Ph.D. thesis presented to the University of Lordon
January, 1966. 3.
A. revision of the lichen genus Liman* in Europe and its taxonomic, affinities
J. qr. Sheard, V.So., D.I.C.,
Department of Botany, Imperial College, London 501.7. 2 t
Abstract
A partial revision of the genus Rinodina in Europe has been undertaken. 96 species have been studied and 41 of these have been reduced to synonomy. The remaining 50 species include speolespalasdtimp) new to science and 2 new combinations. This revision has been carried out with the aid of more refined sectioning and optical equipment than was available to previous workers and with a greater knowledge of the effects of environ- mental modification. The author has also attempted, to apply modern taxonomic principles during the studies and recent dis- coveries regarding the onotgeny and range of form of the apotheeha have also helped in defining the limits of related genera. The subdivision of the genus has been redefined using the spore structure, rather than the spore septation and tballus morphology, as the principle criterion. The relational*p of the genus Rinodim with the other genera of the Bue=awe, end rivaciacea, has also been examined. It has tentatively been suggested that two genera be redesaribed; the genus k4ismajaQi5 to accommodate certain species previously, placed in BuoUi or Rim Klima, end the monospeoifio genus Dita,010141 previously placed under %Lelia, to be transferred to the Plysataoeao. 3.
introduotion 5
The problem 5 Historical review 6 Review of lichen taxonomy with particular
reference to the genus Ameba 6 Classification' within the Suellitaceae and
aainaidatfa 9
Basis of revision and limits of work 15
Materials and methods 16
Herbaria consulted 16 Sectioning and preparation of material for
detailed examination ' 17
Reagents and mounting fluids used 18 Apparatus used. 19
Revision of the genus angana 19
Presentation of the species 19 Artificial key to the British species of the
genus Nadi1151 21
The genus Natant 21
Sect. Oroular.4. Species i 26
Sect. Evrinodina. Species 2 - 33 29 Seat. itischoblasti4. Species 34. 108
Scot. PlacesthPllto. Species 48 51 145 4.
Discussion 156 Modern concepts of lichen taxonomy 156 The dual nature of lichens and their nomenolature 156 Classification of lichens 158 Taxonomic status of 'chemical species' 160 The species concept 162 Subapecific taxa 167 The ontogeny midrange of apothecial structure within the licheoised fungi 169 Nomenclature of apotheoial tissues 169 The ontogeny of the lichen apotheoium 171 Range of apotheolal structure 172 Morphology and anatomy of the species 178 Thallus morphology 178 Apotheoia - morphology 187 Apotheoia - anatomy 192 Taxonomy of the genus Rinodixia 227 The species and taxa of lower rank 227 Subdivision of the genus 235 Relationships with other genera 242 Phylogeny within the Buelliaosas andayiamit 248 Summary 253 Acknowledgements 255 Bibliography 256 5.
Introduiotion
The Problesi Until the present decade liohenologists have largely limited their studies to the publication of floristio works and have almost totally neglected the monographic treatment of genera. Davis and Heywood (1963) have pointed out that, in respect of the Angiosperms, this is most unfortunate for two reasons: firstly because floras could be written much better if more monographs were available and secondly because systematics cannot advance without monographio research. The same undoubtedly holds true for the lichens and any other group of plants. Following the treatment of the genus Buellia in the British Isles (Mesa, 1960 the author became interested in the delimitation of the two genera Buellia and Rinodina. In the absence of any monographic treatment of all the species of Apladina it was thought desirable that such a work should be commenced in the hope that it would do something to elucidate the problem. This thesis gives the results of the first two years work on the monograph and establishes the blueprint for the remainder of the studies. 6.
Historical Review
Review of lichen taxonomy with particular reference to the genus Rinodina.
The first species belonging to the genus Masa were described after the introduction of the binary system of nomenclature (Linnaeus, 1753) by Aoharius (1798). Two species were described, Lichen sonhodes andis gas. Aoharius was the first worker to devote his attention entirely to lichens and shortly after this time he introduced a system for the classification of the group (Acharius, 1803). In this arrangement greatest importance was given to the form of the thallus but the structure of the apothecium was also taken into account. The species of the genus were placed under,Parmelia Sect. Lecaneria in which the tballus was orustose and the apotheoia partly formed from the thallus. In a later publication (Acharius, 1810) this section was renamed Lecanora subgenus Rinodina and two more species, Lecanora colobina and L. milivina were added to the list, although numerous other subspecies'and varieties were also described that are now regarded as separate species. Gray (1821) was the first worker to use Rinodina as a generic name but it was still used in the same sense as Acharius' Sect. Lecanaria, to include the modern genera Lecanora and Caloolaca amongst others. 7.
The great works of Plerke (1815-19), Pries(1831) and Sohaerer (1850) all followed the classification of Acharius although their species conception varied considerably. The use, by De Notaris (1846), of spore characters in classification constituted a great advance in liehenology. Such a system had previously been suggested by Fee (1824) but never utilised.
The value of this system of classification was quickly realised and the works of Massalongo (1852), Koerber (1855) and Mudd (1861) all employed spore characters in defining genera, while morphological characters were retained for the classification of Families. Thus, in Veda (loc. cit.) the genus,Rinodin, is found with Calloeismq (palotlaca), /meanie, Lecanor and Itaggdosa in the Tribe (Family) jeoanoraceeel the genera being separated primari4 by spore characters. Not all authors were of the same opinion however, and !Wander (1854) rejected the use of ascospore Characters except where they were of use in species diagnosis. The occurrence of green cells in lichen 0414 had been known for a considerable time and had, been intensively studied by iallroth (1825) and. Koerber (1839), but it remained for Schwendener (1867) to announce the composite nature of the lichen plant. The association of fungi and algae within lichens came to be regarded as a high/7 developed form of symbiosis and the taxonomic views of most liohenologists of the El • time maybe summed up in the words of Smith (19211):- "AA, system of classification that claims to be a natural one, must, while following as far as possible the line of plant development, take into account the double origin of lichens both from algae and fungi, the essential unity of the class however being proved by the recurring aLalarity between the thalline types of the different *viler. This was the basis for the classification of lichens constructed by Reinke (189496) and elaborated by Zahlbruckner (1926). Although it is now regarded as having severe limitations it is still used in the majority of taxonomio works. In this system the Ascolichena are divided into Series and Subseries based on apothecial structure and morphology. The Families are variously defined, by thallus morphology, algal cells and apotheium structure. The genus Rinodina is thus found in:- Subclass Asooliohens Series Gynmooarpeae Subseries Cyolocarpineae Family Buelliaceae The Family PhYsoiaceae is distinguished from the Buspiacete by its foliose thallus. The genera Buellia and Rinodina constitute the Buelliaceae and are distinguished by their lecideine and lecanorine apothecia respectively. The Physolaceae inoludes the genus PVxine separated from,Phvsoia and Anaotychla in a similar manner by the absence of a thalline margin. The latter 9.
two genera differ in the structure of the thallus. A later olassification byliratoon(1928) oombined these two Families as the Phrscipiocae and included the genus Rhizocayson, placed by Zahlbruokner in the Livitmeeae. In many ways this arrangement is a retrograde step and has been shown to be erroneous, at least in part, by Lamb (1940) ana,Sheard (1961.). Unfortunately it is the basis of the current classification of British lichens (Watson, 1953),
Classification within the Buelliacese and Physciaoeae.
The classification of the Buelliaceae by Zahlbruckner (1926) may be summarised as follows:- Buellia De Notrs. Sect. I Rubue11414 Sect. II Diralotomma Th. Fr. Sect. III D of Stzbgr. Rinodina S. Gray Sect. I Oroularia Malme Soot. I/ Etrinodinft Stzbgr. Sub3oct. Paanvenoria Malmo Conradia Malmo II/ Nischoblastia Mame IV ilacothallia Buelliq Sect. DiuloiciA and MOIL& Sect. Placothallia are distinguished from the other sections of their genera by their squamulose thallus forms. 4310.14 Sect. Diulotomma is distinguished from Sect. Rubuellia by its pseudomuriform or 3-septate spores and spurious thallino margin. Speoles with 3-septate spores are however, found 10. in Sect. Eubuellia but those lack a spurious margin to the apothecia. There are indications (Sheard, 196?.) that soot. AuMaki should be divided into subsections based on spore characters and insertion of the apothecia on the thallue. Thus the Btteilia ditectiformis group of species have pointedly ellipsoid spores differing in structure from the bluntly ellipsoid spores of the At jgadat§ and groups of species. The B. aethelea, group differs from the other groups in having innate apotheota and areolate thalli. The position of those species in the genus with polarilooular spores (Imshaugh: 1955, and Malmo, 1927) has not yet been considered from the tatenamio point of view. Classification within the genus Molina is equally complex and owes much to the -.7ork of Name (1902). The genus is again sub- divided by the use of spore characters. sect..W.4=Q contains those species in which thickening of the spore Ilan is limited to the septum (placodionorph), 38ot. Lug has typical polarilooular spores caid sect. tolischoblastia those species with mischoblastionerph spores. Subsects. Pachmorig and Conradia of :=sot. 3vrinodinq are distinguished by their 1-septate and 3-septate spores respectively. Some or the squamulose speolos of sect. Aggsaglisla nay also be distinguished by their unthiokoned spore walls. These spore typos are illustrated. in Figure 1. In making this classification Zehlbrucknor chose to ignore the work of Wainie (1909) by placing the genus Lie,lansa4a1.344 in Liman& This genus was created to accommodate those apecies 11.
b
d
Figure 1. Spore types of the genus Rinodina. a) Spores with unthiokened walls, R. nulleri; b) plaoodionorph spores, R. insuerato; c) polarilooular spores of R. turfaoes; and d) mischoblastionorph spores of R. consnersa. Magnification x (Prom Milne, 1902). 12. closely related to both Bualia and Rinodina and which apparently fell between the two genera. It therefore contained those species with innate apotheoia and much reduced and pale excipuli. The genus was further divided into three sections by the oolour of the hypotheoium. Thus Sect. SemibueMia contained those species with light brown bypothecia or with hypotheoia which were pale above and brown below or vioa versa. Seat. Buellioosis had a dark brown hypotheoium and Sept. Rindodincosis a colourless I]ypotheoium. The only other major taxonomic work within the Buelliaceas is that of Magnusson (1947) on the non-saxioolousBinodinet species of Europe. The taxonomic arrangement used is that of Malme except that Subseot. ,Pachvsoaria is split into three series based on the thickening of the spore" wall. Series SouhcdAnge is characterised by possessing unthickened spore walls at the apices and septum; Series Archaeinae by having the spore walls thickened at the apices and septum but thin laterally; series Roborinae by the spore walls also being thickened laterally and by the central lamellae being indistinct. These three spore types are illustrated in Figure 2. The classification of the Phxlciospeae adopted by Zahlbruckner is as follows:-
=LS Fr •
Pharssia, Ash. Sect. I Dirinaria I/ Euchveoia Th. Pr. III nvoaroftvacia Zahlbr. Anactychiel K8rb. a
Figure 2. Spore types of Rinodina Subsect. Pachysooria. a) R. sophodes, spore wall uniformly thin b) R. archaea, spore wall thickened at apices and centre, laterally thin; c) ,R. roboris, spore wall uniformly thick. Magnification x 1,700. (After Magnusson, 1947). 14.
Recent studies on the genusWA22 by Imshaug (1957) has resulted in the genus being divided into two sections. Seat. agat and Seat. Pbramemine, by the use of spore characters. The former section has 1•septate, misehoblastiomorph spores; the letter 5..septate spores. The development of a red stipe was considered to be of sUbsectional importance. Of fundamentally greater importance was the discovery that the genus does not have a true lecideine apothecium and that some species contain algal cells in the margin so that they approach the genus Phvacia very closely. Physcia Sect. pirinarie differs from Sect. Euehvsoia in the dark hypotheoium and in this respect it is similar to,Praine. Lynge (1924) believed it could be placed in Pvxine or Phvsola with equal justification but lmshaug suggests that it ought to be regarded as a genus in its own right. It not only differs from Pvxine in the structure of the apothecial margin but also in that it lacks an aerugineous epithecium with a distinctive KOH+ violet reaction. ayggia Sect. Hyrerehyscia is separated from the other two sections of the genus by the muriform nature of spores at maturity. The most recent concepts of classification within the Bielliaceas and Phveciaosae therefore primarily employ characters of apothecial morphology and anatomy to define the genera. Within genera, characters of spore structure and septation are used in delimiting Sections and Subsections. The morphology of the thallus 15.
is also used at this level of classification although it is more often retained to separate Families. Basis of revision and limits of the work It is quite clear from the historical review that the work of Zahlbruckner (1926) should form the basis of the taxonomic revision of the genus and that the species listed by Zahibruokner (19224+0) the basis for the species to be studied. This work has recently been brought up to date by Lamb (1963) and the new species of Rinodina listed in this work were added to Zahlbruokner's list. In addition, the previous work of the author (Sheard, 1964) had indicated that a number of species of Buellia would be more correctly placed in Rinedile and these species were also added to the list to be studied. The number of European species listed in this way came to over 150 and it was therefore decided to limit the study in the first instance in the belief that the remainder of the world's species would be relatively easy to revise once order had been brought to the European species. To attempt to study all the world species at one time would only serve to confuse the issue further. Examination of the herbarium sheets of the genus at Kew and the British Museum, however, immediately posed another problem. The specimens mounted on each sheet labelled under one species were, more often than not, found to represent a number of species with no indication as to which might be the correct determination. It was therefore realised from the outset that the only possible method of attempting the revision 16.
was to locate and study the type material of each species or failing this, authentic material. This in turn imposed a further limitation on the work as it was found that the type material of many of the species was not readily available and the search for these is still continuing.
Materials_and Methods Herbaria consulted Type and authentic material has been obtained from the following eight herbaria:- British Museum (Natural History), London (BM) Royal Botanic Gardens, Kew (K) Institute of Botany, Lisboa (LIEU) Botanische Staatssammlung, Munchen (M) Universidade do Porto (M) ftturhistoriska Rikemuseum, Stockholm (S) Botanical Institute of the University, Turku (TUR) Institute of Systematic Botany, University of Uppsala (UPS) The letters in brackets refer to the herbarium abbreviations listed in Index Herbariorum(Regnum Vegetabile, j, 1965). Additional material for the British Isles has also been studied from the following national herbaria:- National Museum of Ireland, Dublin (DBN) Royal Botanic Garden, Edinburgh (E) National Museum of ".Sales, Cardiff (NM7) 17.
and the private herbaria of:- Dr. K. L. Alvin, London Mies U. K. Duncan, Arbroath Dr. K. A. Kershaw, London J. W. Sheard, London Dr. T. D. V. Swinscow, London
The study of the material from these herbaria has taken place in three parts 1) the sectioning, preparation of semi-permanent mounts and the critical examination of the type or authentic material, 2) the identification and critical examination of further material of each species to determine the range of variation within the species and 3) a detailed and complete revision of the genus in the British Isles involving the study of relevant material in the national and private herbaria listed above.
Sectioning and preparation of material for detailed examination. Experience has shown the following procedure to be the most convenient when making detailed examinations:- 1) The thallus of the specimen is softened with a 50% solution of ethyl alcohol in water. This solution is superior to water alone because the alcohol acts as a wetting agent and speeds the
PrO0888 •
2) When all surface moisture has disappeared two apothecia are dissected away from the thallus and placed in 50 alcohol for further softening. 3) The cortex and medulla of the thallus are tested for chemical 18. reactions with potassium hydroxide (K), hypoohlorite solution (C), aqueous iodine (I) and paraphenylendiamine (P) using micro-droppers and any colour reactions noted. 4.) One apotheoium is mounted and correctly orientated in a wax block by melting the wax with a hot needle. 'Glitterwax' is said to be the most convenient (Swinscow, 1962) but as this is no longer obtainable, paraffin wax (L P. 52°C) was used in its place. 5) The thallus is again examined for any further change in the colour reaction.
6) After the wax has solidified the apothecium is sectioned at 30 - 50p and one section is mounted in each of the following reagents:- a) Gum chloral b) Cotton blue a) K, d) I, e) nitric acid (N). 7) Immediate examination of sections o), a) and e) for colour reactions. 8) Labelling and sealing of sections a) and b) to make semi- permanent slides for subsequent examination.
Reagents and mounting fluids used K 5% solution of aqueous potassium hydroxide C Concentrated hyl?ochlorite solution - Parasone
1% aqueous iodine in 1% potassium iodide
P 1% solution of p-phenylendiamine in ethyl alcohol or stable solution (Scott, 1959). 19. N 1070 solution of nitric acid.
Cotton Blue 0.067, solution of cotton blue in lactophenol.
Gum chloral As supplied by G. T. Gurr Ltd.
Nail varnish Clear nail varnish used for sealing semi- permanent slides.
Apparatus used Morphological examinations and dissections were carried out under Charles Perry "Prefect" binoculars giving a magnification of 12-20x. Apothecial sections wore obtained with a Yung sledge,- microtomo using a flat-ground microtome knife. A Gillett and Sibert "Lablynx II" binocular microscope was used for studying all sections. Spore and hyphal measurements wore carried out under a magnification of 1,000x and all other measurements at 400x.
Revision of the genus Rinodina
Presentation of the species The species have been numbered and described in the same sequence as the taxonomic arrangement of species on page 238. An artificial key has also been constructed but is limited to the
British species of the genus as time has not allowed the complete revision of all the European species. The following sections appear under each species:- Synonomy
The list of synonomy for the species is not necessarily 20.
complete as little benefit would have been derived by copying the lists in Zahlbruckner (1932-40). The lists are therefore limited
to changes of nomenclature that have been effected in the present
work and also include any other synonyms that have caused confusion
in the past. Abbreviation of journal names has been carried out in accordance with the World List of Scientific Periodicals and
the citation of referencesfollows the International Code of
Botanical Nomenclature (1961), The abbreviation of authors' names follows the work of Grumman (1963).
Exsiccati
A list of exsiccati has been given for those species where it has been possible to locate and study them. The abbreviation of exsiccati names has been carried out in accordance with Lynge (1915) and abbreviation of the herbaria in which they occur follows
Index Berbariorum (Regnum Vegetabile, 31$ 1964). The name of the species by which they are referred to in the exsiccati is given and where possible the fullest details of locality, collector and
date of collection have been added.
Species descriptions
The species descriptions are largely self explanatory but
mention should be made of the possible error in the measurements.
The accuracy of the dimensions of the areolae and apotbecia is limited by the smallest units of the measuring scales used which were, under a magnification of 30x, 0,036mm. in sise. These
measurements era therefore accurate to the nearest 0.03mm. but have 21. not been rounded off to this figure at the present stage of the work. Measurements of tissues carried out under the microscope at a magnification of 400x are theoretically correct to the nearest 1.5p but the error of observation is far greater than this because it is rarely possible to delimit the tissues accurately. These figures have therefore been reduced to the nearest 5p. The hyphal and spore measurements are accurate to the nearest 0.05p but the figures have again not been reduced at this stage of the work.
