Abstract: A compilation of records of photobionts that are associated in lichens of the order Lecanorales shows that mycobionts, when seen at the levels of suborders, families, and genera, are strongly selective towards their photobionts. The recently proposed split of the genus Trebouxia into two genera, Asterochloris and Trebouxia, is reflected by the choice of mycobionts. While Asterochloris is mostly found in the suborder Cladoniineae, Trebouxia is mainly confined to the Lecanoriineae. The symbiosis with coccoid green algae other than Trebouxia or Asterochloris might indicate the basal position of a lichen family within Lecanorales. For instance, Dictyochloropsis and Coccomyxa are confined to the Lecanoralean families Biatoraceae (Lecanorineae), Megalosporaceae (cf Teloschistineae), Peltigeraceae and Nephromataceae (Peltigerineae), and Catillariaceae (fam. inc. sed.). The most important cyanobacterial photobionts in Lecanorales are Nostoc and Scytonema as primary and accessory photobionts, while Gloeocapsa and Stigonema are more frequent as accessory photobionts in a few suborders and families of Lecanorales. The co-occurrence of cyanobacteria with Trentepohlia as a primary photobiont in lichen symbioses is not known from families of lichenized ascomycetes. This may reflect differences of ecological preferences of both photobiont groups and of the mycobiont groups that belong to different natural relationships. These examples show that the knowledge of photobionts is valuable information for systematics of lichens and therefore, exploration of photobiont diversity in lichens needs to be expanded. Within the last years, lichen photobionts have attracted increasing interest that has led to the unraveling of their diversity and evolutionary relationships using microscopy, ultrastructure, and molecular techniques (Btidel 1992; Ettl & Giirtner 1995; Friedl 1995; Friedl & Biudel 1996; Melkonian & Peveling 1988; Tschermak-Woess 1988). In the green algal genus Trebouxia, the most common photobiont, the recently demonstrated congruence of morphological and ultrastructural characters with rDNA sequence data resulted in a revised systematic concept of the genus (Friedl & Rokitta 1997). However, the question whether photobionts are important markers for the evolution and systematics of lichens, has been controversially discussed (Ahmadjian 1993). The capability of lichen forming fungi to specifically select their appropriate partners among the algal taxa common in subaerial habitats may be an important phylogenetic trait. From a closer examination of the distribution of algal taxa symbiotic in lichens it may become obvious that only a single algal species or a very narrow spectrum of algal taxa may cause transformation of a fungus into a lichen thallus. Chodat (1913), an early pioneer in the study of lichen algae, felt that his investigations indicated specificity of lichen-forming fungi with respect to their algal symbionts. Later, observations of Jaag (1933) showed that lichen species belonging to the same genus (e.g., Cladonia, Parmelia, Solorina and Umbilicaria) always contain the same certain type of 'gonidia.' He thought that this phenomenon may also hold true at the level of lichen families. In the following, however, the investigators of lichen photobionts stated that specificity of lichen-forming fungi with respect to their algal symbionts is rather poor. Wang-Yang and Ahmadjian (1972) found "different species or strains of algae in morpholog1 This paper was presented as part of a symposium entitled "Bridging the Gap Between Phylogeny and the Classification of Lichen-forming Ascomycetes" organized by Frangois Lutzoni in Montreal, Canada at the August, 1997, ABLS annual meeting. 0007-2745/98/392-397$0.75/0 This content downloaded from 157.55.39.249 on Wed, 03 Aug 2016 05:52:48 UTC All use subject to http://about.jstor.org/terms 1998] RAMBOLD ET AL.: PHOTOBIONTS IN LICHENS 393 ically identical lichen thalli and conversely different subspecies of a particular lichen may contain the same type of algal symbiont." The authors felt that the contribution of knowledge of photobionts to lichen systematics might be poor. Recently, Ahmadjian (1993) stated that "it is unlikely that taxonomists can find much value in the types of photobionts that occur in lichens, at least at the species

Abstract: The internal transcribed spacers (ITS) of the nuclear ribosomal DNA were sequenced for 16 specimens of the lichen Dendrographa leucophaea (Tuck.) Darb. (Roccellaceae) in order to determine whether the sterile and fertile specimens within the species (also called a species-pair) form two distinct monophyletic groups. Forty-four informative sites were obtained. Phylogenetic trees were calculated from aligned sequence data, using Dendrographa alectoroides Sundin & Tehler as an outgroup. The cladistic analysis produced two most parsimonious trees, from which a consensus tree was calculated. Well supported clades containing both sexual and asexual specimens were found in the tree, confirming the theory that asexual specimens of socalled species-pairs are not monophyletic. The concept of "species-pairs" (Poelt 1970, 1972) is well known and widely used in lichenology. It refers to closely related, morphologically indistinguishable lichens that differ from each other by their dispersal strategies only. The so-called "primary species" produces fruiting bodies and sexual spores, while its counterpart, the "secondary species," is vegetatively dispersed by soredia, isidia, or fragmentation. It has also been suggested that microconidia could function as asexual dispersal propagules to establish pycnidial anamorphs (Tehler 1988). Aside from the divergent dispersal strategies, such counterparts differ little if at all, morphologically, but may have different geographical distributions and ecological tendencies (e.g., Culberson 1973; Culberson & Culberson 1973). The taxonomic treatment of species-pairs differs from one lichenologist to another, but the two counterparts are traditionally treated and named as separate species. For thorough reviews, see Tehler (1982) and Mattson and Lumbsch (1989). Poelt (1972) suggested that the counterparts of speciespairs should be given species rank. Moberg (1977) used the terms primary and secondary species in the case of the genus Physcia. According to Culberson (1986), the asexual condition among lichens cannot have been constantly regenerated and further, the asexual morphs show such a uniform morphology that they can not have polyphyletic origin. Mattson and Lumbsch (1989) suggested that the sexual and asexual parts of species-pairs should not be treated as taxonomically different when sexual and asexual morphs are sympatric and intermediate forms occur. When having mostly allopatric distribution and intermediate individuals only at the overlapping distribution areas they would be regarded as separate subspecies. Allopatric species that do not have intermediate forms would not represent a real species pair. Mattson and Lumbsch also suggested that sterile lichens could be ancestors to new, sterile taxa. Other workers (e.g., Robinson 1975; Tehler 1982) suggested that the asexual forms of speciespairs may have a polyphyletic origin and should not be treated as taxa of their own. Robinson (1975) assumed that sympatric sorediate and fertile morphs could recombine with each other, because microconidia of the sorediate lichens might fertilize the ascocarp forming ones. Tehler (1982) suggested that the sorediate morphs of a species-pair could frequently arise from the fertile ones. This would explain the (e.g., chemical) variation between the 0007-2745/98/404-41 1$0.95/0 This content downloaded from 157.55.39.45 on Thu, 01 Sep 2016 05:51:38 UTC All use subject to http://about.jstor.org/terms 1998] LOHTANDER ET AL.: THE SPECIES PAIR CONCEPT 405 sterile counterparts of some species-pairs. According to Tehler (1982) the only case when an asexual morph could be considered as a species of its own, is when the clone has completely lost its ability to reproduce sexually, and no longer has living sexual relatives that might give rise to new asexual clones. Both Poelt (1970) and Tehler (1982) considered asexual morphs as evolutionary dead ends. Our goal in this study is to determine whether the asexual specimens of the species-pair Dendrographa leucophaea (Tuck.) Darb. evolved as a single evolutionary lineage separate from their corresponding sexual morphs, and thus are monophyletic; or if they have originated on several independent occasions and thus are polyphyletic. If the asexual stage had a single evolutionary history, asexual specimens would form a monophyletic clade in the phylogenetic tree. Until now, it has not been possible to experimentally verify any hypotheses concerning lichen species-pairs. This is the first attempt to examine them by applying cladistic methods to DNA sequence data. We have also examined whether gross morphological features were congruent with our molecular results. The genus Dendrographa Darb. contains two species, Dendrographa leucophaea and D. alectoroides Sundin & Tehler (Roccellaceae). Both are fruticose lichens that occur on trees, shrubs, and north-facing vertical rocks in the coastal habitats of California, U.S.A. (Sundin & Tehler 1996). Dendrographa leucophaea has a white, byssoid medulla; short microconidia; and complanate branches, while D. alectoroides has a brownish, coalescent medulla; long microconidia; and terete branches. A recent morphological study of the genus Dendrographa has been carried out by Sundin and Tehler (1996). Sexual specimens of both Dendrographa species have apothecia, while the asexual specimens disperse by thallus fragmentation. Neither soredia or isidia have been found (Sundin & Tehler 1996). Sundin and Tehler (1996) treated the asexual morphs as forma, but suggested that the terms anamorph and teleomorph as used in fungal taxonomy would be more appropriate (see also Tehler 1988). We have used the term forma for convenience, although it does not correspond to groups in a phylogenetic sense. Besides lacking ascomata, D. leucophaea f. minor differs from its sexual counterpart D. leucophaea f. leucophaea by having short internodes and more frequent lateral branchlets. It also occurs in more shaded microhabitats, and has a more restricted distribution than the sexual morphs. The sterile counterpart of D. alectoroides f. alectoroides is D. alectoroides f. parva. Both Dendrographa leucophaea and D. alectoroides sometimes have transitional morphs with ascomata and Pt. Reyes *

