define it here (excl. lamesoniel­ loideae, Leiocolea, Cephaloziaceae, Cephaloziellaceae, Blepharidophyllaceae and Delavayellaceae) includes 303 species. Some of the species are badly known and may turn out to be synonyms with other, more well-known, species. We are also convinced that there are undescribed species in the complex. However, we think that this will not have any major influence on the patterns we find with the present data set. The classification of distribution ranges follow mainly Brummit (200 I) and are scored on 3 levels. Level I is basically the continents and has 9 units. Level 2 is regions within continents and has 51 units (4 in Europe, 10 in North America, 10 in Africa, etc.). Level 3 are basically countries except that large countries (e.g. Russia and Canada) are separated in smaller units and that very small countries are included in a neighbour (e.g. Andorra in Spain and Liechtenstein in Austria). We also recognize the following countries at level 3 al­ though not recognized in Brummit (2001): Estonia, Latvia, Lithuania, Czech Republic, Slovak Republic, Slovenia, Croatia, Serbia-Monte Negro and FYR Macedonia). This gives a total of 383 units (49 in Europe, etc.). In this paper we use the term continent for level 1 units, region for level 2 units and areas for level 3 units. We have scored the number of units at each level a species occur in, mainly by scanning the literature. We calculated a rarity index using the same formula as the Shannon-Wiener diversity index (Zar 1984) as

H'=-I,Pi lnPi where P is the proportion of areas occupied in each region. We also calculated the index with proportion of areas in each continent, and proportion of regions in each continent. The three measures were highly correlated. The areas in regions showed most resolution and is thus used here. With this definition, the H' value is an index on how large the chance is to find the same species in the next region visited. There are many more regions in the northern hemisphere than in the southern thus giving higher maximum H' value for species confined to the north than for those confined L. SODERSTROM & A. SENECA: World distribution patterns in the Lohoziaceae/Scapaniaceae complex 433

Table I. Number of regions north of the Tropic of Cancer (N), north of the Tropic of Capricorn (NT), south of the Tropic of Cancer (TS), south of the Tropic of Capricorn (S), between the Tropics (T), outside the Tropics (NS) and total number (NTS).

Distr. group Number of regions Distr. group Number of regions

N 23 NTS 51 T 17 TS 6 S 5 NS 28 NT 27

to the south. Therefore we calculated the evenness component (1) as H' IH' max for each species. H' max is defined as log(r) (cf. Zar 1984) where r is the number of regions. We di­ vided the globe in three latitudinal groups, north of the Tropic of Cancer (N), south of the Tropic of Capricorn (S) and between the tropics (T). For each species occurring only in the Nr is the number of regions in the N, for species occurring both in Nand Tr is the number of regions in N + T, etc. The r value used for different groups of occurrences is shown in Table I. We then compared pair wise the 1 value for the distribution groups to see if species are more evenly distributed in any of the groups. For this we used the Mann-Whitney U­ test (Zar 1984). To be able to evaluate the importance of different areas for species with restricted dis­ tribution, we calculated an index as

RI=p~nr where Pr is the proportion of the species occurring in the area that has a restricted distribu­ tion and nr is the number of rare species in that region. This index downgrades the effects of rare species in areas with very few species. Statistical analysis was performed in SPSS version 14.0. The variables were not nor­ mal distributed and all correlations were for this reason performed with Spearman correla­ tion coefficient.

RESULTS 3.1. Distribution ranges The correlation between distribution at the three levels (continents, regions and areas) are all strongly positive with a high correlation coefficient (0.880-0.968; p

Table 2. The 158 species with H'> I and/or occur in 4 or fewer areas.