Variability and identification, and taxonomic remarks
These two sections are again self explanatory and in a few cases are inevitably somewhat repetitive.
Distribution of species
No comments have been made on the ecology of the species because it has not been possible to carry out field studies on them. However, it was possible during the course of the work to - map the distribution of the species occurring within the British
Isles on a 10km. square basis and the maps are included,under the relevant species.
Artificial key to the British species of the genus
Rinodina.
1) Terricolous or muscicolous 2)
Lignicolous or corticolous 4)
Saxicolous 9) 22. 2) Spores with predominantly 2 lumina, thallus IC+ yellow. 3) Spores with predominantly 4 lumina, thallus R. conradi p.128 3) Thallus with coralloid isidia, spores (20.9-31.1) 26.7 x 14.2(11.4.47.1)µ. Plant of mossy trees. R. isidioidem p.132 Thallus granular but never with isidia, spores (18.5-22.8)18.7 x 10.7(8.8..14.2)p. On decaying Armeria maritime, of maritime cliffs. RA maritime p.135 4) Apothecium margin concolourous with thallus, hypothecium colourless or pale brown, spores polarilocular or mischoblastiomorph. 5) Apothecium margin concolourous with disc, hypothecium dark brown and spores placodlomorph. R. biloculata p.26 5) Thallus light to dark grey, epithecium brown or red-
brown, K-, N-. Thallus blackish of minute granules, epithecium blue- black, K+ and N+ violet. R. colobina p•62
0) Spores mischoblastiomorph or lumina rounded with thick walls. 7) Spores always polarilocular at first, lumina sometimes
becoming rounded but then with thin walls. 8)
23.
7) Thallus apothecia sub-innate, spores (20.5..31.9) 26.3 x 14.8(11.4,49.8)p, dilated meridiorally. R. teichorhila p.117
Thallus K+ yellow, arothecia sessile, spores (14.3-21.7)18.6 x 10.0(7.7-.12.1)p, not dilated meridioralIy. R. roboris p.142
8) Speros with thin, equally thickened walls, (13.2-X9.8)15.7 x 7.9(6.6-9.1)p. Plant of young twigs of smooth barked trees especially Ash. R. sorhodes n.I05
Spores with thick, unequally trickered wells, (11.4,22.8)160 x 7.5(5.7-9.7)µ. Plant of rough bark or decorticate wood. R. axiom p.66
9) Apcthecium marvih corcolourous with diso. 10) Apothecium margin corcolourous with thnllus. 11) 10)Spores mischcblastiomorph, (16.5-25.3)20.7 x 11.5 (8.5-16.5)p. R. oxydata p.112 ;:peros polarilocular, (12.1-18.2)14.3 x 7.4(5.7-8.5)p. R. occults p.80 11) Thallus C+ red. 12) Thallus C-. 13)
12) Thallus light grey, F+ yellow, apothecia seevile, spores with rounded lumina and thick walls. R. atrocirerea p.120 `Phallus dark grey, X-, apothecia sub-innate, spores polarilocular with unequally thickened walls. R. luridescers n.76 24.
13) Spores misohoblastiomorph or with rounded lumina and thick walls. 14.) Spores polarilooular, with thin walls if lumina' rounded. 15) 14)Phallus apothecia sub-innate, spores (20.5-31.9) 26.3 x 14..8(11.4.-19.8)p, dilated meridionally. R. teichotthile. p.117 Thallus yellow, apothecia sessile, spores (16.5-22.8)18.7 x 10.7{8.5-14.2)p, not dilated meridienally. R. maritima p.135 15)Thallus K-, spores usually broadly ellipsoid but if narrowly ellipsoid then o. 15µ long. 16) Phallus yellow, spores narrowly ellipsoid, (16.0-22.0)19.7 x 9.0(7.7-10.3)µ. R. confragosa P-34 16)Phallus rarely developed, spores (14.3-19.9)16.8 x 11.3(9.9-12.5)p. Plant of calcareous rocks and apothecia sometimes in pits. R. bischoffii 1).38 Phallus usually developed, spores (11.0-19.8)15.1 x 8.5(6.8-11.0)p. Plant of siliceous or calcareous rooks but apothecia rarely in pits.'R. salina .P•82
Rinodina S. Gray in Natur. Arrang. Brit. Plants, 1: 448 (1821) pro parte. Parmelia Sect. Lecanaria Ath., Method. Lich., 154- (1803) pro parte; Leoanora Subgen. Rinodina Ach., Liohenogr. 25.
Univers., 344 (1810) pro parte; ,Berengeria, Travis., Rivista Period. Lavori Acoad. Padova, 265 (1851-52); Melanaspicilig Vainio, Ark. Bot., 8 (4): 76 (1909) pro parte.
Description Thallus thick, thin or evanescent, light to dark grey, ochraceous or brown, continuous, rinse, areolate or subsquamulose, plane, granular or verruoiform, matt or rarely waxy, rarely pruinose, sorediate or isidiate. Thallus determinate or not, with or without a limiting, entire or fimbriate, dark hypothallus. Apothecia innate or sessile, usually frequent, contiguous or not. Disc brown or black, plane or convex at maturity rarely pruinose; margin ooncolourous with disc or thallus, entire or orenulate, persistent or excluded at maturity. Exoipulum thallinum with or without a distinct cortex; cortex cellular, occasionally pigmented, sometimes columnar below or structure occluded. Peripheral hyphae usually pigmented. Exipulum proprium hyaline or rarely brownish, prosoplectenchymatous. Hypotheoium colourless, rarely brownish or dark brown, stipe not usually present, colourless or dark. Theoium colourless. Paraphyses expanded at apices, fusoo-oapitate, forming a brown or red-brown, rarely a dark brown or blue-black epitheoium. A1101 elevate, inner walls uniform or not. Spores 2- or 4-celled, septum well developed at maturity or not. Pycnidia rarely present, conidia baelliform. 26. I Rinodina Seat. Oroularia Malme in L. Svenska VetenskAcad. Handl., 28: 44 (1902).
Description Apothsoial cortex pigmented, hypotheoium dark brown and spores placodiomorph.
1) RinodinsI biloculatanov. comb. Leoidea bilooulata Nyl., Flora, Regensburg, ha: 460 (1877). Leoidea rolospora Leight., Trans. Linn. Soo. Lond. Hot., 1: 241 (1878). - Lecanora biloculats;Hyl. ex Hue, Revue de Botan., 29 0886-87). - Buellia biloculata Oliv., Expos. Lid:. Ouest France, 2: 150 (1901).
Authentic material Isotype:- Nyl. 10437 (H) Leoidea biloculata, IVlemore, Larbalestier.
Description Thallus thin, light grey to oohraoeous, continuous or rarely cracked and plane. Thallus determinate or indeterminate, without perceptible hypothallus. Apothecia sessile, never contiguous, to c. 0.29mm. diem. Disc black, persistently plane; margin concolourous with disc, o. 0.04mm. wide, entire and persistent. Exoipulum thallinum 204.5p wide; cortex pigmented, 10-15p 27., wide, medulla 5-35p wide. Peripheral hyphae o. 446p wide, medullary hyphae 1.7-2.3p and algal oells to c. 9p diem. Excipulum proprium, o. 10p wide laterally, expanding to 10-15p above, hyphae c. 1.1p wide. Hypothecium dark brown, 45-60p deep. Theoium 40-70p high; paraphyses a. 1.1p1 wide, apices to 2.3.4.6p wide, forming a dark brown epitheoium. Aeci a. 45 x 19p, inner walls not uniform. Spores plaoodiomorph, (13.7-17.1)15.6 x 7.6(6.8-8.5)p, apical walls o. 1.2pt thick, lateral walls c. 1.7p, porus o. 1.1p wide and septum not developed. Thalline reactions:- K=, c:, I=, P=. Hypothecium and theoium I+ blue. Plant corticolous.
Variability and identification Nothing is known about the range of variation shown by this species as only two specimens have been examined. The species has a superficial resemblance to a diminutive specimen of Buellia disoiformis and this species has often been mistaken for Rinodina bfiloculata in the past. It is of course readily distinguishable by the placodiomorph spores.
Taxonomic remarks There can be no doubt that Buellia polosoora belongs to this species although no healthy spores were found in the sections examined. Both species have, in any case, been previously described under the name Lecanora bilooulata by Nylander (1878). Despite the fact that Nylander realised the species possessed a lecanorine margin 1.11
• Ri nod ina 9 1 • occulta (Koerb.)
nov. comb. • .
® Ri nod ina 114,.. • 8 biloculata (Ny I. ) •
nov. comb. q 5,' .. - , . 4. 001: ...... -_,-, MILES: 1 , A
tr Is.. -
fr
I
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2 4
BOTANICAL SOCIETY OF THE BRITISH ISLES DISTRIBUTION MAPS SCHEME
newt Ketalbuttott of jraitle 111 Ow British tale*
29. to the apothecium, the species has since appeared in British lists in the genus Buellia. The species is the only British and possibly the only European member of the Sect. Oroularia. The dark coloured hypothecium and the dark apothecium margin lead Malmo (1902) to state that the relationship of this section with BuelliA and Rinodint was ambiguous. It appears to bear the same relation with these genera as the genus Dirinaria does with WL...ne and ,Physcia.
II Rinodina Sect. Eurinodinac Stis. in Bericht. fiber die Thatigk. St. Gallisoh. natures. Gesellsch., 169 (1862). Beret:Aerie subgen. Rinodina Travis., Revista Period. Lavori Acoad. Padova, 265 (1851-52): Rinodina Sect. Paohmoria Malme, K. Svenska VetenskAcad. Handl., 18 (1902)pro parte.
Description Apotheoial cortex colourless or rarely pigmented, hypotheoium colourless or rarely brownish, spores polarilocular, lumina sometimes rounded and spore wall uniformly thick or thin septum usually well developed.
2) Rinodina arohaea Arn. in Flora, Regensburg, 195 (1881). Parmelia sonhodes var. arohaea Aoh., Method. Lich.: 30. 156 (1803): -, 311110disla =means Vain. ex Ahiqu., Ann. Aced Sci.
Penn., Ber. A, a (6b): 83 (1928). -pinedina liena:106 Nign, (pro parte), Svensk. Bot. Tidskr., 30: 259 (1936). -amok sibiric,H. Nagai, Bvensk. Bot. Tidskr., 30: 261 (1936). ...2dMitbet arab:el:dam H. Magna, Acta Herta Gothoburg., 17: 278 (1947) - Rinodina etcchediedes H. Naga., Acta Hort. Gothebtal., 17: 226 (1947). Rinodina qubfluscuk H. Naga., Acta Hort. Gothoburg., 17: 240 (1947). - Ili:Iodine "13.U:beau:A H. Nam:., Acta Hort. Gothoburg., 17: 279 (1947).
Bssieceti Ansi Lioh. Lang. 460 (X) mniarnka var. ,cinnamomea f. minute; Lich. 'ann. 608, 923 (K) - Rinodinasame; Lojka Univ. 75 (E) Lecanora qouhodes.
Authentic material Doh. B.N. 409 (BM) - Lecanora soul:odes var.,arehaea.
Descriation
Mall= thin or evanescent, sometimes ladhing or rarely thick, light to dark grey, usually brownish, continuous or rims°, plane or granular, often verruciform, verrucae sometimes discrete. 'Phallus determinate, more usually indeterminate, rarely with a thin, fimbriate hypothAllus. Apothecia sessile, sometimes continguous to 0.36-0.72mm. diam. Disc black, rarely dark brown, plane sometimes becoming convex; margin concolourous with thallus, 0.04-0.11mm. wide, entire and persistent. 31. Excipulum tholkinum 40-.95p wide; cortex cellular laterally, to o. 20p wide or absent, sometimes columnar and below expanded/to 15•40p wide, medulla 35-75p wide. Peripheral hyphae rarely not pigmented, 4.6-6.8p wide, medplinry ;mime 2.34r.6µ wide and algal cells to 12-20p diam. Excipulum proprium 10-20p wide laterally, expanded to 15-40p above, hyphae 1.1•1.7p wide. Hypotheoium 20-85p deep. Theoium 75-110p high; paraphyses 1.7-2.3p wide, apices to 3.0-6.8p wide, forming a red-brown epitheoium. Aso/ (40-70) x (15*26)1.14 inner wails uniform or not. Spores polarilocular, thick walled, walls rarely becoming uniformly thickened, (13.1-25.6)22.8 x 9.7(6.8-11.04 apical walls ct. 2.2-3•7p thick, lateral walls c. 1.2-1.7p, porus c. 1.1-2.8p wide when present and septum prominent. Thalline rsaotionss- C:, /=, P=. Hypothecium and thecium I+ blue. Plant corticolous, lignicolous or musoicolous.
Variability This species shows the wide range of variation, both of morphology and internal structure and more particularly in the structure of the apothecial cortex, usually associated with such a common and wide ranging species. It may be recognised by the structure and sine of the spores, the great height of the theoium relative to the hypotheoium and by the red-brown opithecium. The speoies should not be confused with Pineal= stlbana which 32. lacks a prominent spore septum, usually has a brown apothecial
disc and has a well developed, determinate thallus with con- tiguous apothecia rather similar to those of R. sophodes, or with R. trevisiana which has a thin continuous thallus and convex apothecia.
Taxonomic remarks Magnusson recognised this species es having widely attached apothecia and lacking an apothectal cortex. Thus he separated it from species having apothecia constricted at the base and from other species having a cellular or columnar cortex.
The present work has shown however, that in many Rinodina species is the thickness of the cortex may vary considerably and it/often
difficult to know whether or not to recognise a cortex in some of those species reported to possess one. It has further been shown
that the development of a lower, columnar cortex can be correlated, in R. archaea, with the lateral growth of the apothecium and hence
with the constriction of tho base of the apothecium. Some of Magnusson's observations would therefore appear to be incorrect as he separated species on the basis of two characters now shown to be linked.
Magnusson states that Rinodina archaeoides differs from
R. archaea in having a thick, columnar lower cortex to the apothecium which the writer's personal observations have failed to
detect. On the other hand R. sophodiodes has been shown to have 33. a thick, columnar cortex which Magnusson specifically states it does not possess. It must therefore be concluded that this character is extremely variable, a conclusion which must be
substantiated by a section of the type specimen of R. subfusca which was seen to have a thick, columnar cortex on one side of the apothecium and a thin, cellular cortex on the other. On
the basis of this evidence these three species have been placed under R. archnea together with R. granulane, R. sibirica and R. subobscure, all of which show a similar variability of cortical structure. Mention should also be made of two species, Rinodina laevigate and R. septentrionalie, which Magnums= indicates are common in sub-arctic Europe. No authentic specimens of these species have been viewed in the present work but as they are said to differ from R. arehaes only in the possession of a cortex, the structure of the apical spore walls and the constriction of the apothecium base, they cannot be regarded as good species until they have been more critically examined. The positive reaction of the hypothecium with iodine is of particular interest in this species as the reaction may be very faint or absent in specimens with a very deep hypothecium. Most
species of Rinodina have a tissue, deeply staining with iodine, that lies between the hypothecium and thecium proper and which in R. archnea apparently extends much further down into the hypothecium
than Is usual. This raises the question as to the nature of the tissue which cannot be distinguished morphologically. It seems 34. possible that it is analogous with the endogenous tissue referred to by Gelting (1954) although it is difficult to be quite sure as he does not define the tissue sully.
3) Rinodina confragosa Koerb. in Syst. Lich. German.; 123 (1855). Parse-11a confragosa Ache, Suppl. Method. Lich.: 33 (1803).
Exsiccati Anzi Lang. 394 (K) - Rinodina metabolica; Anal Yen. 45 (K) - Rinodina aggregeta; Ara. 68 (IC) Rinodina confragosa; Rabenh. 78 (K) - Rinodina csesiella, 873 (K) - Rinodina confragosa; Zw. 190 (K) - Lecanore caesiella.
Authentic material Ach. B.M. 387 (BM) - Lecanora atra var. confragosa.
Description, Thallus thick or thin, light grey to ochraceous, rimose end plane or of discrete verrucae, verrucae and margin of thallus subsquamulose. Thallus indeterminate without perceptible hypothallus. Apothecia sessile, sometimes contiguous, to 0.58-0.86mm. diem. Disc black, plane becoming slightly convex; margin concolourous with thallus, 0.07-0.11 m. wide, entire and 35. persistent. Excipulum thallinum 85-115p wide; cortex 15-20p wide, structure occluded, medulla 70-100p. Expanded peripheral hyphae not present, medullary hyphae 268-567p wide and algal cells to 12-18p diem. Excipulum proprium 10-15p wide laterally, expanding to 10-.25p above, hyphae 1.1-1.7p wide. Nypothecium 60-135p deep. Thecium 90-100p high; paraphyses 1.1-2.14 wide, apices to 3.5- 5.10, forming a brown epithecium. Asci (60-75) x (10-19)p4 inner walls uniform. Spores polarilocular, walls thick, sometimes developing thin, uniform walls, (16.0-22.0)19.7 x 9.0 (7.7-10.3)p, apical walls c. 3.1m thick, lateral walls c. 1.4-1.(a, porus
2.0-2.2p wide and septum well developed at maturity.
Thalline reactions:- K+ yellow, C-, I-, P-.
Hypothecium and thecium I+ blue. Plant siliceous.
Variability and identification Nothing can be said about the variation of the species within the British Isles as only two specimens have been recorded to date. The thallus morphology of the Scandinavian material examined is remarkably constant although the spare morphology is more variable. In most of this material the spores are distinctly polarilocular with angular lumina but in a few specimens this structure is only apparent in very young spores, the lumina becoming rounded with age.
Rinodina confragosa is easily distinguished from 3E •
BOTANICAL SOCIETY OF THE BRITISH ISLES DISTRIBUTION MAPS SCHEME
Figure 4.. Pistributior. of Ax0::.47x1 porrr9gpjA trW) ,j.j ,161Laa the 7iritish Tales. in 37.
R. maritime, a distinctly western species in the British Isles, by the spore type. The spores of the latter species are broadly ellipsoid, never with a prominent septum, mischoblastiomorPh or with thick, uniformly thickened walls.
Taxonomic remarks The species belongs to the polarilocular group of species and appears to be more closely related to Rinodina archaea than any other species. The thin outer apical spore wall is reminiscent of Buellia disciformis and perhaps indicates some relationship with Rinodina consocians.
4) Rinodina erumpens H. Magn. in Acta Hort. Gothoburg., 17: 231 (1047).
Authentic material
Syntype:- Brenner 400.1 (S) Rinodina erurnpens,
Yenisejsk: Garinskij Volok, Brenner, 1876.
Description Phallus thin or evanescent, light grey, continuous and plane. Thallus indeterminate, without perceptible hypothallus.