Abstract: In the Oregon Cascades, epiphytic cyanolichens are abundant in old-growth forest canopies, but they accumulate very slowly in young forests. We evaluated whether epiphytic cyanolichens require old growth and/or thick, underlying moss mats to achieve normal rates of growth and mortality. We transplanted over one thousand mature thalli of two old-growth associated species (Lobaria oregana and Pseudocyphellaria rainierensis) into the crowns of Douglasfir trees in thirteen forest stands representing four age classes: old-growth (400-700 yr), mature (140-150 yr), young (35-40 yr), and recent clearcut. Wooden racks were used instead of trees in the clearcuts. Half of the cyanolichen thalli were transplanted onto thick moss mats and half were transplanted onto bare bark. After one year, both species grew at least as well in younger forests as they did in old growth (20 to 30% increase in mass), but growth rates were significantly lower in clearcuts. Mortality rates were very low (<10%) in young, mature, and old-growth forests, but high (50 to 90%) in clearcuts. Pseudocyphellaria rainierensis grew significantly better on moss than on bare bark (30 vs 23% increase in mass). Since epiphytic cyanolichens can survive and grow in a broad range of forest age classes, silvicultural treatments that facilitate their colonization of young forests have great conservation potential. Epiphyte biomass increases slowly during succession in temperate forests (Esseen et al. 1996; Lesica et al. 1991; McCune 1993; Neitlich 1993; Rose 1992; Selva 1994). Old-growth forests support much higher epiphyte loads than younger, managed forests. In the Douglas-fir forests of Oregon's western Cascades, the most striking epiphytic difference between old-growth and managed forests involves cyanolichens. These nitrogen-fixing lichens dominate the old-growth forest canopy (Pike et al. 1975; Sillett 1995a), but are scarce or absent in younger forests (McCune 1993; Neitlich 1993; Spies 1991). Many epiphyte species, including several endemic cyanolichens, are seldom if ever, found in forests less than a century old (Rosentreter 1995; Sillett & Neitlich 1996). Conservation of old growth-associated epiphytes is becoming an important consideration in the Pacific Northwest, where most old-growth forest habitat has been destroyed by logging (Norse 1990). A recent emphasis on ecosystem management (Swanson & Franklin 1992) has stimulated efforts to promote epiphytes in managed forests. Recovery of epiphytic cyanolichens is particularly important because of their potential contributions to forest productivity. Nitrogen fixed by epiphytic cyanolichens can represent an important nutrient input in humid old-growth forests (Denison 1979; Pike 1978; Sollins et al. 1980). Epiphytes also constitute a significant part of biodiversity. For example, the number of lichens and bryophyte species often equals or exceeds the number of vascular plant species in old-growth forests (e.g., Lesica et al. 1991). The importance of epiphytic lichens has been recognized under a comprehensive new ecosystem management plan (FEMAT); populations of many oldgrowth associated species must now be monitored and protected (Rosentreter 1995). Reasons for the slow accumulation of epiphytic lichens in forests are poorly understood. Unsuitable microclimates in younger forests is one of several possible explanations (Sillett & Neitlich 1996). Microclimatic differences between old-growth and younger forests presumably arise from structural differences such as the frequency and size of gaps, abundance of standing dead material, and complexity of canopy architecture. Unlike more open, structurally complex old-growth forests, much of the canopy in dense, young, even-aged Douglas-fir forests, is shaded and sheltered from direct sunlight and precipitation (Kuiper 1988; Spies & Franklin 1991; Van Pelt & North 1996). Furthermore, bryophytes develop in thick mats on branches in large Douglas-fir trees, and many epiphytic cyanolichens are closely associated with these mats (Sillett 1995a). Perhaps these water storing mats ameliorate temperature and moisture fluctuations of epiphytes in their immediate vicinity, making these microhabitats more suitable to desiccation sensitive species (Lawrey 1991). 0007-2745/98/20-31$1.35/0 This content downloaded from 207.46.13.119 on Wed, 30 Mar 2016 05:39:46 UTC All use subject to http://about.jstor.org/terms 1998] SILLETT & McCUNE: CYANOLICHEN TRANSPLANTS 21 TABLE 1. Characteristics of study sites in the Willamette National Forest in western Oregon. Trunk diameter (dbh) and height are listed for the two Douglas-fir trees climbed in each stand. Stand number Age class Elevation (m) Estimated age Tree dbh (m) Tree height (m) 1 old growth 610 700 2.77, 2.08 68.0, 66.0 2 old growth 830 560 1.84, 1.65 82.5, 78.0 3 old growth 490 450 1.56, 1.41 74.0, 66.5 4 old growth 590 400 1.62, 1.31 71.0, 67.5 5 mature 890 150 0.87, 0.76 48.5, 49.5 6 mature 560 140 1.30, 1.26 59.5, 62.5 7 mature 650 140 1.02, 0.93 54.5, 48.0 8 young 780 40 0.40, 0.38 25.0, 23.5 9 young 760 37 0.46, 0.38 25.5, 20.0 10 young 500 36 0.33, 0.30 25.0, 26.0 11 clearcut 670 12 clearcut 460 13 clearcut 790 This study evaluates one explanation for the slow development of this critical component of Douglasfir forest--that epiphytic cyanolichens require the environment created by the structure of old-growth forest canopies. We also consider the possible effects of moss mats on cyanolichen growth and mortality in the forest canopy. Two cyanolichen species