Taxon Areas H' Taxon Areas H'

Anastrophyllum antidens 0.00 Gymnocoleopsis capensis 0.00 Anastrophyllum crenulatum 0.00 Lophozia aenigmatica 0.00 Anastrophyllum divergens 0.00 Lophozia androgyna 0.00 Anastrophyllum ellipticum 0.00 Lophozia anomala 0.00 Anastrophyllum esenbeckii 0.00 Lophozia autoica 0.00 Anastrophyllum fissum 0.00 Lophozia brunnea 0.00 Anastrophyllum hypoc/adopsis 0.00 Lophozia debilis 0.00 Anastrophyllum intermedium 0.00 Lophozia drucei 0.00 Anastrophyllum karstenii 0.00 Lophozia gymnocoleopsis 0.00 Anastrophyllum lancilobum 0.00 Lophozia nakanishii 0.00 Anastrophyllum leptodictyon 0.00 Lophozia nepalensis 0.00 Anastrophyllum minutirete 0.00 Lophozia nivicola 0.00 Anastrophyllum novae-zelandiae 0.00 Lophozia pallida 0.00 Anastrophyllum obtusum 0.00 Lophozia perssoniana 0.00 Anastrophyllum papillosum 0.00 Lophozia pumicicola 0.00 Anastrophyllum paramicola 0.00 Lophozia subalpina 0.00 Anastrophyllum striolatum 0.00 Lophozia subantarctica 0.00 Anastrophyllum tenue 0.00 Lophozia subapiculata 0.00 Anastrophyllum tristanianum 0.00 Lophozia tasmanica 0.00 carinatus 0.00 Lophozia tristaniana 0.00 Andrewsianthus cavi{olius 0.00 Lophozia verruculosa 0.00 Andrewsianthus confusus 0.00 Nothostrepta longissima 0.00 Andrewsianthus kinabaluensis 0.00 Pisanoa chilensis 0.00 Andrewsianthus koponenii 0.00 Pseudocephaloziella epiphytica 0.00 Andrewsianthus mizutanii 0.00 Scapania diplophylloides 0.00 Andrewsianthus papillosus 0.00 Scapania ferruginaeoides 0.00 Andrewsianthus planifolius 0.00 Scapania gaochii 0.00 Andrewsianthus sundaicum 0.00 Scapania geppii 0.00 Andrewsianthus iantenii 0.00 Scapania grandiloba 0.00 Cephalolobus hodgsonae 0.00 Scapania grossidens 0.00 Cephalolobus scabrellus 0.00 Scapania hedbergii 0.00 Cephalolobus sphenoloboides 0.00 Scapania hollandiae 0.00 Cephalolobus squarrosus 0.00 Scapania horaria 0.00 difficilis 0.00 Scapania integerrima 0.00 Diplophyllum andicolum 0.00 Scapania komagadakensis 0.00 Diplophyllum androgynum 0.00 Scapania koponenii 0.00 Diplophyllum angustifolium 0.00 Scapania matveyevae 0.00 Diplophyllum exiguum 0.00 Scapania nipponica 0.00 Diplophyllum gemmiparum 0.00 Scapania pseudocontorta 0.00 Diplophyllum incrassatum 0.00 Scapania rigida 0.00 Diplophyllum novum 0.00 Scapania schljakovii 0.00 Diplophyllum squarrosum 0.00 Scapania spiniloba 0.00 Gerhildiella rossneriana 0.00 Scapania udarii 0.00 maxima 0.00 Scapania valdonsmithii 0.00 L. S6DERSTR6M & A. SEN ECA: World distribution patterns in the Lohoziaceae/Scapaniaceae complex 435

Table 2. (Continued.)

Taxon Areas H' Taxon Areas H'