Apothecia erumpent, sessile and frequent, rarely contiguous, to
0.30-0.47mm. diem. Disc black, plane becoming convex; margin concolourous with thallus, 044-0.07mm. wide, entire becoming exoludeit. 38. Excipulum thallinum 40-45p wide; cortex absent. Pigmented peripheral hyphae absent or to c. 2.8p wide, medullary hyphae 2.3-2.8p wide and algal cells to 13-15p diem. Excipulum proprium c. 15p wide laterally, expanding to 20.45p above, pale brown, hyphae 1.1.4.7p wide. Rypothecium 25.40p deep. Thecium 86.40u high; paraphyses 1.1-1.7p wide, apices to 2.3-3.4p wide, forming a red-brown epithecium. Asci (65-75) x (22-25)p, inner walls uniform. Spores polarilocular, thick walled, the lumina dilated, (18.2.-24.5)22.3 x 10.1 (8.5...12.0)1, apical walls c. 3.8-4.3p thick, lateral walls c. 1.dp, Pcrus c. 1.4-2.1p wide when present and septum well developed. Thalline reactions:- K-, C-, I-, P:. Hypothecium and thecium I+ blue. Plant corticolous.
Identification and variability The specimens of the species that have been examined show no marked divergencies. The gross morphology of Rinodina erumpens shows many features in common with the Ac archaes group of species. It may, however, be separated from these species by the erumpent nature of the apothecia, a character which has never been observed amongst the latter species. R. erumpens may be further distinguished from these species by the dilated spore lumina.
Taxonomic remarks The anatomy of the apothecia of this species shows a 39 •
very close relationship with Rinodina archaea. The possibility of reducing the species to a variety of R. archaea has been considered and rejected as further material would be required to justify such action. The erumpent epothecia and spore morphology have no counterpart in the R. archaea specimens examined to date, providing evidence that R. erumpons should remain a separate species at the present time.
5) Rinodina excrescene Vain ex Ahlqu. in Ann. Acad. Sci. Fenn., ser. A. 27 (8b): 82 (1928).
Authentic material Holotype:- Vain. 8798 (ruR) Rinodina excrescens, Leunsk, Vainio, 1880.
Description Thallus thick, light to dark grey, ereolate. Areolae to c. 1.98mm. wide, granular becoming papillateisidiate. Thallus indeterminate without perceptible hypothallus. Apothecia innate, frequent but not contiguous, to c. 1.26mm. diem. Disc brown becoming black, persistently plane; margin concolourous with thallus, c. 0.18mm. wide, crenulate and persistent. Excipulum thallinum c. 105p wide; cortex cellular, c. Ism wide, medulla c. 90p. Peripheral hyphae not pigmented, to c. 5.7p wide, medullary hyphac c. 4.0p wide and algal cells to 40.. c. 14p diem. Excipulum proprium c, 15p wide laterally, expanding to e. 40p above, hyphac c. 1.7p wide. flypothecium c. 90p deep. 'Meet= c. 130p high; paraphysea c. 2.3p wide, apices to c. 4.6p wide, forming a red-brown epithecium. Asci 60 x 13p, inner walls uniform. Spores polariloculer, thick walled, (13.7-17.1)15.3 x 6.9(6.3-6.0)p, apical walls co 3.4p thick, lateral walls c. 1.7p, porus co 1.7p wide and septum well developed. Thalline reactions:- IC:, C:, I:, P:. Hypothecium and thecium I+ blue. Plant lignicolouso yarisgadntiiL. cation The typo specimen of Rinodina exerescens indicates that little confusion is likely to arise over the identification of the species. The ioldiate thallus is similar to that of R. lsidioides although not as well developed. The polarilocular spores, how- ever, show that R. oxereseens belongs to quite another group of species.
Taxonomic remarks The spore morphology and thallus habit seem to make this a taxonomically isolated species. The overall internal anatomy of the apotheeium suggests some relationship with Rinodina archaea but this cannot be very close. 41 . 0) Rinodina mntarnea Koerb. in cyst. Lich, German.: 128 (1855).
Lecanora mniaraea Ach., Synops. Lich.: 339 (1814). ••• Lecanora mniaroeisa Kyl„ Flora, Regensburg, 53: 33 (1870).
Rxaiccati Ansi Lang. 108 (K) Rinodina amniocola, 513 (K) Rinodina mniaraea f. biatorina; Ansi It. 219 (K) Rinodina mniaraea, 220 (K) Rinodina mniaraea var. cinnamomea; Arn.433 (K) Rinodina turfacea var. miaraes: Krypt. Mind. 2587 (K) - Rinodina mniarea var. normalis; Lich. Fenn. 100, 808 (X) Rinodina orbata f. depauperate; Rabenh. 380, 382 (K) - Psora turfacea; 7w. 914 (K) Lecanora maiaraeiza.
Description
Thallus thick or thin, light to dark grey, continuous, plane to granular or verruciform. Thallus indeterminate, without perceptible hypothallus. Apothecia sessile frequent and often contiguous, to 0.72-1.04, m. diam. Disc dark bra= becoming black, plane becoming convex; margin concoleuroue with thallus, 0.04- 0.11mm. wide, entire, often becoming excluded. Excipulum thallinum 7a-05p wide; cortex absent or to c. 25p wide, structure occluded, medulla 70-00p wide. Peripheral hyphae occluded or 3.4-8.5p wide, modul/ary hyphee 2.3-5.70 wide and algal cells to 11-19p diem. Excipulum proprium 15-49p wide 42.
laterally, expanding to 30-5qa wide above, hyphae wide. Hypotheclum 75.•130p deep. Theciu 100...1401* hit h; paraphyses 1.7-2.8p wide, apices to 4.0-5.1p, forming a red-brown epithecium, Asci (70-95) x (19-26)p, inner walls uniform. Spores polarilocular, thick walled, (18.8-31.3)25.4 x 12.2(9.7-45.0p, apical walls c. 3.5-3.9i thick, lateral walls c. 1.0-1.7p, porus c. wide when present and septum well developed. Thalline reactions:- X.., C-, I-, P=. Hypothecium I-, thectum /4- blue. Plant muscicolous.
Variability and identification The variation of the thollus morphology of Rinodina mninrea lies well within the range expected in this genus. The apothecin, however, appear to be more variable which can make the identification of the species difficult. ninodinn mninrea is generally distinguished from R. turfacen by ito convex apothccia and lack of a columnar apothecial cortex. Unfortunately the first of these characters has been shown to be a poor one because the apothecia may sometimes
remain plane with a prominent margin. Conversely r. turfacea may occasionally develop convex apothecia. Final identification c these two species therefore rests upon tho presence or absence of a columnar cortex.
Taxonomic remarks Rinodina mniareoiza has been included in the present 43. species as it was found to be almost identical with it. Nylander (loc. cit.) seems to have established the species on the basis of a X* yellow thallina reaction. It hen not been possible to find any such reaction in the material examined and even if it had been found it would not have been regarded with sufficient importance to warrant a separate species. There can be no doubt that Rinodina mniarea and R. turfacea are very closely related taxonomically and their separation as species depends solely on the importance attached to the characters of the cortex.
7) Rinodina macronatula H. Magn. in iota Sort. Gothoburg., 17: 293 (1947).
Authentic material ifolotype:- Brenner 301b (S) - Rinodina mucronatula, Jenisejak: Xrasnojarsk, Brenner, 1876.
Description
Thallus thin, light grey to grey-brown, continuous, plane to somewhat verruciform. Thallus indeterminate, without perceptible hypothallus. Apothecin sessile, frequent and usually contiguous, to c. 0.47mm. diva. Disc black, persistently plane; margin concolourous with thallue, c. 0.07m m. wide, entire and persistent. 44.
Exeipulum thallinum o. 75p wide; cortex cellular, o. 25p wide, medulla c. 50p wide. Peripheral hyphae not pigmented, to o. 4.611 yids, medullary hyphae a. 3.0 and algal cells to c. 16p diem. Excipulum proprium o. 15p laterally and above, hyphae 43. 1.1p wide. Hypotheoium o. 15! deep. Theoium o. 90µ high; paraphysee c. 2.3z wide, apices to o. 5.1p, forming alight brown epitheciue. Asci o. 85. x 19p, inner wells uniform. Spores polarilooulor, thick walled, lumina becoming occluded and with thin outer wall forming mucronate apices (23.9-27.4) 25.9 x 9.9 (9.1-11.4)p, apical walls c. 4.411 thick, lateral walls c. 2.1p, porus a. 2.8p wide and septum well developed. Thall♦ne reactions:- E;:, C:, P.:. Hypethecium I-, thecium I+ blue. Plant lignicolous.
Variability and identificatige Nothing is known of the variability of this species. The identification of Rinodina muoronatula is dependent on the recognition of the muoronate spores although the species could hardly be confused with other large spored species such as R. mniareg and R. turfacog. These species have much larger apothecia and differ respectively in having convex apdthecia and eoolumnar apothecial cortex. R. muoronatula is similar to R. arohaea in the morphology of the thallus and size of the apothicia. %axonomio remark,' From the taxonomic point of viewRinodina mucronatula appears to tall midway between the R. archaeit amine jurfaom grow; of species. The thin apical, outer spore walls might also in- dicate some relationship with R. confraRosa and fit. copsoolanst although the latter species is less closely related in other respects.
8) EAmaamalammThi. ire in Lichenogr. Scandin., 1: 193 (1871) Parpelia nimbosn Fr., Lichenogr. Europe Reform.: 129 (1831). RI:100m nisitosa Ignge, Lich. Novya Zemlya: 24.9, tab. 5, fig. 9-11 (1928). - non Leads& obaloonya Sommerf., Suppl. nor. Lappon.: 150 (1826).
Exallocati Ansi Lang. 108 (K) Rinodina araniocolp, 109 (K) • RAnocUna anniocolt var. slabga; Hepp 82 (K) Poore niabosa.
Descriution
Thallus thiok, light grey to ochraceous, subaquamulose, plane and sometimes pruinose. Thallus determinate, without perceptible hypothallus. Apotheoia sessile, frequent and usually contiguous, to 0.61-1.12mm. dive. Disc black or pruinose plane or becoming convex; margin concolouroue with 46. thallus, 0.07-0.14mm. wide, entire and persistent or becoming excluded. Excipulum thallinum 55-155p wide; cortex absent, cellular or structure occluded to c. 30p wide when present, medulla 40-155p wide. Peripheral hyphae 4.0-5.7p wide when present, medullary hyphae 2.3-4.6p wide and algal cells to 15-22p diem. Excipulum proprium 10-15p wide laterally, expanding to 10-25p above, hyphae 1.1-1.7p wide. Hypothecium 50-140p deep, merging with hyaline stipe. Thecium 95-130p high; paraphyses wide, apices to 4.0-5.1p, forming a red-brown epithecium. Asci (50-70) x (15-19)p, inner walls uniform. Spores polarilocular, lumina becoming dilated and later occluded, (17.1-22.8) 19.7 x 9.2(8.0-10.8)p, apical walls c. 2.6p thick, lateral walls c. 1.2p thick before occlusions porus excluded from start, septum well developed. Thalline reactions:- X:, C:, I:, Hypothecium I-, thecium /+ blue. Plant muscicolous.
Variability and identification The material examined indicates that Rinodina nimbosa is a relatively constant species except perhaps for the morphology of the apothecia. It is unusual to find that apparently well developed specimens of a species can either possess plane apothecia with persistent margins or convex apothecia with the margins excluded. 47.
The subsquamulose thallus habit makes identification of the species easy. In some instances where the thallus is poorly developed Rinodina nimbosa could be mistaken for R. mniarea or R. turfacee but can be distinguished from both of these species by the smaller spores with dilated or occluded lumina.
Taxonomic remarks At first sight the spores of Rinodina nimbosa appear tote devoid of lumina but closer inspection of the younger spares indicates that they are of the polarilocular type. It would appear therefore that this species is related to the other large spored polarilocular species, R. mniarea and R.turfacea. The subsquamulose thanes habit should not be given taxonomic precedence over the spore type and the species has therefore been removed from Sect. Placothallia.
9) Rinodina trevisanii Koerb. In Parerg. Lich.: 70 (1859). Psora trevisanii Hipp. Flecht. Europ., no. 80 (1853). - Diploicia trevisanii Mass., Goneac. Lich.: 20 (1854). - Rinodina convexula H. Magn., Acta Hort. Oothoburg., 17: 290 (1947).
Axe i cc at i
Hepp 80 (K) Poore trevisanii; Mass. 293 (K) Rinodina leprosa; Rabehh. 737 - Rinodina trevisanii. 48.
Authentic material Isotype:- Hepp 80 (K),- Poore trevisanii, St. Moritz.
Description
Thallua thin or evanescent, light grey to brownish, continuous or rimoss and plane. Thallus determinate or indeterminate, without perceptible hypothallue. Apothecia sessile, frequent and sometimes contiguous, to 0.35-0.53mm. diem. Disc dark brown becoming black, plane becoming convex; margin concolouroue with thallus, c. 0.04mm. wide, entire becoming excluded. Excipulum thallinum 50-75p wide; cortex absent or cellular, to c. 15p wide, medulla 50-65p wide. Peripheral hyphae 3.4-5.7p wide, medullary hyphae 2.8-3.4p and algal cells to 14-15u diam. Excipulum proprium c. 10p wide laterally, expanding to 15-25p wide above, hyphae 1.1-1.7p wide. Hypothecium 20-90p deep. Thecium 70-90p high; paraphyses 1.1.4.7p wide, apices to 4.0-4.6p wide, forming a red-brown epithecium. Anci (50-70) x (14-23)p, inner walls uniform. Spores polarilocular, thick walled, (13.1-23.4) 17.9 x 8.6(6.6-10.3)p, apical walls c. 2.0-2.6p thick, lateral walls c. 1.4-1.6p, porus c. 1.1.-1.7p vide and septum well developed. Thalline reactions:- 4, C-, 1-, P=. Hypothecium I-, thecium i+ blue. plant corticoloua. 49. Variability and Identification Little is known of the variability of Rinodina trevisiana as few specimens have been examined. The species is very similar to R. anthems in anatomy but may be distinguished by the convex apothecia.
Taxonomic remarks It has not been passible to find any significant differences between Rinodina convexula and R. trevisiana and the two species have therefore been combined. The species shows a
very close relationship with R. archaea and has only been maintained because the convex apothecia and I-hnothecium are characters not Observed in R. archaea.
10) Rinodina turfacea Koerb. in Syst. Lich. German.: 123 (1855). Lichen turfaceus Wahlenb.. Flora Lappon.: 408 (1812). - Non Lecanora sophodee var. otbata Ach., Lichenogr. Univers.: 678 (1810). - Rinodina lignaria R. Magn. (pro parts), Svensk. Sot. Tidskr.. 30: 259 (1938).
Exsiccati Ansi Lang. 108 (X) Rinodina turfacea var. microcarpa, 439 (IC) - Rinodina turfacea var. depauperata; Arn. 452 (IC) - Rinodina turfacea; Fells. 111 (K) - Lecanora scphodes var. 50. succedens; Hepp 83 (K) - Psora turfacea, 84 (1) - Psora turfacea var. yechnea, 85 (K) Poore turfacea var. aicrocarpa; Krypt. Vind. 1868 (K) • Rinodina turfacea; Lich. Fenn. 442 (K) - Rinodina turfacea f. ecrustacea.
Authentic material Syntype:- (UPS) - Lichen turfaceus, Altensis Norvegia, Wahlenberg.
Description,
Thallus thick, thin or evanescent, light to dark grey brown, continuous or cracked, plane or more usually verruciform. Thallus indeterminate, without perceptible hypothallus. Apothecia sessile frequent and often contiguous, to 0.90-1.55mm. diem. Disc dark brown becoming black andipersistently plane; margin con- colourous with thallus, 0.07-0.14mm. wide, entire becoming cronulate and persistent. Excipulum thallinum 85-103p wide; cortex absent laterally or cellular to C. 20p wide, columnar and hyaline below, expanding to 40-70p wide, medulla 50-85p wide. Peripheral hyphae laterally 5.1-7.4p wide, medullary hyphae 2.8-4.8p wide and algal cells to 12-18p diem. Excipulum proprium 1O-20p wide laterally, expanding to 20-45p above, brownish, hyphae 1.7-2.3p wide. Hypothecium 25-90p deep. Thecium 115-150p high; paraphyses 1.1-2.3p wide, apices to 3.4-6.3p, forming a red brown epithecium. 51. Ascii (73-100) x (28•29)p, inner walls uniform. Spores polarilocular, thick walled, (21.4-46.5)28.4 x 13.2(10.3-18.2)Mo apical walls c. 3.3-4.7p thick, lateral walls c. I.7-2.2p, porus c. 3.0-4.0p wide and septum well developed. Thalline reactions:- K:, C:, I:, P. Hypothecium I-, thecium I+ blue. Plant muscicolous or rarely lignicolous.
Variability and identification Morphologically Rinodina turfacea is a relatively constant species but the internal anatomy and particularly the spore sires are rather more variable. The spores are usually narrowly ellipsoid but in the lignicolous specimens examined they have been broadly ellipsoid and shorter. The species bears a strong superficial resemblance to the auscicolous forms of Rinodina archaea but the spores of R. turfacea are generally much larger. Confusion may also occur with ,R. mniaree in the rare instances when the apothecia become somewhat convex. R. mniarea may always be distinguished from ace.. by the lack of a well developed, hyaline and columnar lower cortex in the apothecium margin.
Taxonomic remarks Rinodina lignaria is virtually identical with R. turfacea except in the size and broadly ellipsoid shape of the spores. Because of the lack of any other differences and bearing in mind that 52. the abnormal substratum could account for the variation of spore size, the two species have been united. R. roscida Should almost certainly be included in this species also but it has not been possible to sne any authentic specimen of this species. The species differs from R. turfs*** only in the presence of pruina and oxylate crystals in the thalami. Those Characters havo also been noted in some specimens of 130 =taros and R. niMbosos Rinodina turfaces Is undoubtedly very closely related to ,R. mniarea. There is also a relationship with n. nrchaes but this would seam to be lees close on account of the smaller spores.
11) Rinodina albana Mee. in Ricerch. Acton. Lich.: 15 (1852). ..BReatualeaMass. Verb. zool-bot. Gee. Wien 1: 221, fig. 0 (1351).
Exsiccati P.m. 1553 (X) Rinodinn sophodes f. albana; Repp 883 (K) - Psora horiza vrr. orbiculeris; Koerb. 245 (K) Rinodina horizn var. orbicular's; Krypt. Mind. 161 (K) Rinodina_pyrina; Mass. 21G (K) - Rinodina albana.
Authentic material Mass. 216 (K) Rinodina albana. 53.
Description
Thallus thick or thin, light grey or dark grey-brown, rimose, verruciform and sometimes granular. Thallus indeterminate, without perceptible hypothallus. Apothecia sessile, often con- tiguous, to 0.50-0.65mm. diem. Disc dark brown, plane often becoming convex; margin concolourous with thallus„ c. 0.07mm. wide, entire and persistent. Excipulum thallinum 60-95p wide; cortex cellular or absent, to c. 13p wide, medulla 45.-83p wide. Peripheral hyphae pigmented or not, 4.004.6p wide, medullary hyphae 2.8-4.6p and algal cells to 11-23p diem. Excipulum proprium 15-20p wide laterally, expanding to 15.40p above, hyphae 1.1.4.7p wide. Hypothecium 50-95p deep. Thecium 75-130p high; paraphyses 1.7-2.3z wide, apices to 3.4-4.6p, forming a red-brown epithecium. Asci (65-85) x (15-19)p, inner walls uniform or not. Spores polarilocular, thick walled, (16.5-23.6)20.2 x 0.7 (8.0-11.4)p, apical walls c. 2.9-4.1p thick, lateral walls c. 1.6p, pores c. 2.3..2.8p wide when present, septum not well developed. Thalline reactions:- K:, C-, P-. Hypothocium and thecium /+ blue. Plant corticolous.