Abstract: Phylogenetic analyses on Polytrichales were conducted using morphological characters as well as sequence data from the chloroplast genes rbcL and rps4 and the nuclear-encoded 18S rRNA gene. Our analyses included 22 species representing all genera of Polytrichales, plus eight outgroup species. Sequence data were obtained from 25, 22, and 19 taxa for 18S, rbcL and rps4 genes, respectively. Phylogenetic trees were constructed with parsimony analyses. Results lend support for recognition of Polytrichales as a distinct, monophyletic entity. After successively approximated weighting, Oedipodium griffithianum appears as the sister-group to Polytrichales. Within Polytrichales, Alophosia, Bartramiopsis, and Lyellia have the most basal placement outside a clade including all other genera. Atrichopsis, Dendroligotrichum, Itatiella, Meiotrichum, and Notoligotrichum are distinguished as a resolved monophyletic group while other genera are left as an unresolved entity. Resolution between these genera is achieved by successive weighting of data. After this, Dawsonia is resolved in the basal position within the clade and Polytrichastrum appears as a sister-taxon to Eopolytrichum and Polytrichum. Polytrichales are typically pioneer plants of open, sometimes even dry, habitats. Despite comprising only a small number of species, the order exhibits great diversity of shapes and sizes, from miniature plants such as Pogonatum piliferum (Dozy & Molk.) Touw of SE Asia and P. pensilvanicum (Hedw.) P. Beauv. of E North America to giants of Papua New Guinea like Dawsonia gigantea Geh. with stems reaching up to 80 cm. The most typical features of Polytrichales are the adaxial leaf lamellae and differentiation of leaves into a distinct blade and sheathing base. The hairy calyptra has given the group its name, although most genera have practically naked calyptrae. Sporophytes of Polytrichales normally have a well-developed peristome with at least 16 teeth that consist of whole cells. The epiphragm that covers the mouth of the capsule is a unique character that distinguishes most of the genera from all other groups of mosses. Size and shape of the urn vary greatly among genera (Schofield 1985; Smith 1971). The number of currently accepted genera in the Polytrichales is 19, and the approximate number of species in each genus is given in Table 1. One genus, along with its sole species Eopolytrichum antiquum Konopka, Herendeen, Merrill & Crane, is known only from beautifully preserved late Cretaceous fossils that reveal its structures in fine detail (Konopka et al. 1997). Many of the remaining genera are monotypic and all the others, with the exception of Pogonatum and Polytrichum, are fairly small. Schofield (1985) gives a conservative estimate of about 370 for the number of species in Polytrichales, but on the basis of recent critical revisions (e.g., Hyvinen 1989) it is realistic to assume that the number is actually closer to, or even less than, 200. Some species, like Polytrichum juniperinum Hedw., are almost cosmopolitan, while others, like the Macaronesian Alophosia azorica (Ren. & Card.) Card., are narrow endemics and possibly 1 This paper was presented at the 1997 Montr6al ABLS symposium sponsored by the Green Plant Phylogeny Research Coordination Group (with funding provided by DOE/NSF/USDA Panel on Collaborative Research in Plant Biology, USDA grant 94-7105-0713, Co-Pls Mark A. Buchheim, Brent D. Mishler, Russell L. Chapman). 0007-2745/98/489-504$1.75/0 This content downloaded from 207.46.13.162 on Fri, 01 Jul 2016 05:45:45 UTC All use subject to http://about.jstor.org/terms 490 THE BRYOLOGIST [VOL. 101 TABLE 1. The approximate number of species in the genera of Polytrichales. t-fossil. Alophosia 1 Meiotrichum 1 Atrichopsis 1 Notoligotrichum 10 Atrichum 15 Oligotrichum 10 Bartramiopsis 1 Pogonatum 50 Dawsonia 10 Polytrichadelphus 10 Dendroligotrichum 2 Polytrichastrum 10 t Eopolytrichum 1 Polytrichum 30 Hebantia 1 Psilopilum 2 Itatiella 1 Steereobryon 1 Lyellia 4 even threatened by extinction. Ecologically, the Polytrichales range from xerophytes like Polytrichum piliferum Hedw. to species of peaty, wet habitats like P. commune Hedw. Although their structure appears so obviously adapted to dry environments, the Polytrichales are largely absent from extremely arid regions, and the group exhibits greatest diversity in areas with humid or moist climates like SE Asia and Central America-northern South America. Phylogenetic relationships of the Polytrichales are particularly relevant to considerations of moss evolutionary history since the group is probably among the first of the lineages that diverged from the common ancestor of all mosses (Mishler & Churchill 1984). Smith (1971) presented a dendrogram with assumed phylogenetic trends for Polytrichales, and these ideas were further developed in his study of epiphragm structure and spore ornamentation (Smith 1974). Hyv6nen (1989), in a revision of Pogonatum that included cladistic analysis based on manual Hennigian argumentation, tentatively distinguished three entities in the Polytrichales as an unresolved basal trichotomy. These, and many other authors, viewed Polytrichastrum G. L. Sm. as the closest extant approximation to the common ancestor of Polytrichales. Forrest (1995) presented the first computer-aided analyses of Polytrichalean phylogeny based on a matrix of 50 characters compiled from the literature. The cladogram from her successively weighted analysis of these data is presented in Figure 1 (fig. 3b in Forrest 1995). In contrast to the views noted above, the position of Atrichum as sister to the remaining Polytrichales suggests that relatively simple members of the group more closely approximate the ancestral condition. This is congruent with ideas presented previously by Fleischer (1923). Forrest's (1995) analyses resolved both strongly and weakly supported groups as implied by the number of characters supporting each clade. To test the strength of the phylogenetic hypotheses based on morphology and to determine whether they are congruent with other sources of data we explored Tetraphis

Abstract: This paper includes 19 saxicolous species of Caloplaca from North and Central America that are characterized by distinctive elongated marginal lobes, non-paraplectenchymatous hypothecium, and presence of anthraquinones. In various ways this group has been referred to as Caloplaca section Gasparrinia or even separated at the generic level. The group, however, is not regarded as natural and consequently not given taxonomic status. Descriptions, keys, and distribution maps are provided for all treated taxa. Three new species are described for the region: Caloplaca appressa, C. stellata, and C. texana. Several lectotypes are selected. Caloplaca augustina is a new synonym of C. galactophylla. Color illustrations are provided for most species or references to previously published color illustrations. Additional notes are presented for six related non-American species.

Abstract: Effects of UV-B radiation on fine structure, carbohydrates, and pigments in Polytrichum commune.

Abstract: Three species of liverwort and one species of moss were tested for their ability to tolerate desiccation and subsequent liquid nitrogen (LN) exposure. Riccia fluitans and Helicodontium cappelare were sensitive to desiccation and required either abscisic acid (ABA) or encapsulation in alginate beads with a 0.75 M sucrose pretreatment for 100% survival through drying and LN exposure. There was little effect ofABA on Plagiochila sp. although drying and LN exposure were successful with encapsulation. With Marchantia polymorpha optimal survival through drying and LN exposure required both ABA and encapsulation. These results suggest that encapsulation cryopreservation, in some cases in combination with ABA preculture, might be used as a method for long-term germplasm storage for desiccation sensitive bryophytes. With the decline in the area of natural plant habitats, ex situ preservation can be used as a supplemental tool for maintaining plant germplasm, offering insurance against the possibility of loss of organisms from the wild. While most ex situ preservation programs emphasize higher plants, there is a concurrent need to preserve the germplasm of nonseed plants, including bryophytes (Soderstrom et al. 1992). In the case of hepatics and mosses, there are a number of species which are considered endangered (Geissler et al. 1997; Tan et al. 1994a,b). The fact that biochemicals with antibiotic, neurotropic, insect antifeedant, and antitumor activity have been isolated from various species adds further weight to the argument for maintaining and exploring bryophyte germplasm (Asakawa et al. 1980; Fukuyama & Asakawa 1991; Joshi & Desai 1989; Spjut et al. 1992). Cryopreservation offers the potential for longterm maintenance of living tissues. Techniques for the preservation of cells, shoot tips, embryos, and other tissues in liquid nitrogen (LN) extend the concept of germplasm banking beyond the traditional banking of seeds and spores. Such ex situ preservation of rare plant germplasm can provide a resource for research as well as for reintroduction if a species is lost in the wild. In cases where tissues can survive desiccation, direct exposure to LN can be used successfully, for example, as with many seeds (Pence 1990; Stanwood 1985). If the tissues are sensitive to desiccation, other protective measures are needed, and a variety of techniques have been developed for cryopreserving plant tissues (Bajaj 1991). The following experiments were undertaken to examine the applicability of cryopreservation to the gametophytes of four in vitro grown bryophytes three liverwort species and one tropical moss species. Open drying and encapsulation dehydration were compared as protective measures, and the effect of preculture with abscisic acid (ABA) that has been implicated in conveying desiccation tolerance in other systems (Werner et al. 1991), was also ex-