Scapania zhukovae 0.00 Scapania orientalis 4 0.87 Tetralophozia pilifera 0.00 Scapania pseudocalcicola 3 0.88 camerunensis 0.00 Scapania serrulata 3 0.88 Anastrophyllum denticulatum 2 0.00 yakushimensis 4 0.89 Anastrophyllum involutifolium 2 0.00 Anastrophyllum borneense 5 0.89 Andrewsianthus ferrugineus 2 0.00 Diplophyllum acutilobum 3 0.90 Gottschelia patoniae 2 0.00 Nothostrepta bifida 3 0.90 Lophozia austro-sibirica 2 0.00 Lophozia patagonica 2 0.92 Lophozia crispata 2 0.00 borealis 4 0.92 Lophozia lancistipa 2 0.00 Diplophyllum verrucosum 4 0.93 Lophozia lantratoviae 2 0.00 Lophozia herzogiana 2 0.94 Lophozia leucorhiza 2 0.00 Anastrophyllum subcomplicatum 3 0.95 Scapania ampliata 2 0.00 Diplophyllum dioicum 2 0.96 Scapania esterhuyseniae 2 0.00 Lophozia multicuspidata 2 0.96 Scapania fulfordiae 2 0.00 Diplophyllum trollii 3 0.96 Scapania himalayica 2 0.00 Scapania bhutanensis 3 0.96 Scapania sandei 2 0.00 Scapania contorta 3 0.96 Scapania sinikkae 2 0.00 Scapania davidii 3 0.96 Scapania subnimbosa 2 0.00 Scapanid harae 3 0.96 Andrewsianthus aberrans 3 0.00 Diplophyllum marionense 3 0.98 Andrewsianthus marionensis 3 0.00 Gymnocolea fascinifera 2 0.99 Anastrophyllum intricatum 4 0.00 Scapania gigantea 2 0.99 Gymnocoleopsis cylindriformis 4 0.00 Scapania hirosakiensis 2 0.99 Scapania gamundiae 2 0.44 Anastrophyllum austroamericanum 2 0.99 Scapania sphaerifera 7 0.45 Anastrophyllum lignicola 2 0.99 Diplophyllum africanum 3 0.56 Scapania rotundifolia 2 0.99 Diplophyllum serrulatum 4 0.72 Scapania secunda 4 0.99 Anastrophyllum prionophyllum 2 0.78 Anastrophyllum squarrosum 4 1.00 Andrewsianthus chimbuensis 2 0.78 Andrewsianthus recurvifolius 4 1.00 Lophozia argentina 4 0.78 Andrewsianthus australis 2 1.00 Lophozia badia 3 0.80 Anastrophyllum ciliatum 4 1.00 Anastrophyllum integerrimum 3 0.83 Scapania lepida 2 1.00 Scapania karl-muelleri 3 0.86 Scapania hians 3 1.00 Lophozia setosa 4 0.87 Chandonanthus squarrosus 4 1.28 Scapania griffithii 4 0.87 Andrewsianthus bidens 4 1.47 sphaerifera and Anastrophyllum borneense do have H' < 1 although occurring in 7 and 5 areas, respectively. Scapania sphaerifera do occur in many (6) areas in 1 region (Siberia), but only in one area outside that. Anastrophyllum borneense occurs in just 2 regions but in a low proportion of the areas in each. Therefore, the H' will be lower as the evenness com- ponent (cf. Zar 1984) will be very low. Of the species with H';::: 1, eight occur in 4 or fewer areas and six of them have H' = 1. The other 2 species, Chandonanthus squarrosus and Andrewsianthus bidens occurs in three 436 1. Hattori Bot. Lab. No. 100 2 0 0 6 regions but in only one or two areas in each giving them an H' value of 1.28 and lA 7 re­ spectively (Table 2).

3.2. Geographical distribution ofdistribution range sizes There are 183 species occurring north of the Tropic of Cancer and 148 out of them occur only there. South of the Tropic of Capricorn 76 species can be found out of which 55 are restricted to this region and in between the Tropics 94 species occur (60 only there). The average H' value for all species is 1.50:::t:: 1.48 SO and the evenness (J) is 1.15:::t:: 1.80 SO (Table 3). The species with the highest average H' value are, as expected, the species occurring in both the north, tropics and the south (3.93:::t::0.61 SO) followed by those occurring in two latitudinal regions. Of the species occurring in only one latitudinal region, species in the north have a much higher average H' value than the other two (Table 3). The distribution group with the highest average evenness (J) is those occurring in the tropics and south (TS; 3.28:::t::O.99 SO). Most of the other values follow the pattern for H ' (Table 3). Comparing the average distribution range (H') and the average evenness (1) values,

Table 3. Average H' and J value for the seven distribution groups recognized.

Distr. Distr. n H' (::t:SD) J(::t:SD) n H' (::t:SD) J (::t:SD) group group

N 148 1.93 ::t: 1.44 IAI::t: 1.01 S 55 0.36::t:0.66 0.52 ::t: 0.95 NS 6 2.70::t:1.12 1.87::t:0.77 T 60 OA2::t:0.66 0.34 ::t: 0.54 NT 19 2.98::t:0.98 2.08::t:0.68 TS 5 2.56::t:0.77 3.28::t:0.99 NTS 10 3.93::t:0.61 2.29::t:0.36

Table 4. Pair wise comparison of the differences in mean H' value (upper right) and mean J value (lower left) for the distribution groups. Stars mean that they are significant different at 90% level (*), 95% level (**) and 99% level (***), respectively. n.s.=non sig- nificant.