Variability Nothing is known about the ecological variations of this species as so few specimens have been examined. The species is very similar to Rinodina seehodes in the morphology of the thallus and to R. arches:: in its internal structure. It sky be separated from both species by the brown apotheoial disc and by the morphology of the spores which lack any well developed septa.
Taxonomic remarks The internal anatomy of this species bears a super- ficial resemblance to Rinodina arehaea and some taxonomic relationship might therefore be suggested. However, the lack of a well developed:spore septum probably indicates a closer relationship with R. exigua although the spores are larger than in most species of that group.
12) Rinodina consooians H. lagn. in Acta Hort. Gothoburg., JIPJ 242 (1947).
Authentic material, Syntype:- Brenner la (S) Rinodina consocians, Jenisejak: Nikandrovskij Ostrom, Brenner, 1876.
Description
Thallus thin, light grey to oohraoeous, continuous or rimose, plane or verruciform. Thallus determinate or indeterminate, without perceptible bypothallus. Apothecia 55. sessile, sometimes contiguous, to 0.43-0.58 a. diem. Disc black, plane becoming convex; margin conoolourous with thallus, c. 0.07mm. wide, entire becoming excluded* Excipulum thallinum 55.75p wide; cortex cellular, C. 15st wide, medulla 40-60p. Peripheral hyphae not pigmented. 3.4.4.6p wide, medullary hyphae 2.84.0p and algal cells to c. 14i dies. Exoipulum proprium o. 10i wide laterally, expanding to 20-25p above, hyphae 1.1.1.70 wide. Ifypotheoiurs 40-6011 deep. Meatus 70-90a high; paraphyses a. 1.7p wide, apices to 4.0-5Ap* forming a brown or red-brown epithecium. Asci (60-65) x (15-19)p, inner wall uniform* Spores polarilooular, lumina becoming rounded, (17.1-21.7) 19.3 x 9.1(8.5-10.3)% apical walls c. 3.0-3.5µ thick, lateral walls o. 1.3-1.5p thick, outer apical wall very thin and spores dilated maridionally4 septum rarely developed. Thalline reactions:- X-, C-10 I-, P-* Hypothecium I-, thecium I+ blue. Plant corticolous.
Variability and identification It is evident from the limited amount of material studied that the apotheoia at Rinodina oonsooians may become markedly convex, to the exclusion of the margin. The internal anatomy displays little variation although further specimens should undoubtedly be examined. Rinodina consociang is generally distinguished from 56. lajobest by the lighter thallus and the tendency of the epotheoia to become convex, features not common in the latter species. More particularly it'is characterised by the spores which are meridionally thickened and have very thin outer, apical walls.
Ta on sic re, arrks The rpeoies appears to be rather isolated taxonomically although the spores show features in common with both Sildtifinii and Ig...Stakeia•
13) Ai:iodine oortioola Arm. in Verh. sool.-bot. Ges. lien, 370 (1879). Rinqaina teiohouhila var. corticola Arm, Verh. Boa.- bot. Ges. Wien, 18: 952 (1868).
Ex$100ati Arn. 1787a (K) - Rinodina cortioola; Kxypt. 2079 (K) Rinodina cortical*.
AuthentiQ material Arm 1762a (K) Rinodina cortiool*, Graden - Tirol, Arnold, 1898. 57. Desortutio;
Thallus thin, light or dark grey, continuous or rimose and plane or of discrete verrucae on a dark hypothallus. Thallus determinate or indeterminate, sometimes limited by a bleak, entire hypothallus. Apotheoia sessile, frequent and often contiguous, to 0.50-0.83mm. dies. Diso dark brown becoming black, plans, sometimes becoming markedly convex; margin cenoolourous with thallus, 0.04-0.07mm. wide, entire and usually persistent. Exuipulum thallinum 70-1151 wide; cortical structure occluded or of columnar hyphae, 10-54 wide laterally, eiummme expanding to 40-90:: wide below, medulla 30-65p wide. Expanded and pigmented peripheral hyphae absent, medullary hyphae 2.8-4.6u wide and algal cells to 9-24111 diam. Exuipulum proprium 10-15u wide laterally, sometimes expanded to e. 1512 above, hyphae 1.1-1.70.Aide. Hypotheoium40-95:: deep. Thecium 90-115µ high; paraphyses 1.1-1.7vLyide, apices to 4.C6.5.ipmide, forming a red- brown epithecium. Asoi (70-85) x (15-22)14 inner walls uniform or not. Spores polarilocular, thick walled, (17.1-24.5) 20.9 x 10.0(8.0-12.0)u, apical walls o. 5.5µ thick, lateral walls a. 1.5-1.714 porus a. 1.91i wide when present and septum developed at maturity only. Thalline reactions:- 4 yellow, C:, I:, P. gypo- theoium I-, theoium blue. Plant corticolous. 58.
Variability and identification The thallus ofMasigiajoaktali shows a wide range of variation from continuous or rinse to being composed of discrete verruoae. In the latter case the thallus may have a very dark aspect as a result of a predominating dark hypothallus.
Runemark (1956) has suggested that in the genus socaroon the predominance of the hypothallus nay be influenced by such factors as shade and. humidity. The species may be distinguished from all other members of the Rinodinit archaea group of species by the R: yellow thallue reaction and from individual members of the group by the morphology and colour of the thallus and more particularly by the massive apothecial cortex owsposed of columnar hyphae.
Taxenomic remarks The internal anatogy of the species shows a very Close relationship with Rinodina arehaea although the spores do not develop a prominent septa. The structure of the aortas is one of the most important characters of R. corticola and although a columnar oortem may sometimes be present in R. ardhaea it is rarely well developed and the lateral part of the apothecium is boarded by expanded and pigmented peripheral hypbae. Magnusson (1947) used minute differences of cortical structure to distinguish between species but this does seen to be one of the few cases where the character shows no intergradations and can be used as a feature 59. of major taxonomic importance.
14) Hlnodina orassesceqp Arm. Jai/lora, Regensburg, ID 318 (1880.
Authentic material Nyl. 29015 (H) • Lecanora confrastosik var. ,orassescens, Korpilahti, Lang, 1873.
DescriTAiqn
Thallus thick, ochraceous to light brown, of discrete or oontiguous verrucae, plane and embeqpamulose at periphery. Thallus indeterminate without perceptible hypothallus. Apotheola sessile, frequent and sometimes contiguous, to c. 0.90mm. diam. Disc black, rapidly becoming convex or even half-globose; margin conoolourous with thallus, c. 0.07mm. wide, entire but rapidly becoming excluded. Excipulum thallinum o. i30µ wide; cortex hyaline of columnar hyphae, o. 65p wide, medulla o. 65p wide. Peripheral hyphae not expanded or pigmented, medullary hyphae 0. 2.8p wide and algal cells to o. 11p diem. Exoipulum proprium c. 20p wide laterally, expanding to o. 25p at periphery, hyphae a. 1.7p wide. Hypotheoium o. 130p deep. Thecium c. 100p high; paraphrase c. .1.7p wide, apices to a. 3.4p wide, forming a brown epithecium. 60.
Asa c. 85 x 194x, inner walls uniform. Spores polarilocular, thick walled, (194-24.5)21.7 x10.5(9.1-12.5)P4 Vioal vans a. 3.6p thick, lateral walls a. 1.3p, porus c. 3.1µ wide when present and septum developing late. Thalline reactions:- X yellow, C:, I:, P:. Hypo- theoium X-, theoium I.:: blues Plant siliceous.
XerUbilitr and identification Nothing can be said about the variability ofaselias cram:wens as it has only been possible to examine the one authentic specimen. The species cannot, however, be easily confused with others on acocunt of its subsquamulose habit and massive, oolumnar apotheoial cortex.
Taxonomic remarks The polarilocular spores and subsquamulose habit make this an isolated species in the taxonomic sense. The latter with character is more usually associated/species whose spores entirely lack lumina. The presence of a thick oolumnar cortex might suggest some relationship with Rinodine. aortic:01g.
15) Rinodina bidiella Th. Fr. in Lichenogr. Soandins, 1: 199 (1871). Leonora badiella Nyl., Herber. Ube. Fenn.: 110 (1859). 61. 41uthentie sattrilal Nyl. 28756 (H) Ifecariora badiella, Asikkala, Norrlin, 1863.
Desorixtiotk
Thallus thick or thin, dark brown, ramose, plane or granular, or of discrete verrucae on a black hypothallua. Thal us determinate or indeterminate, sometimes limited by an entire, black hypothallus. Apothecia sessile, often contiguous, to 0.40•0.90mm. diem. Disc black, plane becoming slightly convex; margin ooncolourous with thallua, 0.07-0.14 m. wide, entire and persistent. Excipulum thallinum 6O-65µ wide; cortex cellular, 10-15p wide, medulla 45•55µ wide. Peripheral hyphae 14..6-5.7p wide, medullary hyphae 2.8-5.1p and algal cells to 12-23p diem. Exoipulum proprium 15-20p wide, expanding to 25-40p above, hyphae 1.1.-1.7p wide. Hypothecium 110-130p deep, dark stipe sometimes developed. below. Theoium 90-110p high; paraOyses 1.7-2.31 wide, apices to 3.4-5.7p wide, forming a brown or red- brown epithecium. Asoi (50-65) x (15.49)p4 inner walls uniform. Spores polarilocular, soon developing thick uniform walls, (13.1-20.5)15.8 x 8.8(6.3-11.4)N apical walls o. 2.0-2.41 thick, lateral walls el. 1.5-1.7p, pores very quickly excluded and septum prominent at maturity. 62.
Thalline reactions:- 4 yellow, C:, I:, P:. HYPo" thecium I- or + blue as thecium. Plant siliceous. yggialUdgueLthatinsvittes The thallus morphology of this species varies between thalli composed of thin, small, dispersed verrucae on a bleak hypothallus and the= which are thick, rims* and sometimes granular without any perceptible hypothallus. The spore sise and morphology shows an unusually, wide range of variation but it is possible to distinguish the species from other brown, saxi- oolous species such as Rinodina stiliviaA and R. crassesoens by the spore characters. The former species has thin walled spores and the latter much larger spores.
Taxonomic remprks This species has been plaoed in the polarilooular group because the lumina in very young spores are angular and quite typical of this group of species. It is one of the few species in the group that have thick walls and rounded lumina in mature spores. A tendency towards this state has been noticed in other species.
16) Rinodina oolobina Th. Fr. in Lichanagr. Soandin., is 205 (1871). Lecanoru colobina Aoh., Lichenogr. Univers.; 358 (1810). 63.
Exalocati Ansi Lang, 305 (K) Rinodina leurost; Arra. Ion. 99, 508 (K) • Naglialugjekkal Larb. Herb. 91 (BM) jassamusagtka; Lich. Fenn. 444. (K) Rinodina aolobinv Lo3ka Hung. 22J (K) Wawa ebb; al Mass. 294b (K) Mandl* lenrosa war. leoideina f. ganniochroil; Rabenh. 965 (x) • Rinodina colobinik.
A101100441_ material
None 8$0324 aggssfatarm
Thallus thin or evanescent, dark grey, continuous or more usually rimose, granular and matt. Thallus indeterminate, without perceptible hypothallus. Apothecia sessile, not frequent and rarely contiguous, 0.37-0.47mm. diam. Disc black, becoming only slightly convex; margin concolourous with thallua or lighter, 0.04-4.071m. wide, entire and persistent. Excipulura tballinum, 65..95µ wide; cortex cellular or absent, to o. 10u wide, medulla 65-95µ wide. Peripheral hyphae not pigmented, 5.7-7.4p wide, medullary hyphae 2.8-4.0u and algal cells 12-1811 diem. Exoipulun proprium 10-15p wide laterally, expanding to o. 15u above, hyphae 1.1-1.711 wide. Hypotheoium 70-115u deep. Thecium 60.45u high; paraphyses 1.1-2.31 wide, 64. apices to 4,0.5.7p wide, forming a blue-black epithecium. Asci (50-.70) x(13w19)p, inner walls uniform. Spores polarilocular, thick walled, (12.5-23.4)17.1 x 8.2(5.1-11.4)p, apical walls c. 4.4p thick, lateral walls a. 2.4p, porus a. 1.2u wide when present, septum developing late and spore with thick outer wall. Thelline reactions:- I:, C:, I:, P:. Hypothecium I-, thecium 1+ blue. Epithecium K+ and N+ violet. Plant corti- coleus.
Variability and identification As all the material examined is from the same locality in Britain nothing is really known of the potential variability of the species. It is easily identified by the very dark thallus, the blue epithecium and the polarilocular spores with a thick outer wall.
Taxonomic remarks The British material described above differs from all the Continental material examined to date in a number of ways. The thallus is darker and the apothecia much smaller than the Continental specimens but of greater importance are the spores which are polarilocular with a distinctly thickened outer spore wall in the British material. The spores of the Continental specimens are mischoblastiomorph and without any thickened outer spore wail. • Rinodino ,. ..- ...
sophodes (Ach.) Mass
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BOTANICAL SOCIETY OF THE BRITISH ISLES DISTRIBUTION MAPS SCHEME
Figure 5. Distribution of k terIC4-114 celcirla and jisjetsala 1St the British Toles. 66.
The present evidence strongly suggests that these types should be regarded as two different species. Unfortunately however, it has not been possible to examine the type material of Rinodina colobina and it is therefore not known to which taxa this name belongs. A new species should not be described until this position has been clarified.
17) Rinodina exigua S. Gray in Natur. Arrang. Brit. Plants, 1: 450 (1821). Lichen exiguus,Ach., Lichenogr. Suec. Prodrom.: 69 (1798). Psora exigua var. naculiformis Repp, Fledht. Europ., no. 79 (1853). - Berengeria exigua var. maculiforeis Travis., Flora, Regensburg, 38: 186 (1855). - Rinodina olea Bagl., Men. Acad. sci, Torino, ser. 2, 17: 403, fig. 3 (1857). Rinodina ramulicola, Kernst. ex Arno, Verh. tool.-bot. Geo. Wien, 46: 132 (1896).
Exsiccati Anzi Lang. 107 (X) Rinodina metabolica var. maculiformis, 304c (K) Rinodina horiza var. orbicularis, 377b (K) - Rinodina metabolica var. corticola, 378 (K) - Rinodina exigua var. corticola; Arn. 663 (K) - Rinodina metabolica„ 1381 (K) Rinodina exigua, 1654 (K) - Rinodina ramulicola; Arn. Monne. 101 (K) Rinodina exigua, 157 (K) Rinodina pyrina, 428 67. (X) . Rinodina maculiformis; Flora Hung. 920 (K) -Rpiodina Atm; Hopp 79 (K) ...tismsias var. maculiformis, 207 (K) - igmajtang; Kern. 2749 (K) -Rinctdinaremulioola; Larb. Herb 169 (K) Lecanoxa abodes var. adim,'262 (x) Leoanora roboris; Lich. Penn. 922 (i) Rinodina wrina; Nyl. Par. 128 (K) Locanora somhodes; Rabenh. 453 (K) - liktilaeaaim Reich. 85 (K) LecanoFa axigua.
Authentic naafi Aoh. Bale 1,.07 (BM) - Leoanora DarioleA
Deeoriution
Thallus thin or rarely evanescent, light to dark grey, continuous or rimose, plane or more rarely granular, matt. Thallus indeterminate, without perceptible hypothallus. Apothecia sessile or sometimes sub-innate, frequent and often contiguous, to 0,294.58mm. diem. Disc rarely dark brawn, always becoming black, plane or becoming convex; margin ooncolourous with thallus, 0.04•0.07tm. wide, entire and persistent. Bxcipulum thAllAnum 40-1141 wide; cortex when present either cellular or with structure occluded, to c. 15µ wide, medulla 25.410p wide. Peripheral hyphae absent, 2.8-5.1pwide when present but never pigmented, medullary hyphae 2.3-44.6p wide and algal cells to 11-18p diari. Excipulum proprium 10-15p wide laterally, expanding to 10-30p above, hyphae 1.1-2.311 wide, pale brown or colourless. 68. flypethesilmt 45-115µ deep, pale brown or colourless. Thecium 70.410n high; paraphyses 1.1-2.3p1 wide, apices to 3405.704 forming a dark ,brown or brown epitheolumi Asoi (45-65) x (13-20p4 inner walls uniform. Spores polarilocular, thick veiled, lumina becoming dilated, (11.4..22.8)16.5 x 7.5(54.-9.7)14 apical walls o. 1.5..344 thick, lateral walls o. 1.0.4.5)14 porus c. 0.6-01.8t wide than present and septum usually well developed at maturity. Thalline reaction's- la yellow (sordid) or more usually • 4, I:, Ilypothecium usually I. blue, like thecium. Plant cortioolous or lignicolous.
Tpriailitr and idegtlification eable_ magus is probably the most variable of all the species considered in this work. A. wide variation of thallus morphology is more easily accepted than a similar variation of sporesine and structure. The spore morphology varies from the typical polarilocular type with prominent septum to a morphology verging on the misohoblastiomorph type with rounded lumina and lacking a prominent septum. The species is readily recognised by the small sine of its spores and. is only comparable with AinocliLawrina in this respect. The species may be easily separated by their spore morphology, the spores of )t. ovrinahmving thin walls. 1.P
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BOTANICAL SOCIETY OF THE BRITISH ISLES PISTRIBUTION MAPS SCHEME
-iture 6. Tistributton of :Arc inn crigun it tiw! Tales, 70. Taxonomic remarks The spore sine and morphology indicates a dlose relationship with the saxioolous species Rinodina 000%414. The pigmented apotheoial cortex of that species, however, indicates a less close relationship in other reepeots. There is no doubt that R. satmp has exceedingly close taxonomic affinities with R. (vague, and this is discussed at greater length under the former species.
18) rtinodina furfurafta H. Magna. in Jlat Hort, Gothoburg.„ jt 236 (1947). kinodina sorediatg if. Maw., Acta. Hort, Crothoburgs, Es 292 (1917).
Authenti2 material Holotypes- (LISU) furfuracea (Rinodina golobina), Rstramadura: Santana, Tavares, 1943.