Abstract: Samples of the epiphytic lichen Evernia prunastri collected in an unpolluted area were transplanted to the urban environment of Teramo (central Italy) The influence of the transplant process on trace element content was found to be negligible and after only two months, concentrations of all elements (Cd, Cr, Cu, Pb, and Zn) were significantly higher than in control samples It is suggested that motor traffic was the main source of atmospheric Cr, Cu, and Pb, while for Cd and Zn additional sources from phosphate fertilizers and pesticides used in the surrounding farmlands are hypothesized The possible uptake and accumulation processes of these metals in relation to time of exposure are discussed Lichens are known for their ability to accumulate airborne substances in concentrations far greater than those in the atmosphere (Nieboer & Richardson 1981) Accumulation at ion exchange sites and particulate trapping are two well documented mechanisms by which elevated levels of elements are achieved in lichens (Nieboer et al 1978) Metal cations bind to extracellular anion exchange sites in the cell wall and on the plasma membrane surface (Brown & Beckett 1985) Because cell wall-bound elements are readily exchangeable, extracellular quantities and proportions reflect recent environmental inputs (Brown 1987) However, when elements occur in insoluble particulate matter, they cannot be exchanged through the cell wall and accumulate in the thallus in relation to the environmental availability of the particles and exposure time ie, lichen age (Brown 1987) This last point was documented by Bargagli et al (1987), who found that the central (older) part of the thallus of foliose lichens has higher concentrations of certain metals than the peripheral (younger) part However, elements present in trapped particulates may be dissolved in water or solubilized by the lichen to some degree, and metal ions released by this process may have several fates eg, become potentially capable of occupying cation bind sites in the cell wall or be taken up intracellularly (Kershaw 1985) According to Loppi et al (1997), elements of limited metabolic significance (eg, Al, Pb) have higher concentrations in the central parts, where they are presumably trapped in the medulla, whereas e ements essential for lichen metabolism (eg, Co, Mo, Zn) have higher concentrations in the peripheral parts, but are easily displaced from one part of the thallus to another Nevertheless, the different me abolic fates of the various metals suggest that lichens may selectively accumulate those elements that remain extracellular, whereas those elements that enter into the cell may be metabolized and eliminated or may lead to death of the lichen (Kershaw 1985) In the present paper, the results of a transplant experiment with Evernia prunastri are reported The objective of the study is to investigate the uptake over time of Cd, Cr, Cu, Pb, and Zn after lichen samples were transplanted to an urban envi-

Abstract: A bioclimatic model based on bryophyte species distribution and abundance relative to climate was coupled with climatic and geographical data obtained from Leemans and Cramer (1991) and the Canadian Climate Center (CCC) General Circulation model (GCM) at 1XCO2 in order to reconstruct the present geographical distribution of seven peatland types in the Mackenzie River Basin. The geographical distribution of 195 peatlands previously identified by type were used to test the validity of the reconstructions. The test revealed that the reconstruction using data from Leemans and Cramer was more accurate than the reconstruction using the CCC GCM data. For this reason, the CCC 1XCO2 data was subtracted from the CCC 2XC02 climatic data to produce an anomalies data set which was then added to the Leemans and Cramer data to project the distribution of the seven types of peatlands at 2XCO2. Results of this prediction were then compared to predictions using 2XC02 data obtained from the Geophysics Fluid Dynamics Laboratory (GFDL) GCM. The position of the southern limits of peatland distribution was compared to past distributions resulting from a warming period in the early to mid Holocene. Results of the predictions for the two climate change scenarios indicated a northward migration of the southern boundary of peatland ecosystems of approximately 780 km in the central portion of the Mackenzie River Basin. The model also predicted that Mid-Boreal peatlands would be located along a diagonal running from southeast to northwest from 600 longitude to an area just south of the Mackenzie Delta for both scenarios. High-Boreal and Subarctic peatlands were located to the north of the diagonal, while Low-Boreal peatlands were located to the south. However, the CCC anomalies + Leemans and Cramer predictions did not clearly define the Low-Boreal since Low-Boreal indicators were only located in the Cordilleran Ecological Province. Ecological diversity is anticipated to be maintained in the peatlands because all types were predicted to be found in the Basin at 2XC02 but at different locations. Comparisons between the predicted position of the southern limits of peatland distribution and that during the early to mid Holocene indicate that the model's predictions were reasonable. Global warming resulting from increased concentrations of such greenhouse gases as CO2, CH4, and NOx could produce temperature increases as much as 1.5 to 4.60C by the middle of the 21st century (Mitchell et al. 1990). Northern and continental regions in particular will experience significant warming, especially during the spring and winter months. Increases in annual temperature for northern locations in North America are anticipated to be in the order of 3.7 to 4.60C (Cohen 1993). Elevated temperatures are expected to produce droughts that will affect site water balance and runoff, and cause major shifts in the distribution of ecosystems on a global scale (Houghton et al. 1990; Vitousek 1994). These effects will be quite noticeable in such ecosystems as wetlands, and in particular peatlands, which are especially sensitive to climate and fluctuations in water levels. Peatlands are wetlands that have a minimum accumulation of 0.5 m of peat, and are usually divided into two types: bogs and fens. This division is based on whether the water supply is ombrotrophic (bogs) or minerotrophic (Gore 1983; Sjors 1952). The largest concentration of peatlands is in the boreal and subarctic regions of the Northern Hemisphere (Gore 1983). The northern distribution indicates that peatlands are climatically sensitive ecosystems and that they are primarily found in areas that have wet and humid climates where there are no sustained dry periods (Moore & Bellamy 1974). 0007-2745/98/572-587$1.75/0 This content downloaded from 207.46.13.128 on Tue, 06 Sep 2016 05:18:25 UTC All use subject to http://about.jstor.org/terms 1998] GIGNAC ET AL.: PREDICTED MIGRATION OF PEATLANDS 573 Peatland formation is a function of two climatic variables, precipitation and evaporation, and generally they occur where evaporation does not exceed precipitation i.e., where potential evapotranspiration ratios < 1 (Gignac 1993). Not only does the macroclimate affect the presence of peatlands, but it also affects the distribution of different forms of bogs, and the height that they attain above the surrounding landscape (Ivanov 1981; Moore & Bellamy 1974). Among climatic variables that have been correlated to the different types of bogs are: total precipitation (Eurola 1962; Ruuhijirvi 1960), number of precipitation days (Taylor 1983), and precipitation and temperature together (Ivanov 1981; Moore & Bellamy 1974). Fens can also be found in all climatic zones where peatlands occur, and some types of fens can be placed into different zones based on climatic factors (Gignac & Vitt 1990; Moore & Bellamy 1974). Peatland vegetation, particularly bryophyte species, are very sensitive to changes in the height of the water table (Gignac et al. 1991; Gorham & Janssens 1992; Nicholson et al. 1996). The height of the peat surface above the water table and thus the height of the vegetation are a function of climatic conditions, particularly temperature and precipitation (Gignac & Vitt 1990). Increases in temperature enhance evaporation and if precipitation remains constant or diminishes, it should cause a lowering of the water table. A decrease of the height of the water table will result in subtle changes in the presence, absence, and abundance of bryophyte species (Andrus et al. 1983; Gignac & Vitt 1990; Gignac et al. 1991; Vitt et al. 1975). Thus, peatland vegetation is very sensitive to climate. Changes in peatland vegetation should offer an early warning of large-scale climatic changes before they occur in more stable and larger ecosystems such as the boreal forest. Because peatlands are sinks and sources of the greenhouse gases CO2 and CH4, any modification of the climate could affect the quantities of these gases released to the atmosphere (Bubier et al. 1993; Roulet et al. 1993; Roulet et al. 1994). Accumulating deposits of peat retain water causing local water tables to rise (Ingram 1983) which has lead to large scale peatland formation through paludification in northern landscapes (Nicholson & Vitt 1990; Sjors 1963). Under a warmer drier climate methane emissions will initially increase with melting of permafrost and rising of water tables (Liblik et al. 1997). Once new drainage patterns establish, increased oxidation of the peat will release more CO2 to the atmosphere (Gorham 1991). Organic acids are also produced from decomposing mosses and as a result of increased decomposition caused by warmer climates, local streams may become acidified (Bayley et al. 1992; Hogg et al. 1992). The microclimate of the peatland will also change causing a reduction of evapotranspiration rates because of the shift from areas with free standing water to shaded areas as a result of increased tree or shrub cover (Ingram 1983). Subsurface soil temperatures will also increase with the resulting degradation of permafrost in many north-