Distr. group N NS NT NTS S T TS

N n.s. ** *** *** *** n.s. NS n.s. n.s. n.s. *** *** n.s. NT * n.s n.s. *** *** n.s. H' NTS n.s n.s n.s. *** *** n.s. S *** *** *** *** n.s. *** T *** *** *** *** n.s. *** TS n.s. n.s. n.s n.s. *** *** L. SOOERSTROM & A. SENECA: World distribution patterns in the Lohoziaceae/Scapaniaceae complex 437 species occurring in the south (S) and in the tropics (T) differ significantly from all other distribution groups except themselves (Table 4). The only significant difference not includ­ ing those two are the difference between Nand NT (both on average H' and average J) and between NTS and N in H'. The last pair does not, however, differ in average evenness value.

3.3. Species rich areas and areas with restricted species One or more species occurred in 254 out of the 383 areas. Alaska has the highest

'-

0 1-10 11-20 21-30 31-40 41-50 51-60 61-10 11-SO 81-90 91-100

Fig. 1. Number of species in each area.

Fig. 2. Number of restricted species (H' < 1) in each area. 438 1. Hattori Bot. Lab. No. 100 200 6 number of species (91) followed by Norway (90), Northern European Russia (84), Sweden (83) and West Siberia (82). No area on the southern hemisphere had over 25 species (Fig. I). One or more restricted species (H' < I) occurs in 85 areas. The arctic and the boreal regions do have very few species with restricted distribution (Fig. 2). Nepal and South-cen­ tral China have most restricted species (19) followed by East Himalaya (14) Japan, South Chile, New Zealand South I (all 11), New Guinea and Tasmania (both 9). There were 169 areas without any of their species being restricted. Number of restricted species is positively correlated with number of species (correla­ tion coefficient 0.370, p

DISCUSSION The species of the Lophoziaceae/Scapaniaceae complex is markedly arctic and boreal in its distribution. A large number of the species do have very restricted distributions with about 113 of the species occurring in only I region and 112 the number of species with an H' value < 1. Fifteen of the species occur in only I area and it is noticeable that 13 of them are Scapania species .. It is expected that a badly known group includes several taxa de­ scribed long time ago from remote areas, but never re-evaluated, that could turn out to be conspecific with more common taxa. However, in Scapania this seems not to be the case since Potemkin (2002) has revised the genus worldwide. Although he leaves some question marks, there is no reason to believe that all or even a large part of the 13 species occurring in just one area will be synonymized with a more common species. Then it is more proba­ ble that some of the very restricted species will turn up elsewhere. The number of species occurring in the northern hemisphere is higher than in the tropics and in the southern hemisphere. In addition, as the distribution ranges are in aver­ age very much larger in the northern hemisphere, the individual areas are much more species rich in the north than in the south and the tropics. Of the area in the tropics, the northern Andes are the most diverse (mainly due to the many species of Anastrophyllum) as well as Borneo and New Guinea. However, most of the species occurring there are found on higher mountains. South of the tropics, only southern South America has a higher num­ ber of species. The number of restricted species does not follow the same pattern although the num­ ber of species is positively correlated with number of restricted species. Instead they are concentrated to the southern hemisphere, except that the Himalaya and neighbouring areas of China are exceptionally rich in restricted species. Species occurring only south of the Tropic of Capricorn and those occurring only in between the Tropics do have smaller dis­ tribution areas and, in addition, do not fill up the available areas as well as the species oc­ curring north of the Tropic of Cancer or with a wider distribution. An explanation may be that the northern hemisphere, especially the area north of the Tropic of Cancer, consists of L. SODERSTROM& A. SENECA: World distribution patterns in the Lohoziaceae/Scapaniaceae complex 439

Table 5. Number of species, number of restricted species and RI for areas where re- stricted species occur.