Ilemetitiga
Thallus thin, light grey, continuous or more usually rimose, plane or more usually granular, granules often dark grey or breaking up into lighter soredia. Manus indeterminate without perceptible hypothallus. Apothecia sub-innate or sessile, frequent and sometimes contiguous, to 0.61-0.79mm. 71. diem. Disc dark brown becoming black, plane or becoming well convex; margin conoolourons with thallus, 0.07-0.11mmt. wide. entire, persistent or beooming exoluaed. Exoipulum tiageotnum 50-7511 wide, cortex absent. Peripheral hyphae absent or to o. 4.0z wide, never pigmented, medullary hyphae 2.5-2.8µ wide and algal cells to 12-181: diem. Exeipulum proprium a. 1511 wide laterally, expanded to 15-50y above, hyphae 1.7.02.511 wide. Rypothecium colourless or pale brown, 75-9511 deep. Theoium colourless or pale brown, 75-90µ high; parsphyses 1.7-2.311 wide, apices to a. 4.011 wide, forming a brown epitheoium. Asci (50-60) x (15-17)14 inner walls uniform. Spores polarilocular, thick walled, lumina beooming dilated, (13.7-19.9)15.6 x 7.5(6.8-9.744 apical walls o. 2.1p thick, lateral walls a. 1.444 pores c. 1.7? wide when present and septum well developed at maturity. Thalline reactions:- K:, C:, I:, P. Hypotheoium I-, thecium I+ blue. Plant oortioolous.
Variability and identification The wide range of thalline morphology and more particularly the apotheoial morphology of Rinodina furfuraceg described above is due to a depauperate speoimen of the speoies previously known as R. sorediata. The innate apothecia, brownish hypotheoium and theoium, together with the depauperate 72. spores indicates very strongly that the specimen has been growing under adverse conditions. The general habit of the species shows some characters which could be -oonfused with Rinodina roboris and with R. erumoens, but the small, polarilooular spores convincingly separate R, furfuracea from these two species.
Taxonomio remarkk There must always be some doubt about the status of such species as Rinodina sorediata which are based on depauperate specimens. The species has, however, been included under R. furfuracea. Although the spores are polarilocular and of more or less the same sise as R. furfuracea it is still possible that it represents a separateapecies. Rinodike, furfuracea shows some relationship with R. exigua,on account of the small polarilocular spores with some- what irregular lumina.
19) Rinodina gIebulosit JUma. in Flora, Regensburg, 318 (188t..). var. glebulosa Ara., Flora, Regensburg, 12: 38 (1872).
Authentic) material Arn. 68 (K) -Rinodina glebuloes, var. mamma, Kunatein • Bich/tett, Arnold, 1859 and 1863. 73.
Thallus thick* dark grey, ramose beooming arapolate. Amiss. to o. 0.50mm. wide, plane or verruciform, often cracked. Thallus determinate or indeterminate, without perceptible hypothallus. Apothecia sub-innate, to 0.54- 0.61mm. diem, frequent and often contiguous. Disc black, persistently plane; margin oonoolourous with thallus, 0.04- 0.1ine. wide, entire and persistent. Excipulue thallima 65-115p wide, lacking a cortex. Peripheral hyphae 4.0-4.6p wide, pigmented or not, medullary hyphae 2.8..50441 wide and algal cells to 13-15p diem. Excipulum proprium a. 15p wide laterally, expanding to 15.4.30p above, hyphae a. 2.3p wide. Hypothecium 5540p deep. Theoium 75-851:t high; paraphysea 1.7*2.5pmide, apices to 4.0-4.6p wide, forming a brown to dark brown epithecium. Asoi (4550) x (15-15)14 inner walls uniform. Spores polarilocular, thick walled, lumina becoming dilated and finally occluded, (13.1-17.7)14.6 x 8.0(6.8-9.0p, apical walls o. 2.7-2.8p thick, lateral walls c. porus o. 1.1p wide when present and septum well developed. Thalline reactions:- K:, I:, C:, Hypotheoium and theoium I+ blue. Plant siliceous.
LviabilitY and _identification In view of the fact that only two specimens of 74.
Rinoditia slebulesa have been examined there is no basis for commenting on the variability of the species. The morpholog of the species bears a strong resemblance to R. *marls and it has been confused with this species in the past. R. &merle has, however, large mischoblastiomorph spores. R. Apbuloss can hardly be confused with any other polarilocular species on account of the spore sise.
Taxonomic remarks The species was originally described as a variety of Elmaillamitga and the spore morphology indicates that there is certainly a close relationship with this species. The sise of the thecium in relation to the bypath:Krim, together with the spore aise and morphology may also be taken as showing some relationship with R~ oonfraflosa, if these characters are to be regarded as holding any degree of taxonomic. weight.
20) Rinodina igmerboreq H. Magn. in Acta Hort. Gothoburg., 12a 24.1 (19k7)
Authentic material Holotype:. Brenner 1826e (s) hrperborea, Jenisejsk: Nikandrovskij Ostrovv Brenner, 1876. 75« pep Dti
Thallus thin or evanescent, light grey-brown, rimose or of discrete verrucae. Thallus indeterminate, without perceptible hypothallus. Apothecia sessile, frequent and often contiguous, to 0.46-0.58mm. diem. Disc dark brown, persistently plane; margin concolourous with thallus, 0.01+.-0.11mm. wide, entire and persistent. Excipulum thallinum4.0-70p wide; cortex usually absent but cellular when present, to o. 15 vi wide, medulla 40-70V mid*. Peripheral hyphae never pigmented, 3.4.-6.3p wide, medullary hyphae 2.8-4.0p wide and algal cells to 15-23a diem. with prominent pyrenoids. Exeipulum proprium 10-15p wide laterally, expanding to 15-2511 above, hyphae 1.1.4.7p wide. Hypothecium 4045p deep. Thecium 60-85p high; paraphyses a. 1.7}t wide, apices to 3.4-5.7p wide, forming a brown epithecium. Asoi (60..70) x (17-48)p, inner walls uniform. Spores polarilocular, thick walled, lumina dilated and spores dilated neritiiimmally, (14.2.45.9)17.7 x 8.8(74-10.5)p, apical walls o. 5.0..5.1p, lateral walls o. 1.5-1.6p6porus o. 1.4p wide when present and septum well developed at maturity. Thalline reactions:- K:, C:, I; blue or P. Hypo- thecium I-, theoium I+ blue. Plant corticolous.
Variability atilt identification Specimens of the species so far examined indicate that Rinodina hyperborea is a relatively constant species which might, 76. however, be overlooked as Umiak. It is distinguished from the latter species by the brownish thallws and by the spores which have a tendeney or be dilated meridienally. up err i Ially the species has a strong reseablanoe to a diminutive asjohis but the spore else and structure or this species is quite different. blAgantgaaiiirki There osn be no datibt that AtalliaLlandmas is very closely related to R. caltua, The rust that the spores hare a consistent tendency to be dilated meridionelly indicates that it should be treated as a separate species.
21) lanatalaclatiassa Ans. in Flora, Regensburg, s 39 (1872). duel iuridenens Ansi, Comment. too. Crittoganolog. Ital., (3): 158 (1862). - Lecenora oonioutit rlyl., Flora, Regensburg, Ifis 19 (1873). - 4eganoira solo* Idyl., Flora, Regensburg. s 68 (1873).
Exaianati Cronb. 158 (Bli• Bo '112SWAELIKS1221141
Authadamatarial Isotypes- Ansi Ltr. 20 (Y.) - silaap12112A221a, Monte 'Mean. 77. Pindaan Thallus thick or thin, oehraceous to dark grey-brown, arsolate. Areolae to 1.26-2.70mm. wide, often cracked and plane. Thallus determinate, limited by a dark, entire bypo- thallus. Apotheoia sub-innate, frequent end often contiguous, to 0.79-1.44mm. dime. Discs black, plane becoming slightly convex; margin darker than thallus, lighterthan disc, 0.044. 0.11mm. wide, entire becoming excluded. Excipulum thall4num lacking, at least in young apotheoia. Thalline cortex absent or to c. 20p.wide, cellular, peripheral hyphae 34-4.6p wide, medullary hyphae 2.3..3.411 and algal cells. to 9-1511 diem. Excipulum proprium brownish, 45...90u wide, expanding to 45-90p above, hyphae 1.7-3.4p wide, t oar- bonaGeous. Ilypothecium light brown, 70-155p deep merging with stipe below. Theeium 75-115p high; paraphyses 1.1-1.7p wide, apioes to 3.4-5.1p wide, forming a brown epitheoium. Asoi (50-65) x (18-23)p, inner walls uniform. Spores polarilooular, thick walled, (14.8..22.8)17.8 x 9.1(7.7-12.0)pi apical walls o. 2.3-2.9p thick, lateral walls o. 1.4-1.8% pores o 1.1-2.5p wide when present and septum prominent. Thalline reactions:- X:, Ct red, 0:. Hypothecium I- or I+ blue like theoium. Plant siliceous.
VariabAliti and identiftoation Rinodina luridescens seems to be a very constant species
9 .
Rinodina
loridestens (Anxi.) g_
Arn. •
A r 7 Km _,
NIL.. 9 Is 6 ,_,lcs._._...... ,
Malt /••...-
If
- -1- )
• .9.• 10 1 • -
0
BOTANICAL SOCIETY OF THE BRITISH ISLES DISTRIBUTION MAPS SCHEME
Figure 7. istributior cf 71Mdlre _lurideuccrs it the liritich Isles. 79.
well typified by the sub-innate apothecia and the mouse-brown thallus, Ct red. It seems uni kaly that this speoies could be mistaken for any other examined in the present work.
Taxonomic remark' It is apparent that Rinodina **pipets is synonymous with R. luridescens and Nylander (loc. cit.) states that R. aokodes is closely related to the former species and omits to state Why it was given specific rank. R. luridesoens is a taxonomically isolated species which has previously been placed in the genus BuAlkt in British lists. The apotheoial margin of the species is of great interest. Young apotheoia completely lack an excipulum thallinum and have a variously pigmented excipulum proprium. It appears that the expipulum proprium is capable of returning to the vegetative state because the outer part often has a medullary structure and trapped, viable algal cells are some- times found within it. In other material the hyphae of the exeipulum proprium intermingle with the medullary hyphae of the thallus so that the exoipulum proprium appears to be much reduced in sise. The species has been transferred to the genus Rinodina because the margin has the ability to become thalline, at least in part. 80. 22) Rinodina occulta nov. comb. kigaitiaLaMata Kcerb., Pares. Lich.: 188 (1800). - Lecencra teauiicola Nyl., Flora, Regensburg, 57: 307 (1874). Lecidea Interpolate stirt„ Scott. Nat., 1: 20 (1877-78).- Rinodina verrucariolldos H. Magn., Nyt Mag. Naturv., 87: 219 (1940.
Exsiccati Wight. 217 (K) - Lecidea spur**.
Description
Thallus thin, ochracecus, areolate or rarely rims.. rgl.eolae to 0.21-0.72mm. wide, usually contiguous, plane or rarely verruciform. Thallua indeterminate. with dark, entire and limiting hypothallus. Apothecia sessile, frequent but rarely contiguous, to 0.14.0.28mm. diem. Disc black, plane or becoming convex; margin concolourous with disc, rarely with thallus, c. 0.04mm. wide, entire, perste:tont or sometimes excluded. Excipulum thallinum 40-.85F wide; cortex 10-30F wide, structure occluded by piGmentation, medulla 23-55U wide. peripheral hyphae 4.0-3.7i wide when distinguishable, medullary hypbae 2.3-4.041 wide and algal cells 11-17p diem. Fxoipulua proprium t0-15F wide laterally, expanding to 10-40F above, hyphne 1.1-2.341 wide. Hypothecium 1005311 doop. Thecium 53.8541 high; paraphyoca 1.1-2.3F wide, apices to 4.0.65.741, forming a brown 81. epithecium. Asci (430.3°) x (13.-20)p4 inner walls uniform. Spores poiarilocUlar, thick walled, (12.1-18.2)14.3 x 7.4 (5.7-8.5)µ, apical walls c. 1.9.03.2p thiCke lateral walls c. 1.0-1.8p, porno c. 1.1.02.0p wide when present and septum developing late. Thalline reactions:- K: Cr + yellow, C", I-, P-. Hypothecium I-, theCium + blue, Plant siliceous.
Variability and identification The apothecia of Rinodina occults vary widely both in their size and morphology. They may be so minute that they appear to be perithecia at first sight and hence Magnusson's
R. verrucariodes. In such specimens the disc is usually convex and the margin excluded. Very rarely the margin of the apothecium is not pigmented (R. tequlicola) which could lead to difficulties in identification. The thallus of the species may sometimes consist of dispersed areolae which because of the dark hypothallus, gives a very dark aspect to the whole plant (R. interpolate). Identification of the species is usually made easy by the pigmented apothecial margin. This could horever, lead to confusion with Rinodina discolor although the spores of R. occulta, in contrast to the latter species, are small and polarilocular. in the few instances when the apotheciel margin lacks any pigment the species may be separated from. R. saline, a species with similar spores, by the areolate thallus. For distribution in the British Isles see under R. biloculata. B2. Taxonomic remarks.
Because of the pigmented margin Rinodina occults shows
some affinity with R. discolar, but the spores demonstrate that it
belongs to the polarilocular group of species. The small site
of the spores and the late development of the septum, coupled with
the rather irregular shape of the lumina indicates that the species
is probably quite closely related to R. saline.
23) Rinodine saline Degel,
in Ark. Bot., 30 (1): 55 (1942).
Non Zeora metabolica var. demissa Flk, ex laot., Jber.
Schlos. Ges vaterl. Knit., 27: 124 (1849). Lecanora sUbexigua
Nyl., Florn, Regensburg, 57: 308 (1374). - Lecidea alocizoides
Leight., Lich. Flora Great Brit., edit. 3: 323 (1879). - Rinodina
saline Doge l., Uppsala Univ. Arsskr., 11: 192 (1939) nomen nudum.
Exsiccatl Arn. 1040 (K) Rinodina demissa; Bepp 645 (K) Psora demissa; Johns. 190 (MM, RMW) Lecanora exigua, 367 (nm) - Lecanora laovigata; Krypt. nude 2080 (K) Rinodina domissa;
Larb. Herb, 261 (K) Lecanora oxiva; Mudd 107 (13M, E, K) -
Rinodina exigua.
..MM IE••••••••••••,
Thallus thin, light to dark grey or ochraceous, 83. continuous or more usually rimose, plane becoming verruciform. Thallus determinate or indeterminate, without perceptible hypo.- thallus. Apothecia sessile, rarely sub-innate, frequent and often contiguous, to 0.36-0.81mm. diam. Disc dark brown becoming black, plane becoming convex; margin concolourous with thallus, 0.04-0.07mm. wide, entire and usually becoming excluded. Excipulum thallinum 35-85p wide; cortex cellular or structure occluded to 15p wide when present, medulla 30-75p wide. Peripheral hyphae not pigmented, 4.003.3p wide when visible, medullary hyphae 2.3-4.8p wide and algal cells to 13.•19p diem. Excipulum proprium 15-20p wide laterally, expanding to 30-45$ above, hyphac 1.1 -1.7p wide. Hypothectuli 25-95p deep. Thecium 70-95 high; paraphyses 1.1-1.7p wide, apices to 3.4-4.3112, forming a brown epithecium. \Bei (50-70) x (15-21)'1, inner walls uniform. Spores polarilocular, thick willed, (11,049.8)15.1 x 840248-11.0,p, apical walls a. 2.1-2.5p thick, lateral walls co porus c. 1.4-2.0p wide and septum poorly developed. Thallino reactions:- K-, C-, 1-, P-. Hypothecium and thecium blue. Plant siliceous or more rarely calcareous.
Variability and identification As the description indicates, Rinodina saline is one of the most variable species dealt with in the present work, Despite this great variability, however, the species would be difficult to confuse with other saxicolous species because of the spore type. R. occult4 also has spores which are polarilooular, small with irregular lumina and indistinct septa but my easily be separated fress11. waling by the dark apotheoial margin.
Taxonomic remarks There can be no doubt that Rinodina subs:Aiwa is synomymous withjk.eium and that the sub-innate apotheoia and °ohm:sous is represent a depauperate form of the latter. The internal anatomy and measurements are virtually identical in both species. Rinodina saliva is undoubtedly very closely related to the oortioolous species R, exigua and there must be a strong case for combining the two species as varieties if not as one species. The present work, however, has indicated a number of differenoes, enumerated below, which may or may not be wholly constant.
Rinodina sOina Magae2 —Lai3iii Spores broadly ellipsoid narrowly ellipsoid thiok walled thinner welled Excipulum proprium broad narrow Apotheoium - convex - plane margin - excluded t persistent
Until further material has been studied the question of uniting these two species must be put aside. e5•
0 3 AS,
•
-.. 2
Ronod$no ll solina Degel •• i 0 1 . .111 • 14. ' -4 A
CM , Ci3. '0401911( --....s--- NILES I 6 3 1 Im
. ) 1 1 • •
• ,i- • .
3 4111-- _ •• • • • • • 2
ai•
11:31 .9 0
,i(--- 0 4
BOTANICAL SOCIETY OF THE •RITISH ISLES DISTRIBUTION MAPS SCHEME
Figure 8. Distribution of /Izallazairja in tho British Laos, 86. 24) Rinodinet aecuekula B. de Lesd, in Bull. Soc. bot. Fr. a 516 (1906) Lecenora aeauajula Nyl., Flora, Regensburg, ,§2: 388 (1884).
Exsiocitt Zw. 956 (R-101. P.M. 2888) - Leoanora 40010.1p.
Authentio materkal Nyl. 28619 (H) tecanora aecuatpla, Budapest, ',Oka, 1881.
Description Phallus thin or evanescent, light grey to ochraoeous, of dispersed, sourty granules. Thallus determinate or in- determinate, without perceptible hypothallus. Apothecia sessile* rarely contiguous, to 0.36-0.54mm. diam. Disc black, becoming convex and half-globose; margin concolourous with thallue at first, later with disc, 0.04-0.07mm. wide, entire, rapidly becoming excluded. Beoipulum thallinum 50-80p, wide; cortex cellular, pigmented or not, 15-2511 wide, sometimes expanding to 0. 40µ below, medulla 25-6511 wide. Peripheral byphaa 4.6-6.5p wide, medwonry hyphae 2.8-5.1pnide and algal cells to 14-19p diem. 87. Excipulum proprium 10-15n wide laterally, expanding to 15-30p above, hyphae Ada. *path:le/um 110-115µ deep. Thecium 75w115µ high; paraphyses 1.7-2.3pL wide, apices 5.1-7.4 wide, forming a dark brown epithecium. Asci (4540) x (13.45)14 inner walls uniform. Spores polarilocular at first, walls becoming thin and uniform, later apparently without lumina, (11.4-17.1)1442 x 6.2(5.1-6.9)V4 &Pical and lateral walls less than 1.0n thick at maturity, porus absent and septum well developed. Thalline reactions:- g, C:, I:, P=. Hypothecium I-, thecium feint blue. Plant calcicolous.
Variability The morphology of the apothecium is particularly variable in this species. The margin is initially the same colour as the thallus and the disc plane. However, with increasing age the margin becomes darker and the disc convex until the apothecium is finally half-globose and the margin excluded. The species bears a strong superficial resemblance to Rinodina biso3offii but the else and structure of the spores readily distinguish it from that species.
Taxonomic remarks Old but apparently healthy spores of this species 38. appear to look lumina but the young, and also the occasional nature spore show that the species more truly belongs to the polarilocular group. The morphology and shape of the spores shows a relationship with &Wine mina and pokvsnora.