Abstract: Maintaining numbers of aseptic cultures of bryophytes by serial transfer not only consumes supplies and labor but can result in loss of vigor or the appearance of other abnormalities in long-term cultures. Cryopreservation of cultures solves the problems associated with long-term culture, and is of particular importance for cultures or mutant cell lines incapable of making spores or where spores are short-lived. Dramatic simplification of a published protocol for moss cryopreservation is accomplished by preconditioning cultures for 3-4 days in medium supplemented with 10-5 M ABA and 100 mM proline. This proline/ABA protocol can be successfully used with Ceratodon purpureus, Funaria hygrometrica, Physcomitrella patens, and two species of Sphagnum; cryopreserved cultures remain viable for a minimum of one year at -800C. Experiments demonstrate a differential response by cell types to cryopreservation, as well as differences between species or for particular selected cell lines within a species. Other experiments suggest that successful cryopreservation is not a simple consequence of brood cells in ABA-treated

Abstract: Leucobryum glaucum and L. albidum are generally distinguished by quantitative differences in plant height, leaf length, and transverse sectional leaf anatomy. Although extremely small plants can be readily identified as L. albidum, and large plants can be confidently assigned to L. glaucum, intermediate forms are common and many identifications are arbitrary. We amplified approximately 815 bp of the nuclear ribosomal DNA ITS from plants varying in size, and cut the products with three restriction endonucleases (Hhal, Hinfi, and Taql). Two DNA haplotypes were detected in a sample of 23 plants growing at a forested site in Durham, North Carolina. All plants with leaves 5.0 mm in length or shorter had one haplotype, and all plants with leaves longer than 5.0 mm had the other haplotype. Our results support the interpretation that L. albidum (small plants) is genetically discontinuous with L. glaucum (large plants), at least in the limited area from which we sampled. Leucobryum is one of approximately eight genera in the Leucobryaceae (Yamaguchi 1993), which is comprised of acrocarpous mosses with haplolepideous peristomes and multistratose leaves. Members of the family are separated from the Dicranaceae on the basis of their distinctive leaf structure. The leaf costa in leucobryoid species, which comprises most of the leaf, consists of a central layer of small, often diamond-shaped chlorophyllose cells (chlorocysts) and several layers of hyaline cells (leucocysts) above and below the chlorocysts. Leucobryum includes approximately 180 species (Brotherus 1924; Yamaguchi 1993), some of which are common components of temperate and tropical forest communities. Leucobryum species of Europe and North America are often called "pin cushion" mosses because of the rounded colonies they form. Two or three species of Leucobryum occur in eastern North America (Anderson et al. 1990; Crum & Anderson 1981). Leucobryum glaucum (Hedw.) Angstr. ranges from Newfoundland west to Manitoba, and south to the Gulf of Mexico. The species also occurs abundantly across northern Europe and Asia, east to Japan (Yamaguchi 1993). The other North American species, L. albidum (Brid.) Lindb., is restricted to the eastern United States and Canada, and occurs in the Caribbean region and Central America (Crum & Anderson 1981). A third species, L. antillarum, has been reported from the Gulf coast region of the United States, but Crum & Anderson (1981) exclude it from the flora, arguing that North American plants attributed to this species cannot be distinguished from L. glaucum. Nevertheless, Anderson et al. (1990) included L. antillarum in their most recent checklist of North American mosses, and the problem requires further study. Leucobryum glaucum and L. albidum have been distinguished by size of the gametophytes and the number of leucocyst layers below the single layer of chlorocysts, as seen in transverse sectional view at or near the leaf base (Crum & Anderson 1981). The larger species, L. glaucum, is said to form deeper cushions (2-9 cm high according to Crum & Anderson 1981), and the leaves are 3-8 mm long, with 3-4 layers of leucocysts below the chlorocysts. In contrast, L. albidum is described as forming cushions rarely more than 1 cm high, with leaves less than 4 mm long and having only 2-3 layers of leucocysts below the chlorocyst layer. Sporophytes are considerably more common on smaller plants that correspond to L. albidum than on larger plants that fall within the range of morphological variation generally attributed to L. glaucum. The reason(s) for this difference in sporophyte formation has not been studied. Both species are dioicous and are reported as having dwarf males that grow epiphytically among the perichaetial bracts of the females (Crum & Anderson 1981; Smith 1978; Yamaguchi 1993). In L. glaucum, male plants vary in size from extremely dwarfed and epiphytic to only slightly smaller than the females (Yamaguchi 1993). The situation is less clear in L. albidum because no systematic studies have been conducted, but it has been suggested that the males 0007-2745/98/272-277$0.75/0 This content downloaded from 157.55.39.95 on Sat, 11 Jun 2016 05:04:58 UTC All use subject to http://about.jstor.org/terms 1998] PATTERSON ET AL.: DNA OF LEUCOBRYUM 273 5'>TCGATGAAGAACGCAGCG TCCGTAGGTGAACCTGCGG<3'

Abstract: One hundred-twelve species of lichens are reported for the first time from Arizona. The new combination Lecanora muralis var. brunneola (Mereschk.) Ryan & Nash is made, and the variety is reported as new to North America. The following 19 taxa are also reported for the first time from North America, north of Mexico: Absconditella lignicola, Caloplaca brouardii, Hypotrachyna dactylifera, Lecanora cavicola, L. plumosa, L. swartzii, Mycocalicium victoriae, Pertusaria moreliensis, Placidium pilosellum, Pyxine meissneriana, Rhizocarpon arctogenum, Um- bilicaria subglabra, Verrucaria obductilis, V. trabicola, Xanthoparmelia australasica, X. consociata, X. isidiosa, X. neocongensis, and X. tegeta.