Restr. Restr. Area Species RI Area Species R! species species

South-central China 46 19 7.85 Mauritius 3 0.33 Nepal 46 19 7.85 Victoria 3 0.33 New Zealand South I 17 10 5.88 Society Is 3 0.33 South Chile 21 11 5.76 Tanzania 15 2 0.27 Marion-Prince Edward Is 7 6 5.14 Samoa 4 0.25 Tasmania 13 8 4.92 Greenland 77 4 0.21 East Himalaya 43 14 4.56 Lesotho 5 0.20 New Guinea 24 9 3.38 Madagascar 5 0.20 Southern Argentina 15 7 3.27 Reunion 5 0.20 Venezuela 20 8 3.20 India 5 I 0.20 Crozet I 5 4 3.20 Irkutsk 48 3 0.19 Borneo 26 8 2.46 Cameroon 6 0.17 Hawaii 11 5 2.27 K waZulu-Natal 6 0.17 Kerguelen 2 2 2.00 Northeast Mexico 6 0.17 Japan 69 11 1.75 Veracruz 7 0.14 New Zealand North I 6 3 1.50 Korea 33 2 0.12 Juan Fernandez Is 3 2 1.33 Rwanda 9 0.11 Falkland Is 8 3 1.13 Central Mexico 9 0.11 Colombia 16 4 1.00 Malaya 10 0.10 St Helena 1 1 1.00 Sumatra 10 0.10 Mariana Is 1 1 1.00 Ecuador 11 1 0.09 Java 19 4 0.84 Nunavut 48 2 0.08 West Himalaya 11 3 0.82 Uganda 12 0.08 Cape Provinces 5 2 0.80 Southwest Mexico 14 0.07 New Caledonia 6 2 0.67 Krasnoyarsk 70 2 0.06 Tibet 25 4 0.64 Yakutia 74 2 0.05 Tristan da Cunha Is 7 2 0.57 West Siberia 81 2 0.05 Moluccas Is 7 2 0.57 Wyoming 26 0.04 Solomon Is 7 2 0.57 Tuva 27 0.04 South Georgia 7 2 0.57 Colorado 33 0.03 South China 16 3 0.56 Ontario 47 0.02 Antarctic Continent 8 2 0.50 Buryatiya 49 0.Q2 Chita 2 0.50 Labrador 49 0.02 Lesser Sunda Is 2 0.50 Newfoundland 50 0.02 New South Wales 2 0.50 Washington 51 0.02 Vanuatu 2 0.50 Altay 51 0.02 Mikronesia 2 0.50 Britain 61 0.02 Winward Is 2 0.50 Finland 72 0.01 South Brazil 2 0.50 Sweden 82 0.01 Heard-McDonald Is 2 0.50 North Russia 83 0.01 Taiwan 35 4 OA6 Norway 89 0.01 Kenya 10 2 OAO Alaska 90 0.01 440 J. Hattori Bot. Lab. No. 100 2 006

Fig. 3. Distribution of RI over areas. more land masses (40%) that is more or less continuous with suitable habitats while the southern hemisphere have less land mass (20%) and is more split. In the southern hemi­ sphere the occurrences are restricted by large oceans. To be widespread they will need higher dispersal ability than the species on the northern hemisphere. Thus, in southern South America (south Chile and southern Argentina), Tasmania, New Zealand and some subantarctic Islands, species may have evolved due to long term isolation under moderately stable conditions, which they seems to have been recently (Adams & Faure 1997). Al­ though dispersal of diaspores over longer distances is possible in the southern hemisphere (Muiioz 2004), many species in the Lophoziaceae/Scapaniaceae complex do not seem to have a particularly good dispersal ability (rarely produce spores or gemmae, or the dias­ pores are too large; cf. S6derstr6m & Herben 1997) or diaspores may not survive dispersal over longer distances (cf. Van Zanten 1983). They are thus spreading only short distances, as oceans seem to act as efficient barriers.

ACKNOWLEDGEMENTS We thank Jifi Vaiia for providing us with information on species synonymy. This is a part of a project on "Biodiversity, rarity, life history and phylogeny: a case study in the he­ patic family Lophoziaceae" with grants from the Research Council of Norway and NRF, South Africa (project no. 152297NI 0) given to Lars S6derstr6m and Terry Hedderson.

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