25) Rinodina MAW:off* Mass, in Fromm. Lich.: 26 (1855). Ps9ra bisohoffii Hopp, Flecht. Bump., no. 81 (1853). - Berenaeria bischoffii Travis., Flora, Regensburg, 38: 186 (1855).
Bzeiccati Ansi It. 222 (K) Rizodiria bischoffii; Hepp 81 (K) - Poore, bischoff4; Hass. 113 (K) -Rinodinabischoffilj Rabenh. 84.3 (K) - RI:Jo:dine bisitoffii var. immoral.
Authentic materilal Isotypo:- Hepp 81 (10 Psora bischoffii.
Descridion
Thallus evanescent, light grey, of disoretel scurfy granules, or rarely thin, grey to ochraceous, rimose and piano. Apothecia sessile, sometimes contiguous, to 043-0.58mm. diam. Disc black, plane becoming convex; margin conoolourous with thallus becoming darker and conoolourous with disc, 0.07-0.11mm. u9. wide, entire becoming excluded. Exeipulum thal14mal 40-94 nide; cortex usually present, to 25µ wide, medulla 20e'70µ vide. Peripheral hyphae Pigmented or not 5.7-8.0µ wider medullary hyphae 2.8-51.71 wide and algal cells to 10-192 diem. Excipulum premium 15-30µ wide laterally, expanding to 30-65a above, hyphae 1.1-2.3µ wide. ftothecium 75-14Oµ wide, colourless stipe sometimes developed below. Thealum 85-115µ high; paraphyses a. 1.7p wide, apices to 4.6-6.6µ. wide, apices of lower 1-2 cells also pigmented form.. ing a brown epithecium. Ascii (50-70) x (18-22)v4 inner walls not uniform. Spores polarilocular, thin wailed at apices, (144-19.9)16.8 x 11.3(9.9-12.5))4 apical walls a. 0.7..1.1µ thick, lateral wails c. 1.3-1.414 porus a. 2.7-3.0pL aids, outer wall thickened at septum and septum well developed. Thalline reactions:- go C:, P. Hypothecium theoium 1+ blue. Plant caloicolous.
Variability and identlfloAtign The available data suggests that Rinodina biatherfii is a rather constant speoies in its morphology and anatomy. The epotheoia of some specimens era, however, immersed in the substratum and this oondition has been described as the var. immorsq. This occurs in may other species of caloioolous lichens and it is now thought that the rook substratum is dissolved away by the lichen. acids so that the apothecia sink I
9 2 .
Rinodlna . . • 1 Isischoffit (Trevls ) S Mass
140: 4.1,. IN MILES li .0---- '
,1 •
‘...
II •
•
wY ••
•
• 0 1
d • fa 9 . .
BOTANICAL SOCIETY OF THE BRITISH ISLES RIISTRIBUT OH MAPS SCHEME
Figure 9. Distribution of Rirrelira bischoffil in the British Toles. 91. into pits. This character would therefore seem to be more a description of the rook type than of the lichen. This form of the species has often been misnamed Buellia alocisoides but the latter species has now been shown to be synonymous with Rilledin& saline. R. bischoffii should not be oonfused with R. eeauatula, a rarer species, which differs in spore size, shape and morphology. At maturity the spores OR. aeouetula do not possess any visible lumina.
Taxonomic remarks The darkening of the apotheeial margin which occurs in this species is not due to cortical pigmentation as in other species such as Rinodina oxydata. In R. bischoffil it is due partly to the thalline margin being gradually excluded by the disc but more particularly by the extensive lateral growth of the excipulum proprium at its periphery.
26) Rinodin& oalearea Ara. in Verh. cool.-bot. Ges. Wien, 29: 362 (1879). Rinodina oaesiel3 var. calcarea Repp ex Arm., Flora, Regensburg, 1.3: 69 (1860).
Autheltio material Syntype:- Arn. 161 (K) - Rinodina calcarea, Pappenhcim, Arnold, 1860. 92. pescrintimq
Thallus thick, light or dark grey, areolate. Areolae discrete,. verruciform, to 0.90mm. wide. Thallus indeterminate without perceptible hypothallus. Apotheoia sessile, not frequent ar contiguous, to a. Disc black, plane becoming convex; margin conoolourous with thallus, a. 0.07mm. wide, entire and persistent. Exoipulum thallinum a. l05µ wide; cortex cellular, c. 20m wide and medulla o. 85m. Peripheral hyphae a. 5.7p wide, medioury hyphae o. 5.7p and algal cells to 0. 2311 diam• Exoipulum proprium o. 15p wide laterally, expanding to 0. 25m above, hyphae o. 1.7 It wide. Hypotheatunt a. 1508 deep. Thecium o. 100p high; paraphyses 0. 3.4p wide, apices to 0. 4.6p. wide, farming a brown epithecium. Asoi a. 70 x 26m4 inner walls not uniform. Spores polarilocular, walls thick and uniform, (20.5..25.1)22.7 x 12.8(11.4-14.2)pl apical walls o. 3.4 thick, lateral malls a. 2.0p, with a thick outer wall representing half these measurements, porus rapidly occluded, septum well developed at maturity. Thallino reactions:- R:, C:, I:, P.. flypothecium and theoium I-. Plant oaloicolous.
VariabilitY_and identification Thorn is very little that can be said about either 93. of these topics as only the type material has been examined but as the thallus morphology and spore characters are so unique VOW it would be difficult to confuse this species with any other.
Papno9ic remarks calcarea has been placed in the R. sophodes group of species because the spore lumina are dilated and the spores would appear to be thin walled if it were not for the thick outer spore well.
27) Ring=86.42aP101% 16nge in Lich. Nev. Zemlya: 251 (1928). RILL:ekstmniarests, var.211dmat Th. Fr., K. Svenska VetenskAdadi Mandl., 2 (2): 25 (1867).
Authentic material Syntype:-. (UPS) - Rinodiina mniarese var. oaloigena, Wahlensbergbay, Malmgren, 1861.
Description
Phallus thick, dark brown, areolate. Areolae to C. 1.26mm. wide, plane. Phallus determinate or indeterminate without perceptible hypothallus. Apothecia innate, rarely contiguous, to 0.72mo. diem. Disc black, persistently plane; 94. prominent thalline margin not evident. Szatpulum thaLlinum a. 751* wide; cortex cellular, a. 15p wide, medulla a. 60p. Pariplunmahyphas to 0. 5.7µ wide, medullary hyphae a. 2.3p and algal cells to a. 15p diem. Exoipulum proprium 8. 15p wide laterally, expanding to a. 50p above, hyphae o. 1.1p wide. Hypotheoium o. 150p deep, merging with a hyaline stipe below. Theoium o. 115p high; parephyses a. 2.3p wide, apices to e. 4.6p wide, forming a brown epithecium. Asoi a. 60 x 23p, inner walls not uniform. Spores polar Jocular, walls thin and uniform (19.4-22.8)21.7 z 13.4(12.5-14.8)P, apical walls a. 1.3p thick, lateral walls a. 1.5p, pores c. 5.7µ wide when present, septum well developed. Thalline reactions not known. Hypothecium I-, thecium I+ blue. Plant oalcareous.
Variiglikr qnd identification Nothing is known of the variation within the species as only the type material has been studied. The speoies is very similar to Rinodina castanomelq and can only be distinguished from it by the truly innate apotheoia and a small difference in the size of the spores, the outer wall of which does not appear to be thickened at the septum.
Taxonomic remarks The relationship of Ririodina caleigena and R. oastanomele, 95, is oertainl,y very *lose and the possibility of uniting it with that species has been considered. The motives for making it a variety of the latter species are strong but until mare is known of the variation within the taxa it would seem wiser to maintain the species to prevent any future confusion.
28) Rinodina castanosiela Ara. in Verh. sool.-bot. Ges. Wien, II 121 (1887). - Lecanora oaetanomela Nyl., Flora, Regensburg, 01.: 99 (1886).
Buicoati Arn. 1226 (H-Nyl. P.M. 2838) - Rinodina castanomela.
Authentic material Holotype Nyl. 28966 (H) - Lei:snore oastanomela, Panneveggio, Arnold, 1884.
Description
Thallus thick, dark brown, areolate and radiate- plicate at margin. Areolae contiguous, sometimes cracked, plane or verruciform, to c. 1.26mm. wide. Thallus determinate, without perceptible hypotbellue. Apothecia.sessile, often contiguous, to 0.90-1.01mm. diem. Disc bleak, plane becoming slightly convex; margin ooncolourous with thallus, c. 0.11mm. wide, entire or somewhat crenulate, persistent. 96.
Exoipulum thallinum 105-150p wide; cortex cellular, 20-4071 wide, medulla 85..90p. Peripheral hyphae 4.6-5.7)14de, medullary hyphae 2.8.4.0p and algal cells to 14-16p diem. Excipulum proprium 15-20p wide laterally, expanding to 25-114F above, colourless or pale brown, hyphae 1.1-1.7p wide, Hypo- thecium colourless ar pale brown, 75.85p deep. Thecium a. 90p high; paraphyses c. 1.7p wide, apioes to 4'.0-4.6p wide, forming a brown epithecium. Asa (50-65) x 19p, inner walls uniform. Spores polarilocular, thin and uniformly walled, (14.2-19.9) 17.0 x 8.6(6418-10.5)p4 apical and lateral walls c. 1.0F thick, outer wall often thickened at septum, pores absent and septum well developed. Thalline reactions:- K:, C:, i:, P:. Hypotheoium I-, thecium 1+ blue. Plant siliceous.
Variability and identification As for many other species there is not enough data available to discuss variation within the species. The species bears a close morphological resemblance to Rinodina, milivina„ as do most of the brown saxicolous species, but differs in the radiate- plicate thellus margin. The structure of the spores separates the two species without difficulty.
Taxonomic ramarka The thickened outer spore wall at the septum suggests a 97. close relationship withltinodinabisohoffii and the rounded lumina and thin walls with R. milivina. The relationship with R. calcilena has been discussed under that species.
29) Rinodina controversa Naas. in Rioerch. Auton. Lich.: 16, fig. 23 (1852).
=mai Arn. 35 (10 - Rirtzdj.rui controvert*.
Authentic material Mass. 295 (E) - Rinodina controversa.
Desoriotion Phallus thick or thin, light or dark grey to ochraceous, areolate. Areolae cracked, plane, to 0.61-1.00mm. wide. Phallus determinate or not, without perceptible hypothallus. Apotheoia ! persistently innate, often contiguous, to 0.47-0.58mm. diam. Disc black, persistently plane; margin concolourous with thallus, c. 0.07 mm. wide, entire and persistent. Excipulum thallinum 65-115rt wide; cortex cellular, 10-30p wide, medulla 40-85p. Peripheral hyphae 4.6-6.8e wide, medullary bYPhee 2.84i.0velds, algal cells to 14-20ji diam. Excipulum proprium 15-20 wide laterally, expanding to 20-30et above, hyphae 98.
1.1-2.3p wide. H.ypotheoium pale brown, 65-9011 deep. Thecium 75-11511 high; peraphysea 1.7-2.8p wide, apices to 4.0-.5.70- wide, forming a brown epithecium. Asoi (40-50) x (19.42), inner walls uniform. Spores polarilocular when young, walls becoming thin and uniform, (12.0-16.0)13.0 x 8.5(7.9-10.3), apical and lateral walls less than 1.0p thick, porus o. 44611 wide When present, septum prominent and outer wall sometimes thickened at septum, characteristic pigmented band around the upper half of each lumina. Thalline reactions:- Km:, C:1, Zs, P-. Hypothecium I-, theoium I+ blue. Plant calcareous.
Variability and identification The spores of Rinpdina controversa show a good deal of variation but this can probably be equated with changes due to age. Young spores have distinct lumina of the polarilooular type but the porus is soon occluded. At this stage a characteristic, darkly pigmented band can usually be observed around the upper parts of each lumina. The pigmentation gradually becomes more uniform and the lumina swell to fill the cells so that the spores become distinctly waisted. This species has previously been confused with &twain& crustulata a species with narrowly ellipsoid spores which lack any lumina. The spores of the species are similar in structure to R. bisohoffii but much smaller. R. controversa differs from both these species in its light brown hypothecium. 5.
Taxonomic remarks Although the spores of Rinodina controversa appear to lack a porus the shape of the spores indicates that the species has a. close relationship with R. bisdhoffil. The bands of pigmentation found on the spores of these species are of great interest. In the latter species the band is found round the septum and is due to a thickening at the outer cell wall. In R. oontroverse; however, the bands appear to be due to a greater deposit of pigment than to any thickening of the cell wall.
30) Rinodina milivina Th. Pr. in Nova Acta Soo. Sci. Upsal., ser. 3, 224(1861). Parmelia milivina 77ahlenb. ex itch., Method. Lich. Suppl.: 34 (1803).
Exsicoati Ansi Lang. 45 (K) Rinodina atro-oinerea; Arn. 830 (K) - flinodina solibodes; Hepp 208 (K) - Psora caesiella; Loika Hung. 33 (K) Lecanora milivina; Scheer. 608 (K) - Parmelia obsoura Var.caesielli.
Authentic material Ash. B.M. 425 (BM) - Lecanora milivina.
Description
Thallus thick, brown and areolate. Areolae 1.00-2.23m. 100. wide and minutely verruoiform. Thallus determinate or indeterminate, on a dark, entire hypothallus. Apothecia sub-innate becoming t sessile, frequent and usually contiguous, to 0.58-0.86mm..diam. Disc black, persistently plane; margin concolourous with thallus, 0.07-0.11mm. wide, entire and persistent. Excipulum thallinum 65.100p wide; cortex absent or cellular, to a. 15 wide, medulla 50-90p wide. Peripheral hyphae 4.6-6.8p wide, medullary hyphae 2.8.04.6p wide and algal cells to 9-18p diam. Excipulum proprium o. 15p wide laterally expanding to 25-30p above, hyphae o. 1.1p wide. Dypotheoium 100.140p deep. Thecium 75-1101 high; paraphyses o. 1.7p wide, apices to 4.0-5.1p vide, forming a brown epithecium. Asoi (50-60) x (19-20)p, inner walls not uniform. Spores polaril- ocular becoming thin walled, (12.1-26.8)20.5 x 11.4(7.7-12.5)p, apical walls 0. 1.5-2.2,1 thick, lateral walla c. 1.5-2.0p, porus c. 2.0-4.4p wide when present and septum well developed. - Thalline reactions:- K-, C-, I-, P-. Hypotheoium and theoium I+ blue. Plant siliceous.
Variability and identification The external morphology of Rinodina milivina is relatively constant being typified by the areolate and minutely verruciform thallus with donsley contiguous apothecia. 101.
The spores, however, show a very great range of sire although their structure is quite constant. The species may be separated from the other brown, saxioolous species by the thin walled polarilooular spores. Rinodlna oastanamela which has a similar spore structure may be distinguished by the banded pigmentation of the spores.
The spore type indicates that abodina millvi has a close relationship with R. Apphodes but differs in lacking a columnar lower cortex.
31) Rinoanzi colyscorg Th. Fr. in Nova Acts Soo. Soi. Upsal., ser. 3, .1: 26 (1661).
Rxsiooati Ansi It. 221 (X) Rinodina polvscara; Ansi Lang. 304b (K) -Rinodina horisa var. orbioularis; Ansi Van. 43 (K) Rinodina sonhodes var. oaerulesoena; Arn. 1617 (K) - Rinoding colgscorf; Hepp 78 (K) Psora sonhodes var. caerulgoomp; Krypt. Sind. 662 (K) Mass. 237 (K) -Rinodina sophodea; Rabenh. 755 (K) Rinodina sochodes var. ooerulesoens; Scheer. 314. (K) - Parmelia sclphodes; Th. Fr. 59 (K) Trev. 102.
21 (K) -Herengeria nolysnora.
Authentic material /sotype:- Hepp 77 (UPS) -Poor& sophodes, 78 (UPS) -psorq ,sophodes, var. caeruleecens.
Description
Thallus thin or evanescent, light grey, continuous or rimose, plane. Thallus indeterminate, without perceptible hypothalluss Apothecia sessile, frequent but rarely contiguous, to 0.25-0.54m diam. Disc brown, sometimes becoming black, rapidly becoming convex; margin concolourous with thallus, 0. 0.01 n. wide, entire becoming excluded. Excipulum thallirum 25-65p wide, cortex lacking. Peripheral hythae not pigmented, to 2.8-5.1 wide, medullary hyphae 2.34.0u and algal cells to 10-23p diem. Excipultan proprium 10-15F wide laterally, expanding to 15-40p wide above, hyphae 1.1-1.771 vide. Hytothecium 50-90p deep. Thecimm 70-100p high; paraphyses 1.1.1.7e wide, apices to 3.4,4.6p, forming a light brown epithecium. Asci (50-65) x (17-19)p, inner walls uniform. Snores pclarilocular, 8- 16 in aims, thin walled, (11.4-19.8)16.3 x 7.4(6.641./014 apical walls a. 2.2p thick, lateral walls c. 1.9r and porua a. 1.7p wide when present, septum well developed. Thallino reactions:- 4 feint yellow or I:, gypothecium I-, thecium 1+ blue. Plant corticolous. 103. Variability and identification
• Rinodina Polysvora appears to be a relatively constant species characterised by the small, brawn-and convex apothecia which may, hew... ever become darker and sometimes black with age. The species may be readily separated from the other species having similar apothecial characters by the small spores which Ere up to sixteen in the accuse i'maromic remarks
The thin walled, polarilocular spores may indicate that the species has sone relationship with Rinodina eorhodes although the similarities go no further. The number of spores in the ascus is of interest but should not be given any treater taxonomic status than the presence of 3-septate spores in some other - inodira snccies. 32) Rinodira oyrina Arr. in Flcra, Regensburg, 11: 196 (1881). Lichen uyHrus Achs, Lichenogr. Suec. Prodrom.: 52 (1798). Pscra horizavar orbiculeris f. rhododendri Repp, Flecht. Furop., no. 884. (1867). Exsiccati
Ansi Ven. 43 (I1) Rinadina souhcdes var. caorulescerm Arm 109 (K) -2412auj_z_galoi f.jilododeuit1;, 994N - Rinodina pvrina; Arm Mon. 25y 429, 4.302 469 (K) ••• Rinodina mane; Hopp 77 (X) - Psora soyhcdea, 8E4 (X) - Psora horiza v r. orbicularis f. rhodendri; 104.
Lich. Fenn. 443 (F) RInodina pyrina.
Authentic mate__rial
Ach. B.M. 410 (IN) - Rinodina soplgodes var. pvrina.