Abstract: A model was developed that classified and projected the distribution of seven different types of peatlands in the Mackenzie River Basin. The model was based on the relationships between bryophyte indicator species, the types of peatlands they characterize, and regional climate. The model used the presence, absence, and abundance of 15 bryophyte indicator species to classify 81 peatlands in the study area into seven groups. Abundance values were calculated for each of the indicator species along three climatic gradients-Mean Annual Temperature (MAT), Mean Annual Total Precipitation (MATP), and Length of the Growing Season (LGS). The percent cover of all species were then ascribed to appropriate combinations of MAP, MATP, and LGS. The result produced a matrix consisting of 4,560 grid nodes where each node was identified by values for each of the three climatic variables and the types of peatlands that could be found at that climate. An independent data set consisting of climatic and ecological values and vegetation cover for 115 sites was used to test the ability and accuracy of the model to classify and project the climatic distribution of the seven peatland groups. The model correctly classified 106 of the 115 sites and of those, correctly projected the distribution of all but five of the test sites. The model accuracy was 70% for six of the seven groups, and > 90% for three of those. The accuracy for the remaining group was 50% and errors were mostly caused by the failure to project the distribution of three of the test sites. Other errors include: the inability to classify lichen dominated peatlands; the inclusion of wet lawns in bogs into one of the groups which caused a southward extension of that group by approximately 200 km. The overall model accuracy was 88%. Peatlands, primarily bogs and fens, cover extensive areas of the boreal and subarctic zones particularly in the northern hemisphere (Gore 1983). Several studies have demonstrated that peatland types and peatland vegetation are to a large extent controlled by macroclimate, physiology, and local gradients. The macroclimate plays an important role in the regional distribution of the different types of peatlands (Moore & Bellamy 1974). Among climatic variables that have been correlated to the different types are total precipitation (Eurola 1962; Ruuhijiirvi 1960), number of precipitation days (Taylor 1983), and precipitation and temperature together (Ivanov 1981; Moore & Bellamy 1974). The present distribution of peatlands in North America is a function of two climatic variables, precipitation and potential evaporation. Generally peatlands only occur in areas where evaporation does not exceed precipitation (Gignac 1993). Because bryophytes have a high fidelity to such environmental gradients as moisture and pH, they are particularly good indicators of different types of peatlands at the local level (Gignac & Vitt 1990; Horton et al. 1979; Nicholson et al. 1996; Vitt et al. 1975; Vitt & Slack 1975, 1984). The distributions of several of those species are also limited by climate and thus become indicators of climate. For example, in western North America the distribuion several such species as Sphagnum austinii, S. papillosum, S. rubellumn and S. tenellum are limited by climate to areas that have > 1,000 mm total precipitation and are thus indicative of oceanic peatlands (Gignac et al. 1991a). Others, such as Drepanocladus fluitans, Sphagnum lenense and S. riparium, are limited to areas having mean annual temperatres lower than O0C and are indicative of high boreal and subarctic peatlands (Nicholson & Gignac 1995). Also, several geographically widespread species can be indicators of peatlands based not on their presence and absence but on their abundance. The primary division of peatlands into bogs and 0007-2745/98/560-571$1.35/0 This content downloaded from 157.55.39.225 on Tue, 13 Jun 2017 18:01:52 UTC All use subject to http://about.jstor.org/terms 1998] GIGNAC ET AL.: PRESENT DISTRIBUTION OF PEATLANDS 561 fens is based on the source of surface water. The supply of water in ombrotrophic peatlands (bogs) is entirely from precipitation and because of this, the pH of bog surface water is low (Sjirs 1952). In contrast, a portion of the surface water on minerotrophic peatlands (fens) has been in contact with mineral soil. Depending on the nature of the mineral substratum and the quantity of water in the peatland that has flowed over the substratum, the pH of surface water in fens can vary considerably. Bogs are usually classified according to their landform patterns into raised bogs, plateau bogs, eccentric-domed bogs, concentric-domed bogs, and blanket bogs (Glaser & Janssens 1986; Moore & Bellamy 1974). Fens are usually divided according to the species richness of the vegetation into poor and rich (Sjirs 1952). Associated with species richness are differences in pH of the surface water: poor fens have pH values usually below 5.6, while rich fen surface water pH values are usually circumneutral or higher. There are further subdivisions of the rich fen category into moderate and extreme, again depending on the species richness of the vegetation (Sjors 1952, 1963). Other types of fens are aapa and palsa fens, which are classified according to landform and can be either rich or poor. Aapa fens have patterns composed of strings (ridges) and flarks (pools), while palsa fens have mounds (palsas) that contain a core of permafrost (Moore & Bellamy 1974). The distribution of the different types of bogs is related to macroclimate (Moore & Bellamy 1974). Ivanov (1981) for example, determined that the maximum height of a bog is in part a function of the moisture surplus during the driest month. Moisture surplus is the excess of precipitation over evapotranspiration (Damman 1986). The convexity of the dome is also a function of the water balance for the driest month (Ivanov 1981). The surface of bogs in hyperoceanic and oceanic areas may be raised several meters above the water table because there is always a large monthly moisture surplus (Gignac & Vitt 1990). Conversely, continental bogs are only slightly raised above the water table because there are several months during the growing season when there is relatively little precipitation. Although fens can be found in all climatic zones where peatlands occur, some types of fens can be placed into different zones based on climatic factors. Aapa (string) fens are found in continental areas where precipitation is low and peatlands form in basins and are thus more susceptible to influence by mineral soil water. Palsa fens are located in high boreal and subarctic regions that have discontinuous permafrost (Moore & Bellamy 1974). Generally, rich fens, particularly extreme-rich fens, are located in subcontinental and continental areas and are rare or absent in hyperoceanic and oceanic areas (Gignac et al. 1991b; Gignac & Vitt 1990; Vitt et al. 1990). The Mackenzie River Basin provides an excellent geographical area in which to study the effects of cl mate on the distribution of peatlands. It is a large basin that is restricted to continental areas and has wide variations along several climatic gradients, particularly temperature. Also, and most importantly, its southern boundary generally corresponds to the southern limit of peatland development and, with a few exceptions, its northern boundary coincides with the northernmost development of extensive peatlands in western Canada. A previous analysis of the vegetation of peatlands in the Mackenzie River Basin (Nicholson et al. 1996) indicated that the distributions of several peatland types and their bryophyte indicator species were closely related to climate. Those results led to the development of a model that reconstructs the present distribution of peatlands in the Mackenzie River Basin based only on climatic variables. The purpose of this study is to delimit the climatic distribution of each type of peatland that was found in the study area. The following steps were used to model peatland distribution 1) relate bryophyte indicator species to different types of peatlands and the climate in the Mackenzie River Basin; 2) project the present climatic distribution of those types of peatlands; and 3) test the model's validity and projections using an independent data

Abstract: Entosthodon durieui Mont. and E. pallescens Jur., two species with Mediterranean distributions are compared and are identical. A description and illustrations of E. durieui, incorporating spore morphology, are given. Entosthodon durieui was described by Montagne (1849) and E. mustaphae by Trabut (1886), both species from Algeria. They have also been reported from Israel and Spain. Bizot (1945) considered E. mustaphae synonymous with E. durieui. Allorge (1960) published descriptions and illustrations of both taxa. Juratzka (in Unger & Kotschy 1865) described Entosthodon pallescens from Cyprus, but details about calyptra and lid were not provided. Subsequently, this species has been recorded from many Mediterranean countries and islands (Balearic Island, Crete, Egypt, Greece, Italy, Israel, Sicily, Turkey, and Spain), central Asia, and Tenerife. Casares-Gil (1915) reduced E. physcomitroides CasaresGil & Beltrin reported from the Iberian Peninsula (Casares-Gil & Beltriin 1912), to a variety of E. pallescens. Loeske (1929) suggested the epithet mitratus because of its mitriform calyptra, even though details about calyptra were not provided in the original description of E. pallescens. It seems to have been understood that E. pallescens had a cucullate calyptra, which corresponds to the characteristic shape found in the genus Entosthodon; however, the presence of a mitriform calyptra can be found in the description of E. pallescens by Abramov et al. (1989). Comparative studies of E. durieui and E. pallescens have only been carried out by Loeske (1929) and Bilewsky (1965). Loeske doubted that there is any difference between these two species. Bilewsky tried to distinguish between them, but his conclusions are ambiguous and contradictory. Loeske also related E. durieui and E. mustaphae with E. commutatus Dur. & Mont., a species also described from the north of Africa. Thus, these three species appear as synonymous in Index Muscorum. In the Israel checklist (Herrnstadt et al. 1991), E. commutatus, E. durieui, and E. mustaphae are considered synonyms of E. attenuatus (Dicks.) Bryh. In the distribution maps of the bryophytes of the Iberian Peninsula (Casas et al. 1996), E. pallescens, E. durieui, and the varieties mitratus and mustaphae, have been shown in one single map, due to difficulties in distinguishing them. C reful research shows that E. durieui, E. commutatus, and E. attenuatus are not synonymous. Entosthodon commutatus has leaves tapering to long apiculate apices and long-lanceolate peristome teeth; these features serve to clearly distinguish it from E. durieui. Additionally, E. attenuatus can be differentiated by its cerise rhizoids, which are light brown in other taxa (Fife 1987). It is also worthwhile noting that E. durieui differs from other closely related species from the north of Africa which are included in the genus Funaria, such as F. saharae Trab., F. deserticola Trab., and F. nilotica Broth. Long lanceolate peristome teeth have been observed in the above species. Funaria mouretti Corb., from the North of Africa too, has denticulate leaves with excurrent nerve. Specimens classified as E. pallescens or as E. durieui present a set of characters that are not found in other species of the genus Entosthodon. The leaves are entire, elliptic to obovate or spathulate, marginal cells not differentiated, with a weak short nerve and an acute to obtuse apex. The pyriform capsule has a neck as long as the sporangium, the peristome teeth are rudimentary and short, with only 2 or 3 cell segments overhanging the capsule mouth. The spores are coarsely baculate-insulate and the mature calyptra has inflated and lobed bases with 3 or 4 slits. Entosthodon durieui and E. pallescens share the same habitat. All specimens have been collected on calcareous rocks and artificial walls in dry environments at low elevations. We conclude that E. durieui and E. pallescens are the same taxon. The name E. durieui takes priority. Entosthodon durieui belongs to the subgenus Entosthodon (Fife 1985): the exothecial cells are oblong with thick, radially cuneate walls and the mouth has the same diameter as the moist capsule. However, the calyptra is mitrate, as characteristic for the genus Physcomitrium. Spores are coarsely baculate-insulate, similar to those of E. tucsonii, which is considered by Fife (1985) as a species with anomalous spores, but rightly placed in subgenus Entosthodon. This ornamentation pattern is 0007-2745/98/133-136$0.55/0 This content downloaded from 207.46.13.109 on Wed, 22 Feb 2017 19:02:57 UTC All use subject to http://about.jstor.org/terms 134 THE BRYOLOGIST [VOL. 101

Abstract: Six species of the lichen genus Strigula are reported from New Zealand: S. delicata Sdrus. sp. nov. S. fossulicola P. M. McCarthy et al., S. nemathora Mont., S. novae-zelandiae (Nag Raj) Sirus. comb. nov. (Basionym: Discosiella novae-zelandiae Nag Raj), S. oceanica P. M. McCarthy et al., and S. subtilissima (Fde) Mill. Arg. In 1983, the late J. K. Bartlett deposited a large set of foliicolous lichens he had collected in New Zealand in the LG herbarium. Although poorly curated, the material was prolific and has yielded several new taxa: the type collection of Enterographa bartlettii S6rus. (1984), Badimiella serusiauxii Malcolm & V6zda a monotypic genus with spectacular campylidia (Malcolm & V6zda 1994; S6rusiaux 1986), and the recently described Strigula kaitokensis S6rus. & Polly (1996). For some time, I was aware that several other undescribed species of Strigula, not mentioned in the monograph of Santesson (1952), were present in the material. Two have now been described by McCarthy et al. (1996) from Lord Howe Island (Australia) and are here recorded for the first time from New Zealand; another species appeared to be identical with the anamorphic lichenized fungus Discosiella novae-zelandiae Nag Raj (1981) and is here transferred to Strigula, and an additional one is here described as new. A new species of Mycomicrothelia, amazingly similar to a genuine Strigula, has also been found in the collections and is described as new in a separate paper (Sdrusiaux & Aptroot 1998). New Zealand appears to be rich in species of Strigula, with several new species and new records reported (Hafellner & Kalb 1995; Harris 1995; McCarthy 1995; McCarthy & Malcolm 1996; S6rusiaux & Polly 1996). However, the genus is still poorly known at the world level so it is too early to claim that New Zealand is a center of diversity. More work is needed before a global assessment can be made. The taxonomic concept of the genus Strigula has been reorganized by Harris (1975, 1995) to include all foliicolous species referred by Santesson (1952) to that genus, Raciborskiella Hohnel, the so-called Porina phyllogena group (= Phylloporis Clem.), and several corticolous and saxicolous species, previously included in Arthopyrenia Massal. and Porina Mtill. Arg. All these species have the same type of asci, paraphyses, and conidia. This concept is accepted here as such a genus is clearly monophyletic, even if one expects that further detailed studies may lead to the segregation of several smaller genera within that clade. All collections mentioned in this paper have been gathered by the late J. K. Bartlett (t1986) and are housed in LG. Representative samples are deposited in B, CANB, CHR, and herb. R. Luicking. STRIGULA DELICATA Sdrusiaux sp. nov. (FIG. 1-2 & 7, a-b) Thallus epiphyllus, subcuticularis, plus minusve circularis, 2-5 mm diam., pallide viridis vel olivaceo-brunneus, nitidus, laevigatus, lobis angustis (ca 0.1 mm latis) et dichotomis formatus, interdum connatis et reticulum formantibus; margo cum continua vel interrupta, nigra linea et numerosis, parvulis, elongatis nigrisque papillis. Algae ad Cephaleuros pertinentes. Perithecia in parvis, circularibus thalli maculis crescentia, ad marginem locatis, hemisphaerica, 0.2-0.25 mm diam. et 0.10-0.18 mm alta, nigra et nitida; paraphyses numerosae, ramosae et leviter anastomosantes; asci clavati, 55-60 x 10-14 pCm; ascosporae 8/ascus, biseriatae, ellipsoideae ad fusiformes, 1-septatae, 15-16(-18) x 4.5-5.0 pm, pariete crasso (0.7-1.0 pEm). Macroconidia oblonga, 1-septata, appendicibus mucosis, 3-5 Ipm longis et terminalibus praedita, 13-16 X 3.0-3.5 Ipm; microconidia ellipsoidea, ca 3 X 1 im. Thallus epiphyllous, subcuticular, occurring or not along the nerves, margins, and scars of leaves, ? circular, 2-5 mm in diam., usually quite abundant, pale green or olive brown (white when dead), shiny but without any metallic tint, with smooth surface, formed of dichotomously branched, narrow (ca 0.1 mm wide) lobes that can fuse laterally and form a reticulum; lobes margins with discontinuous or continuous black line and usually with numerous, rather regularly scattered, small, elongate, black papillae. Photobiont a species of Cephaleuros (Trentepohliaceae), with ? rectangular or rounded cells, 8-15 x 4-7 pm, yellowish green, + arranged in radiating rows. Perithecia usually present and abundant, typically occurring in center of small, circular (0.3-0.6 mm in diam.) patch of thallus with small and rounded lobes located at or near lobes tips, single or paired (rarely in 3s), typically hemispheric, never (ob-)conical, with rounded apex, 0007-2745/98/147-152$0.75/0 This content downloaded from 157.55.39.107 on Wed, 30 Mar 2016 06:34:55 UTC All use subject to http://about.jstor.org/terms 148 THE BRYOLOGIST [VOL. 101