Description
Thallus thin or evanescent, light grey, continuous or rimose and plane. Thallus indeterminate, without perceptible hyothallus. Apothecia eesrile, frequent and usually contiguous, to 0.36-0.50mm. diem. Disc dark brawn or black, plane becoming convex; margin coricolourous with thallus, 0.047-0.071rn. wide, entire and usually pertiistent. Exciguium thallinum 30-85p wide, cortex lacking. Peripheral hyphae pig -rented or not, tc 3.4-6.3g vide, medullary hyphae 2.8-4.6p wide and algal cells to 8-24p diem. Excipulum proprium 10-15p wide lateral3y4
expanding to 15-30p above, hyphae 1.1-2.3p wide. Rypothecium 25-60p deep. Thecium 70-90p high; paraphysts 1.1-2.3p vide, apices to 4.0-6.3p, forming a brevn epithecium. Asci (45-65).x (13-19)p, inner walls uniform. Speres polariloculer, thin walled, (9.7-17.7) 13.6 x 6.3(5.1-8.5)14 apical walls a. 2.0-2.5p thick, lateral walls c. 0.9-1.44 porus c. lap vide when present at first and septum well developed. Thalllne reactions: - X:, C:, T:, P. Hypothecium and theOium I+ blue. Plant eerticolous and lipnioolous.
Variability and identification
Rinodina rvrina appears to be a rather variable species part- icularly in its internal anatomy. Morphologically the species is similar in appearance to a diminutive R. exigua although the apothecia 105. are probably more uniformly convex than in the latter species. Anatom- ically the spores are extremely variable both in size and structure and fall between those of R. raysnora and R. exigua. Rinedina wrina may be readily distinguished from R. polv9pora by the dark brown, contiguous apothecia and eight spored ascus. Separation from R. exicua is more difficult and can only be carried out by careful examination of the spore size and structure. The spores of R. exioaa are thick walled and unequally thickened Mile those of R. Invrina are 'chin walled tand equally thickened.
Taxonomic remarks
The spore type of Rirodina pyrina indicates that it is closely related to P, norhodes, fl, Polvsopora and other specien 7.-ith thin walled, polariloeular spores. In other resnects the species is close to R. witzmit and uas in fact placed in this species by Th. Fries (1871).
33) Rinodina sophodes Bass. in Ricerch. Acton. Lich.: 11. (1852). Lichen sophodes Ach., Lichenogs. Suec. . Proarcm. : 67 (1798).
F.xsiccati
Anti Long. 304.a, b (K) Rirodina horiza var. crbicularis; Arr. Mon. 24 Rirodino nonhodes; T'xypt. Vind. 2288(0 - Rinodina sophodes; Lich. Fenn. 759,760 (K) Rinodira sonhodes var nonnalis laudd 109 (BM, E) Rinodina exigua var. horiza; Rdbonhe 508 (x) - 106.
Rinodins albana; Scheer 314 (K) - Parmelia sophodes; Stenh. 132 (x) - Parmelis sophodes.
Authentic material
itch. B.n. no. 408 (J3) Leoanora saohodes.
Description
Thallus thick or thin, light grey to brown, rimose, plane or somewhat verruciform. Thallus determinate, otter with thin, entire, dark hypothallus. Apothecia 5db-innate, sometimes becoming sassile, frequent ard usually cortirucus, to 0.43-1.15mm. diem. Disc dark brown becoming black, persistently plare; margin mneelourens with thallus,
0.07-0.14mm. wide, entire and persistent.
"Excipulum thallinum 75-130p ride; cortex cellular laterallY,
15-20p wide, columnar below and expanding to 25-50p, medulla 65-115p wide. Peripheral hyphae not pigmented, 3.4,.5.70 viuc, redullarY hyphae 3.44.3p and algal cells to 13-19p diem. Excipulum proprium 10-20p wide laterally, expanding to 20-30p above, hyphae 1.1-1.7p wide.
Hypothecium 60-130p deep. Thecium 85-130p high; pararhyses 1.7-2.3p wide, apices to 2.8-5.7p, forming a brown epithecium. Asci (50-70) x (15-20)p, inner walls uniform. Spores polarilocular, thin walled, (13.2-19.8)15.7 x 7.9(6.6-9.1)p, apical c. 1.1-2.0p thick, lateral walls c. 1.2-1.7p, porus c. 1.4p wide but r-uickly excluded, septum well developed.
107.
Thalline reactions:- X:, 0::, T:t T r. Trypothecium and thecium /4. blue. Plant corticolous.
Variability and kleqtification
Rinedina seph9des is a very constant species typified by the thin walled, polarilocular spores and by the columnar apothecial cortex. The latter character separates the species from others such as R, rvrina and R. rolvsommilltdch have a similar snore structure. The spore shape and size, and the number of spores in the ascus readily separate these species in other respects. There hcs been much confusion between this snecies and Rind fr4ituas particularly in the British Isles. R. exirua is easily dis- tinguished from R. eaphodesk by its thick wailed spores and very often by its habitat. The former is usually found on rough bark cr dec- orticate wood while the latter grows on the twigs of such smooth barked trees as Fraxirms. The distribution of this species has been mapped with Rinodina oolobina. Taxonomic remarks
Rirodina milivina with its thin walled, polarilocular spores, cub- irrate spothecia and brown thallus shows a very close relationship with the corticolous R. soPhodea. The spore shape of these two species differs from the narrowly ellipsoid spores of R. nvrina and R. rob,- spore which, althcugh they have a similar spore structure, indicates a less close relationship.
108.
TIT Rinedina Sect. HischeblestialIalme in F. Svenska VetenskAcad. Mandl., 28: 20 (1902). Mischoblastia ors., Rieerch. Auto!). Lich., 40 (1852); Rinodina Sect. Paehvsnoriallalme, Y. Svenska VetenskAcad. Handl., 28: 18 (1902) pro parte; Rirodina Sect. Conradia Maine, E. Svenska VetenskAcad. Handl., 2k 18 (1902).
Description
Apothecial cortex colourless or rarely pigmentedv hypotheeium colourless, spores mischoblastiomorph or with rounded lumina and thick uniform balls, septum poorly developed.
34) Rirodina arenaria Th. Fr. in Lichenogr. Scandin., 1: 197 (1871).
Psera atrocinerea var. macrosnora f. arenaria Hopp, Flecht. Euron., ro. 646 (1860). - Rinedina caesiella var. erenaria .Arn., Flora, Regensburg, .41: 69 (1860). - Rirodina arenaria var. cana Arm, Verh. tool. - bot. Gee. A.en, /2.: 269 (1872)
Exsiccati
Arne 494 (s) — Rinedina areraria var. earn. HePP 646 ( — Psera atrooimerea ver. macrosncra f. arenaria.
Authentic material
Isotype:- Hopp 646 (r) Psora atrocinerea var. macresnora f. arenaria. 109.
Descriptitop
Thallus thin, light grey to ochramous, areolate. Areolae to 0.32-0.54ma. wide, plane or = verruoiform sometimes cracked. Thallus determinate or indeterminate, sometimes limited by a thin, entire and dark hypothallue. Apothecia innate or sessile, contiguous or not, to 0.430-0.50mm. diem. Disc black, = persistently plane; margin con- colourous with thallus, 0.04-0.11mm. wide, entire but becoming excluded. Exeipulum thallinum 4C-49µ wide; cortex absent or cellular, to wide, medulla 30-40p wide.' Peripheral hyphae not pigmerted, 2.8-4.0p wide, medullary hyphae 2.8-3.4p and algal cells to 13-14a diem. Excipulum proprium 10-15p wide lat-rally, expanding to 30-40p wide above, hyphae 1.1-2.3p wide. Rypothccium 60-100p deep. Thecium 75-110p high; raraphyses 1.1-1.7p wide, apieen to 3.4,-3.5p wide, fornr. ing a red-brown epitheciu" Aeci (65-70) x (19-20)p, inner walls uniform. Sncres mischoblastiomorph often becoming thick and uhiformly walled, (17.1-23.4)19.5 x 9.7(8.0-12.0)p, apical walls cr. 4.3-4.5p thick, lateral walls c. 1.8-2.1p, pores c. 1.8-9.6p wide when present and septum not well developed. Thalline reactions:- F.-, I-, I-. irypothecium I-, thecium /+ blue. Plant siliceous.
Variability and identificatior
Little can be said about the variability of the species except that the thin, somewhat scurfy thallus and innate apothecia of the 110. specimens previously named Rirodira cana represent a depauperate state. The generally smaller size of the spores and other internal characters are a frequent result of such a condition. The species might be min.- taken for R. atrocirerea at first sight but the lack of any thalline reactions and the broadly ellipsoid, non-mischoblastiomorph spores of that species make it readily distinguishable.
Taxonomic reinarks
In spite of the close resemblance of this species to Rirodina atrocirerea, noted above, the spore morphology indicates that it is more closely related to R. discolor. It differs from the latter species in lacking any cortical pigmentation and in not having a dark brown epithecium although it seems adviseable not to attach too much taxonomic importance to these characters.
35) Rinodina dissimilis Ansi in Comment. Soo. Crittogemolog. Ital.., 2 (1): 9 (1864.).
Authentic material /sotypes- Anzi Lang. 322 (7) - Rirodina dissimilis, Boerio - prow. Sondriensis.
Description
Thallus thin or evanescert, light grey, continuous or cranked, or of discrete verrucae, plane. Phallus indeterminate, without percept- ible Iwthallus. Apothecia sessile, frequent but not contiguous, to c. 0.43=6 diem. Disc dark brown, convex or often half-globose; margin concolourous with thallus, c. 0.04mm. vide,- entire but rapidly becoming excluded. Excipulum thallinum c. 50p wide; cortex cellular, 0. 10p wide, medulla a. 40p. Peripheral hyphae not pigmented, c. 440p wide, medul- lary hyphae c. 2.6p and algal cells to el. 9pdiam. Excipulum proprium c. 15p wide laterally, expanding to c. 25p above, hyphae c. 1.7p wide. Hypcthecium liEht brown, c. 50p deep. Thecium c. 95p high; para physes c. 1.1p wide, apices to c. 4.0p, forming a brown epithccium. Asoi c. 60 x 19p, inner walls uniform. Spores mischoblastiomorph, walls becoming equally thickened, (15.4-22.2)18.3 x 8.3(6.3-9.1)N, apical walls c. 3.5p thick, lateral walls c. 1.6p, pores c. 1.4p wide when present and septum developed at maturity. Thalline reactions:- X-, 0-, I-, Hypothecium I-, thecium /+ blue. Plant siliceous.
Variability and identification
Without further material it is rot possible to give any indication of the variability of the species. Separation of the species from other species growing on siliceous rocks is relatively easy. The characters of the brown, half-globose disc with excluded margin taken to-gether typify the species. The species might conceivably be con- fused with Rirodina saline or R. arenaria. The spore size and structure would separate it from the former and the apothecial morphology from the latter. 112. jaX0/1011110 remark.
The possibility of uniting this species with14_20INEAghas been considered. The various differences in character referred to above, although none of them hold much taxonomic weight individually, taken in total appear to indicate that the species should remain separate. Examination of further material, however, might indicate that the species should be regarded as a variety of,R. arenarists
36) Rinodina oxvdateMass. in Genesi". Lich.; 19 (1854). Berengeria oxydata Travis., Spighe e Paglie; 6 (1853). - Lecidea, discolor Hepp, Fleoht. Europ., no. 319 (1857). - Buellia discolor Ansi, Catal. Lich. Sondr.; 87 (1860). - Lecanora oontribuens Nyl., Flora, Regensburg, 2.1.: 198 (1873). - Lecanora griseofusca Nyl., Flora, Regens- burg, 12: 360 (1875). Lecanora confragosa var.,lecidotropa, Nyl., Flora, Regensburg, 60: 232 (1877). - Lecanora intuta Flora, Regensburg, 68: 296 (1885). Rinodina subarenariq A.L. Sm., Monogr. Brit. Lich., 1: 255 (1918). alL22.1111 Ansi Lang. 193 (K) - Buellia discolor var. candida.
Authentio Neterial
Isotype:- Trev. 22 (K) Berengeria oxydata, Mt. Closo, S. Miohiele. 113.
Desert :11m
Thallus thin, more rarely thick or evanescent, light grey to oobraceous, rimose or sometimes of discrete of contiguous verrucae, plane. Thallus determinate or indeterminate, sometimes limited by a dark, entire hypothallus. Apothecia sessile, frequent and sometimes contiguous, to 0.29-0. 4.7m. diem. Disc dark brown becoming black, persistently plane; margin concolourous with disc, 0.04•0.11mm. wide, entire and persistent. Exoipulum thallinum 3O-75p Wide; cortex cellular, outer pig- mented part 10-45p wide, inner colourless portion 10-20p and medulla 1044p wide, Peripheral hyphae obscured or to 440-6.3p wide, medullary hyphae 2.3-4.011 and algal cells 9-17p diem. Excipulum proprium less than 10-15p wide, expanded to less than 10-25p wide above, hyphae 1.1-2.8p wide. Hypothecium 2O-90p deep. Thecium 70-115p high; pare- physes 1.1-2.8rvide, apices to 3.4-5.7p, forming a brown epithecium. Asci (60-90) x (18-26)p, inner walls uniform or not. Spores mischo- blastiomorph or with thick, uniform walls, (16.5-25.3)20.7 x 11.5 (8.5-16.5)p, apical walls c. 3.7-6.0p thick, lateral calls 0. 1.6-3,214 porus 1.5-3.2p wide and septum not well developed. Thalline reactions:- yellow (often sordid), C:, /:, HYPothecium I-, theoium /+ faint blue. Plant siliceous. 114.
Variability and identification
The external morphology of Rinodina ox data is remarkably constant although the thallus may be well or poorly developed as in many other species. The spores are of the mischoblastiomorph type but the lumina often expand so that the spore walls become more or less equally thickened. This has been noted for many other species with mischoblastiomorph spores and seems to occur with the ageing of the apothecium. The amount of pigment deposited in the apothecial cortex, as seen in section, varies between quite broad limits and in every few instances it is not strictly correct to describe the external aspect of the margin as being concolourous with the disc. However, even in these few cases the exoipulum thallinum is much darker than the thallus and confusion would not occur with other species completely lacking pigmentation. It is evident, as the list of synonyms of the species shows, that there has been a great deal of confusion regarding this species in the past. There is no doubt that much of this confusion has arisen from the fact that the external appearance of the apothecia is lecideine and many of the species were therefore originally des- cribed as belonging to the genus Lecidea. Knowing the true nature of the apothecia, it is this character that makes for the easy identification of the species as only one other saxicolous species, Rinodina occulta, has a dark coloured apothecial margin. R. oxvdata is easily distinguished from this species by its large, mischoblast- iomorph spores. 4 /
Rinodino
oxydolo (Travis.) o 8 B
P
.a o. 4- ' ' ..-.....4-1 ...,!. . 1i 3 1 In ii 1 dB—
Mali /1/6.6. - ) • -11' ,
.... r if.
3
•
_ 2
• _ lk
ta o ....--j
.... . /::....: 2 6
BOTANICAL SOCIETY OF THE BRITISH ISLES DISTRIBUTION MAPS SCHEME
Figure 10. i:istributior of R1toaVie mifinto it the t-itish Isles. 116.
Taxonomic remarks
The spares of Rinodina oxrdatjs are a fine example of the mischoblastiomorPh type and there can therefore be no doubt as to which section of the genus the species belongs. The type specimen of R. ozvdsta differs rather markedly from the other synonymous species in having a darkly ochraceous thallus. It is now known that this pigmentation is caused by iron compounds in the substratum and cannot therefore be of any taxonomic significance. The pigmented cortex of the excipulum thallinum is of greater interest. As the thalline cortex is not also pigmented it is apparent that the margin of the apotheoium closely parallels some of the more advanced species of the gerusrvxine. However, this character is also present in the sections of the genus with polarilocular and placodiomorph spores, c.f. ItimateLossultaand R. biloculata respect- ively, and although it is of great importance from the developmental point of view it is not thought that it Should be givenumdte taxonomic weight. In a similar way Imshaugh (1957) rejected the use of characters of the excipulum thallinum in favour of spore characters for erecting subsections within the genus Pvxine. If spore morphology is to be taken as the most important character in the taxonomy of the species then it would appear that Rirodine arenaris and R. dissimilis bear the closest relationship to R. exydata. 117.
37) Rinodina teichophila Arm in Flora, Regensburg, l16: 329 (1863). Lecanora teichophilaNyl., Flora, Regensburg, hd5 78 (1863). - Lecanora suberumpens ryl., Lich. Pyren. Orient.: 7 (1891).
Exejecati
Arn. Mon. 155 - Rinodi6a calcares var. obscurata; Johns. 197 (Iv, En) - Lecanora milivina; Larb. l'erb. 25 I: ) - LecarArt 4Aivina; Mudd 108 Mt E) - Rinodint exisua var. metabolida.
Authentic material
/soty-pe:- Mudd. 108 (73M, 1:)7 - Rinodina exima var. metabolica, Ayton, Cleveland.
Description
Thallus thick or thin, dark grey or rarely light grey to ochraoeous, rimose, plane or usually granular. Thallus determinate or irdetermiratc, without perceptible hypothnllus. Apothecia sub-irrate, frequent and often contiguous, to 0.6e-1.26mm. diam. Disc dirk brown becoming black, persistently plane; margin con- colourous with thallus, 0.07-0.122m6 wide, entire and persistent. 118.
Exoipulurn thallinum 60-95p wide; cortex cellular when present, to c. 25p wide, medulla 40-85p wide. Peripheral hyphae pig- mented or not, 440-6.3p wide, medullary hyphae 3.4-5.1p wide and algal cells to 10-10 diam. Expipulum proprium 15-20p wide, expardirg to 20-50p above, hyphae 1.1-1.711 wide. Rypothecium 55-100p deep. Thecium 80-145p high; raraPhyses 1.1-14,7µ wide, apices to 3.4,-5.7p, forming a brown epithecium. Asci (75-90) x (20-32)p, inner walls uniform. Spores mischoblastiomorph or with thick, uniform walls, dilated at septum, (20.5-31.9)26.3 x 14.6(11,4-19.8)14 apical walls c. 4.7-6.11p thick, lateral walls c. porus 0. 2.6-4.1p vide and septum very pocrior develored. Thalline reactions:- K:, 0-, I-, P-. Rypothecium I-, thecium 1+ blue. Plant siliceous or rarely corticolcus.
Variability and identification
Rinodina teichophila is a relatively constant species well chvracterised by its large, mischoblastiomorph spores, sub-innate apothecia and granular thallus. Depaurerate specimens that have been subject to adverse environmental conditions may hove light grey or some- what ochraceous thalli which may also lack grcrules. This species is one of the very fcw saxicolous species that has also been found growing on bark. •
2
o
Rinodinn
teichophila (Nyl.)Arn. g 8 wl c a
b A 7 ...., ---2-"'? MIL":
) AIP
....
• •
2 _..... 2
Q. 1Z3
BOTANICAL SOCIETY OF THE BRITISH ISLES DISTRIBUTION MAPS SCHEME
Figure 11. .'stributil..1- in the british Isles. 120.
Corticolous specimens have been misidentified as Rinodips
sophodes but 11: to e really bears ro resemblance to this species.
In the British Isles the species has been widely confused with
R. mi1ivina which is now known to absent, or at least not recorded,
from these islands. R. teichophila has also been confused with R. confrasosa probably because of the similarity of scre size but it
is easily distinguished from the latter species by the !TIT= thallus reacticr and the mischoblastiomornh spores.