Abstract: The phylogeny of the genus Diploschistes is investigated using parsimony analysis with morphological, chemical, and ecological characters. The scruposus group, including D. ocellatus, is supported as monophyletic, whereas the actinostomus group is shown to be paraphyletic. The predicted trend from perithecioid to urceolate ascomata within the genus is supported. The lichen genus Diploschistes Norman is a small genus in the Thelotremataceae Zahlbr. containing approximately 30 species (Lumbsch & Guderley 1996). It is usually distinguished from other genera in that family by the combination of the following characters: presence of a Trebouxia Puymaly photobiont (most other genera contain Trentepohlia Martius), blackish pigmentation of the pseudoparenchymatous excipulum, presence of lateral paraphyses, and absence of a columella (Guderley et al. 1997; Lumbsch 1989). The genus is widely distributed in arid and semiarid areas and occurs mainly on rocks and soil, although facultatively corticolous species are also known. The related genera having Trentepohlia as the photobiont have their centre of distribution in the wet tropics and are often corticolous. Monographic studies on this genus were initiated in 1982 by one of us (HTL) and the revision of taxa from several regions and species groups were prepared together with several colleagues (Abu-Zinada et al. 1986; Guderley & Lumbsch 1996; Lumbsch 1987, 1988, 1989, 1993; Lumbsch & Aptroot 1993; Lumbsch & Elix 1985, 1989; Lumbsch & Mayrhofer 1990; Mies & Lumbsch 1990). In addition, a revision of the species of Diploschistes in India was published (Pant & Upreti 1993), and a revision of the Australasian species is in preparation by Lumbsch and Elix. The results of these taxonomic studies are summarized in a multi-entry key to Diploschistes species (Lumbsch & Guderley 1996), available via Internet (http://www.botanik.biologie.uni-muenchen.de/ botsamml/lias/modules.html). For an exhaustive treatment of the systematic position, characters, and history of taxonomic exploration, the reader can refer to other papers (Lumbsch 1989; Lumbsch et al. 1997). Cladistic methods have been used in systematic studies of different groups of the lichenized ascomycetes (e.g., Hyvionen et al. 1995; Kmrnefelt et al. 1992; Lutzoni & Brodo 1995; Tehler 1983, 1990, 1993a, 1993b, 1994b, 1995, 1997; Tehler & Egea 1997; Wedin 1993) and an overview of the application of cladistics in ascomycete systematics has been prepared by Tehler (1994a). However, no phylogenetic analysis in the Thelotremataceae has been published. Over a decade ago, at an IAL meeting in Miinster, some preliminary ideas about evolutionary trends in the genus Diploschistes were presented (Lumbsch 1986). These evolutionary trends were deduced from a phylogenetic tree prepared manually. Since then, computer programs developed for cladistic analysis have completely altered the basis for reconstructing phylogenies. To gain a better knowledge of the characters in the genus and with modern computer programs now available for the search of most parsimonious trees, a cladistic analysis of the genus has now been carried out. The object of this contribution is to discuss the relationships in the genus Diploschistes on b sis of the results obtained from the present cladistic analysis as compared to the results formulated by Lumbsch (1986). MATERIALS AND METHODS The study is based on herbarium material of Diploschistes and related genera in the Thelotremataceae from numerous herbaria, including AK, ASU, , BCC, BM, BRI, CANB, CHR, COLO, DUKE, E, ESS, FH, FI, G, GZU, H, HBG, HO, IMI, KOELN, L, LD, M, MB, MEL, O, OTA, PC, S, STU, TUR, U, UPS, US, W, WELT, WRSL, WU, and the private herbaria of A. Aptroot (Baarn), K. Kalb (Neumarkt/Opf.), H. T Lumbsch (Essen), and A. Vezda (Brno). One of us (HTL) has also collected and studied material of most Diploschistes species in the field. For details on methods, refer to earlier publications (e.g., Lumbsch 1989). Cladistic analysis.-All morphological, anatomical, and chemical character data used for the cladistic analysis were assembled from direct examination of specimens; no unconfirmed literature data were used. Unfortunately, no materi l of D. awasthii Pant & Upreti or D. nepalensis 0007-2745/98/398-403$0.75/0 This content downloaded from 157.55.39.91 on Wed, 21 Sep 2016 04:58:16 UTC All use subject to http://about.jstor.org/terms 1998] LUMBSCH & TEHLER: CLADISTICS OF DIPLOSCHISTES 399 TABLE 1. Characters used in the analysis and character states. Habitat and nutrition

Abstract: One new species, Lejeunea convexiloba, is described and figured. Lejeunea proliferans Herzog and L. patens Lindb. var. uncrenata K. C. Chang are treated as new synonyms of L. cocoes Mitt. and L. parva (Hatt.) Mizut., respectively. The former is figured in detail. Three species, Lejeunea konosensis Mizut., L. magohukui Mizut., and L. tuberculosa Steph., are reported from China for the first time and critical comments are provided. Lejeunea is certainly one of the most difficult genera of Hepaticae. Piippo (1990) listed 30 species and one variety from China. Lin and Yang (1992) described a new species, Lejeunea chaishanensis, from Taiwan. Qiu (1993) reported Microlejeunea roundistipula Steph. var. pallida Hatt. (now as Lejeunea pallide-virens Hatt.) from Guangxi. Recently, So and Zhu (1996) added three species, L. otiana Hatt., L. curviloba Steph., and L. sp. from Hong Kong. A total of 36 species and one variety have been recorded from China. This paper describes one new species, treats two synonyms, and reports three species new to China. LEJEUNEA COCOES Mitt., Jour. Proc. Linn. Soc. London 5: 114. 1861. (FIG. 1:1-18) Lejeunea proliferans Herzog, Jour. Hattori Bot. Lab. 14: 51. 1955. syn. nov. The distinguishing features of Lejeunea cocoes include 1) small size of plants (less than 0.6 mm wide with leaves), 2) only 3(-5) medullary cells of stem in transverse section, 3) remote to contiguous leaves distinctly longer than wide, 4) large leaf cells (marginal cells 10-18 x 13-24 jim, median cells 22-38 x 18-28 jim) with thin walls and indistinct trigones, 5) deeply bilobed underleaves with two lanceolate lobes, 6) dioicous sexual condition, 7) presence of regenerants from leaf margins, 8) presence of male bracteoles only at the base of androecium, 9) small compound oil bodies (2-6 per cell, mostly oblong-fusiform, 1.5-2.1 x 4.2-6.3 im), 10) slight tendency towards caducity, 11) well developed leaf lobules with a non-constricted apex, and 12) gynoecium always with a lejeuneoid innovation usually on one side. Lejeunea cocoes is somewhat related to Lejeunea subg. Heterolejeunea (Schust.) Grolle (Grolle 1995), because of the presence of regenerants and its slight ability to produce caducous leaves. Besides Lejeunea cocoes, L. subacuta Mitt. is also a species with strong ability to produce caducous leaves. However, the other characteristics of the two species, such as texture, the first tooth, oil body, leaf lobule, and marginal cells of leaf lobe are typically lejeuneoid. The distinction between Rectolejeunea Evans and Lejeunea has been clarified by Grolle (1995). According to him true Rectolejeunea s. str. is confined to the Neotropics. Lejeunea cocoes is often confused with Lejeunea anisophylla Mont. and Lejeunea subigiensis Steph. These three species have large leaf cells and similar morphology of underleaves. However, L. cocoes can be separated from these two species by being dioicous, having 3(-5) medullary cells of stem in transverse section, and the presence of regenerants from the leaf margin. Sometimes some poorly developed forms of L. cocoes are somewhat similar to Microlejeunea Steph. However, the former differs in the following features: 1) large leaf cells, 2) small, uncurved lobular tooth, 3) absence of basal ocelli in leaf lobe, and 4) presence of regenerants from leaf margin. Particularly, the presence of regenerants can be used to separate L. cocoes from other related species. Regenerants are common in Taiwanese material, but only a few occur in the type collection and in the Guangxi material, and fewer still in other specimens. The known range of L. cocoes now includes China and Sri Lanka (Fig. 3). Descriptions and illustrations are in Mizutani (1963) and Zhu and So (1996). The species has been found on tree trunks, rotten logs, and rarely on living leaves. Specimens examined.-CHINA. GUANGXI. Xingan, Xiaomiaoershan, Gao & Zhang 1754 (HSNU, IFP); HAINAN. Bawangling Nature Reserve, 1,100 m, Zhu 89304 (HSNU); HONG KONG. Tai Mo Shan, 700 m, So & Zhu 95424L2 reported by Zhu and So (1996) as Lejeunea sp. (HKBU, HSNU); TAIWAN. Botel Tobago, Schwabe 90 (holotype of Lejeunea proliferans JE); ZHEJIANG. Baishanzu Nature Reserve, Chegen, 800 m, Zhu 901135 (HSNU). SRI LANKA. Balagama, "Ad truncos Cocos nucifera", Gardner 1399 [1499] (isotype of Lejeunea cocoes BM). LEJEUNEA CONVEXILOBA M.-L. So & R.-L. Zhu, sp. nov. (FIG. 2: 1-30) Planta autoica, caulis irregulariter ramosus. Lobus folii triangulo-ovatus, concavus, apice obtuso-roundato. Lobu0007-2745/98/137-143$0.85/0 This content downloaded from 157.55.39.127 on Mon, 27 Jun 2016 05:26:09 UTC All use subject to http://about.jstor.org/terms 138 THE BRYOLOGIST [VOL. 101