TRXDPOMIC marks
Rinodina teiehonhila Shows a close relationship with, R. arenaria and R. discolpr in spore type although the pigmented epothecial margin of the latter species makes this relationship less close.
There can be ro doubt that Rirodina suberumnens represents ro more than a depauperate form of the present species as the anatomical
features are identical.
38) linodina atrocinerea Koerb.
in Syst. Lich. German.: 125 (1855).
Mich r. etrocirereus DicIts., Paso. Plant. Cryptog. Brit.,
14, fig. 9 (1793). - Rinodipa alba Metzl. ex ;r2a., Flora, Regensburg, LI: 35 (1872). - Lecanora plumbella ryl" Bol. Soc. Broterian., 6: 227 (1888). 121.
Exsiccr.ti
Ansi Lang. 321 (K) Rinodina caesiella; Arai Von. 44 (K) - Rinodina le aronipa var. traohvtica; Arm 493 (K) - 2knoaina . trachytica; Johns. 235 (MO - Lecanora taichorhila; Larb. Cams. 28 (lc) Lecanorq sorhodea var. strocinerea; Larb. Lich. 302. (K) -
Lectrora (3.trooinerea; Leight. 146 (X, E) - sonhodea; Lojka Hung. 124 (K.) Lecanore. atrooinerea.
Authentic material
Leototype:- Herb. 4ithering no. 59 (Y) Lecanora atroeinerea,
DescrAutim
Thallus thick or thin, light grey to ochmceous, arcolate, areolae plane, 0.43-1.26=6 wide, discrete on a black hypothallus or contiuuous, sometimes rimose and verruciform.. Phallus determinate, limited by a black entire hypothallus. 1pothecia sessile, to 0.54- 0.83mm. diem., somotimcs contiguous. Disc blach, 1-erzistently plane or becomirg convex; margin corcolourous with thallus, 0.07-0.11m• wide, entire and persistent.
Exeipulum thallinum 50-1241 wide; cortex 15-25i wide laterally, structure occluded, sometimes expendirg to c. 30v below, 122. medulla 40-100p wide. Medullary hyphae 2.8-5.1p wide, algal cells to 9-151 diam. Excipulum proprium 10*2.0p wide laterally, expanding to 15.4.0p above, hyphae 1.1-1.7p wide. Hypothecium 45-14Op deep. Thecium 75-140p high; paraphyses 1.1-2.3p wide, apices to 2.3-4.714 forming a brown or red -brown epithecium. Asci (50-90) x (19-23)p, timer walls uniform. Scores uniformly thick walled, (16.C-21.1)18.1 x 11.2(8.5-13.7)14 apical walls c. 2.5-3.6p thick lateral walls c. 2.4- 3.4p, porus usually absent sometimes c. 2.7p wide, sot tum rot prominent. Pycridia often present, innate nt first. Coniaa bacilliform, c. (4-6) x lu.
.1111 Thalline reactions:- X+, yellow cr C.t or + red', I-, P-. Hypothecium I-, thecium 1+ blue. Plant saxicolous.
Variability And identification
RiroaIra. plutlbella characterised by its small, aspersed areolae, renting on a black hynothallus wets found to fall within the normal range of variation of R. atroolnerea. it is proboble that Pylander via strongly influenced by the ::011 reEative reaction cf the thallun -.7hen he erected the species but this reactirr is now known to be variable in ma=y species of Rinodiva. rather extreme =maple of th- colour variation of the species is represented by :Z. alba. Thin form has a markedly onhraceoua thallus but there is evidence that 12.3*
-u,f
Rinodino 9 62 otrocinerea iDicks.)
Koerb.
may,• a N
7
,....1-'7 pm66:
6.1.1
\\! • ..... \ II-
"411" •
A. Id s9 0— —4-
6
BOTANICAL SOCIETY OF THE BRITISH ISLES DISTRIBUTION MAPS SCHEME
Wiyure 12. tctributtcn ofilrIctdirl.0 utroeinerzolk in the British isles. 124.
this colour develops, at least in part, in the herbarium and it cannot be regarded as a good Character for speciation. Rinodina atrocinerea resembles R. arlpariq in some respects but may be distirguiShed from that species by its broadly ellipsoid' and rather shorter spores.
Taxonomic) remarks
As with many species that Dickson erected it has not proved possible to locate a type specimen. An authentic specimen from herb. Withering (no. 59) in Kew, labelled Lecanora atrocinereus, has therefore been selected as loototype.
39) Rinodina cintrana Samp. in Broteria, ser. bat., 20: 161 (1922). Lecarora rruinella var. cintrana Samp., Broteria, ser. bot., .7. 11 75 (1916).
Authentic material
Syntype:- (IC) Rinodina cintrana, Cintra, Sampaio, 1916.
Description
Thallus thick or thin, light grey, continuous or cracked, plane 125.
or more usually granular. Thallus determinate, limited by a thin, entire, black hypothallus. Apothecia sessile, sometimes contiguous, to 0.61-1.01mm diem. Disc black, persistently plane and sometimes pruinose; margin concolourous with thallus, 0.07-0.11m. vide, granular like thallus, entire and persistent. Esoipulum thallinum 60-70p wide; cortex, 10-15p wide, structure occluded, medulla 45-60p. Medullary hyphae 2.34+.0p vide and algal cells to 11-15p diam. Exoipulum prorrium c. 15p vide laterally, expanding to 15-25p above, hyphae c. 1.1p wide. Hypotheoium 90-140p deep. Thecium 75-130p high; paraphyses c. 1.1p wide, apices to 3.4- 4.6p wide, forming a brown epithecium. \sci (60-70) x (14-18)p, inner walls uniform. Spores mischoblastiomerrh or later uniformly thick walled, (14.8-23.4)18.5 x 9.3(8.0-10.8)p, apical walls c. 3.7-4.0i thick, lateral walls 1.6-1.5p, pores c. 1.3-1.7p wide when present and septum absent, spores pointed at apices or widened at centre. Thalline reactions:- P- yellow, C-, /-, P-. Hypothecium I-, thecium/+ blue. Plant corticolous.
Variability and identification
The morphology of the thallus is remarkably constant in this species although the spore structure and size is probably more variable than this would lead one to expect. The species might possibly be mistaken 126.
for Rinodina roboris but usually has a thinner, wore regular thallus. The spores of R. cintrana can be separated from those of R. roboris by their pointed apices, thickened centres and more narrowly ellipsoid Shape.
Taxonomic remarkk
Rinodina cintrana is undoubtedly a close relation of R. roboris as the spore morpholokk shows. Yegnuszon (1947) states that the species is synonymous 10,1.th R. truinella although he saw no authentic specimen of the latter species. Until such time as the type of R. Druinella has been viewed the name R. cintrana should be retained.
40) Rinodina confinis Samp.
in Bolet. Socied. Broterian., ser. 2,p 19 (1924)
Authentic material
Syntype:— (TO) — Rinodina confinis, Povoa de Lanhoso,
Sampaio, 1919.
Description
Phallus thih, light grey to ochraceourt, continuous or cracked, plane or often granular and sometimes t subsquamulose. Thallus indeterminate, without perceptible hypothallus. Apothecia sessile, 127.
often contiguous, to 0.50-0.79mm. diem. Disc dark brown or rarely black, plane becoming slightly convex; margin conoolourous with thallus,
0.07-0.11mm. wide, entire and persistent.
Excipulum thallinum 70-1141 wide; cortex subcellular or absent, to c. 15p wide, medulla 55-110p. Periphertl hyphae not pig- mented, 3.4-4,4 wide, medullary hyphae 2.8-4.6p cm& algal cells to 10-135 diem. with prominent pyrenoids. Fxoipulum proprium c. 10p wide laterally, expanding to 20-40p above, hyphae c. 1.1p wide. HyTothecium 45-90p deep. Thecium 115-140p high; paraphyses 1.7-2.3p wide, apices to 2.8-4.6p, forming a light red-brown epithecium. Asci (85-90) x (26-32)p, inner walla uniform. Spores thick, uniformly walled,
(20.5.35.3)27.0 x 14.0(12.0-18.8)p, apical walls c. 4.9-5.4p thick, lateral walls c. 3.2-3.5p, pores hot present and seetum developing very late. Thalline reactions:- K:, C:, I:, P:. Hypothecium and thecium 1+ blue. Plant corticolous,
Variability and identification
The variability of this species is mainly aTrarent in the development of the thallus. The usual granular condition of the thallus has a tendency in some material to develop into minute squamules, here termed subsquemules. They aprear to form by the coalescence of 128.
a ruMber of Frorules in much the same way as the isidia of Rinodina isidiodes fuse to form squamules. Rinodina =finis cannot be confused with any other species. The large spores with thick, uniform walls are characteristic of only one other species, R. isidiodes$ which is distinguished by its isidiate thallue.
Taxonomic remarks The spore morthology iraicates that this nrecies is very closely related to Rirodira isidiodes ard R. corradi. This view is further sUtstartiated by the similarity of the thalli, the large spores, the relatively hith thecia and by the red-brown epithecia.
41) Ripodina ccnradi "Fverb, in Syst. Lich. German.: 123 (1853). Lecanora dinlinthiaryl., Acta Soc. Fenn., 444 (1863). - Rinodiipa intermedia Bagl., Cocment. Soc. Crittogam. Ital., 1: 315 (1863). - Rinoal.ra intermedia Gomment. Soc. Crittogam. Ital., 2: 315 (1863).
Exsiccati
Arn. 67, 1551 (K) - Rinodina corradi; Larb. Gees. 78, Larb. Herb. 263, 302H - L-carora mreniosrora; Lojka Hung. 123 (I") - Lecorora conradi; Raberh. 381 -Peore turfacea, 81.0 (lc) Rinodina conradi. 129.
DescrintiN
Thallus thin or evanescent, light grey, continuous, plane or cracked or of t discrete verrucae. Thallus indeterminate, without perceptible hypothallus. Apothecia sessile, often contiguous to 0.29.. 0.83mm. diem. Disc often dark brown, soon becoming black, plane some- times becoming convex; margin concolourous with thallus, 0.04-0.07mm. wide, entire and persistent. Exoipulum thallinum 45-90p vide; cortex absent or rarely present and then structure occluded to 15p wide, medulla 35-75µ wide. Peripheral hypherae rarely distinct 2.8-5.7p wide, medullary hyphae 2.3-3.4i and algal cells to 11-20p diem. Excipulum proprium 10-20p wide laterally, expanding to 15-50p above, hyphae 1.1-1.7p wide. Hypothecium 25-75p deep. Thecium90-13op high; paraphyses 1.7-2.3p wide, apices to 3.4-5.7p, forming a brown or more usually red brown epithocium. Asci (65-95) x (23-28)!*, inner walls uniform or not. Spores thick, uniformly walled, 1-3-septate, rarely with a longitudinal septum, (18.2-30.2)25.4 x 11.7(8.5-14.2)p, apical walls c. 2.9-3.5p thick, lateral walls c. 2.3-3.5p, porus absent and septum poorly developed. Thalinel reactions:-X :, C:, 1-, P-. Aypothecium I-, thecium /+ blue. Plant terricolous, muscicolous or lir_nicolous. 130.
Variability and identification
The variation of thallus morphology in Rinodina conredi is tot great, the thallus being either well or poorly developed. The internal anatomy is more variable, particularly in the presence or absence of a cortex and its structure when. present. Young spores may often be observed with two lumina but are rarely seen in this state at maturity. Spores with a longitudinal septum may also be rresent in some material. The species cannot be mistaken for any other on account of its predominantly 3-septate spores although some other snecies such as Rinodina isidioaes do occaisionally have a tendency to produce 3- septate spores.
Taxonomic remarks
It has not been possible to study authentic material of Rinodina intermedia whidhVagnuston (1947) separates from R. conradi by its 5-6-celled spores. However, authentic material of R. lusitarica, A synonym of R. inteziedia according to Magnusson, has been studied and no significant difference could be found between this species and R. corradi. It is therefore very probable that R. intermedia should be regarded as being synonymous with R. corradi. Type material of R. dial,Anthia showed that the species was certainly synonymous with
9
Rinodi no conrodi n1
4
NB
4
BOTANICAL SOCIETY OF THE BRITISH ISLES DISTRIBUTION NAPS SCHEME
re 13. istributior of •,4tc:s4;6 tr the 132.
R. eorradi for the same reasons as stated above. Spore septation is no longer considered as important in lichen taxonomy as previously and there can be no good reason for putting Rinodina conradi in a separate section within the genus as Malmo (1902) and Magnusson (194.7) have done. It has already been pointed out that there appears to be a close relationship between this species and R. isidiodes and R. =finis, neither of which species comonly possess 3.-septate spores.
42) Rirodina isidioidpg Cliv. in diem. Soc. Nat. Sai. Cherbourg, 2: 186 (1909). Parmelia isidicidu Born, ex Hook et Sowerb., Suppl. fngl. Botai6,1: tab. 2808 (1843).
Exsicoati Salve. 155 (M) Parmelia isidioides.
Authentic material
Isotype:- On) - Pormelia isidioidea, Corm Bychan, Salwey, 1835.
Description
Thallus made up of numerous coralloid, branched and thin, 133. cylindric isidia. Isidia light grey, sometimes admit:: to form plicate squamules. Thallus indeterminate without perceptible hypot- Apothecia sessile, frequent and sometimes ccntiguouB, to 0.90- 1.12mm. diem. Disc black, persistently plane; margin concolourous with thallus, 0.11-0.14m. wide, entire or crenulate and persistent. Excipulum thallinum 55-90p wide; cortex absent laterally or with structure occluded, t o. 25p wide, columnar below and 40-60p wide, meGulla 30-90p wide. Pigmented peripheral hyphae absent, medullary hythae 2.3-4.6p wide and algal cells with prominent pyrenoids, to 13-151 diem. Excipulum proprium 10-20p wide laterally, expanding to 25-40p above, hyphae 1.1-1.7p wide. Tixpothecium 3C-50p deep. Thecium 130-160p high; paraphyses 1.1-2.3p wide, apices to 4.0-5.7i wide, forming a red-brown epithecium. Asci (75-110) x (22-35)µ, inner walls uniform. Spores thick, uniformly walled, (20.9-31.3) 26.7 x 14.2(11.4-17.1)p, apical walls c. 4.0-5.5p thick, lateral walls c. 3.4-3.5p thick, pores c. 1.1-2.6p thick when present and septum well developed at maturity only. Thsllibe reactions:- 4 yellow, C:, I:, P. Hypothecium I-, thecium I+ blue. Plant muscicolous on bark or corticolous.
Variability and identification
Rirodira isidioides is a very constant species end would 0 .....
9 2
Rinodina o . sidioides (Bord Oliy. a nt 0
7 0 AN .
may, 6
4 r.ii .166...- ) 13,
ill .._ s.... „A..., 4,-
111111""tr
2
0 .!
C3
0
BOTANICAL SOCIETY OF THE BRITISH ISLES DISTRIBUTION MAPS SCHEME
Fiore 14,, istribution of Bie lifla uladam in the nritIsh Tales. 135. be easy to identity even in the sterile state. The light grey isidiate thallus of the species is quite characteristic and the species could not be mistaken for any other within the genus.
Taxonomic remarks
The large spores and rounded lumina indicate that Rirodina isidioides is closely related to R. conradi in spite of the fact that the latter species has 3-septate spores. R. isidioides does show some slight tendamy to form four lumina within the spores.
43) Rinodthamarittma nov. sp.
Exaiccati
Larb. Cass. 28 (r, K) - Locanora sophodea var. atrocinerea, La Coupe - Jersey, Larbalestier.
Description
Thallus thick or thin, light grey, continuous and granular, rimose omit pyre or of discrete verrucae. Thallus determinate or indeterminate, without perceptible hypothalluss Apothocia innate or more usually sessile, sometimes contiguous, to 0.72-1.26--. diem. Disc black, t persistently plane; margir. corcolcurcus 7iith thallus, 0.07-0.18=4 tide, entire bccomirg arenuiatc, rcrsistont. 136.
F.xcipulum thallirum 70•95p wide; cortex 10-30p vide, structure occluded, medulla 60.750. Expanded peripheral hyphae not ?resent, medullary 11,701110 3.44446p wide and algal cella to 9-11p diem. Excipulum proprium 10-20p wide laterally, expardirg to 13-2512 above, hyphae 1.1-1.70 wide. nypothecium 11c)-140p deep. Thecium 90.400p hilth; paraphysea c. 1.70 nide, apices to 5.1...C.0O3 formirc a red- brown epithecium. Asa (65-95) x (15-2;:4)0, inner walla uniform. Spores rarely mischoblastiomcrph„ more usually thick and uniformly walled, (16.5-22.8)18.7 x 10.7(8.5-14.2)0, apical walla o. 4.6µ thick, lateral walls c. 2.50, -perus o. 2.4' wide Whon rresent and septum rot yell developed. Thailire reactions:- I-, yellow, C-, I-, Hypethecium and thccium blue. rlant siliceous cr terricolous.
Variability Pnd identification
The variability of this rew species arrears to be limited by the substratum on which it grows. The specirens from aokholm island that have crown on decaying Armeria_rarittima have a thiec, con. time= thallus which is densely rranular. Other raterIal, more typical of the species, Eathered fro:n a s%aded cliff nearby had a
rimose thallus an,7 trarules. Thcl intern:A aratorw of thane two typo is elicst identical. For the distribution of the species in the British Isles see under Rinodina corfragosa.
tAtime 15. Thallus '1.nodint, myrittl.a. rtgrifloodon x 7. 138.
This western species has formerly been inoluded under Rinodina oonfr4gosa in British lists, of which only two speoimens are now known to have been recorded and those from Scotland. R. maritime is distinguished from this latter species by the broadly ellipsoid spores, lacking a well developed septum and by the thick, uniforal,v thickened spore walls.
Taxonomic remarks Rinodinampritisa belongs to the eiaohoblastioaorph group of species and is therefore no relation to R. confragosa with which it has formerly been confused. The spore morphology would seem to indicate that it might be related to the R.. isidioides group of species. The possibility of separating the two thallus types referred to above as separate varieties has been considered but it is thought that this should await more data on the morphology and distribution of the terricolous material.
44) Rinodina melanooarri Mull. Arg. in Flora, Regensburg, 2: 434. (1867).
Authentic material Isotype:- (x) anodize melanocaros:, nr. Geneva, Muller Argoviensis. 139. Description
Thallus thick, light grey and areolate. Areolas plane becoming verruciform, to o. 1.44mm. wide. Manus determinate or indeterminate, without perceptible hypothallus. Apothecia sub- innate, frequent and sometimes contiguous, to c. 0.43mm. diem. Disc black, plane or slightly convex; margin oonoolourous with thallus, o. 0.04mm. wide, entire but quickly becoming excluded. Exeipulum thallinum a. 80p wide, cortex absent. Peripheral hyphae to c. 6.3p wide, medullary hyphae a. 2.8p and algal cells to a. 13p diem. Excipulum proprium o. 5pt wide laterally, expanding to a. 10p wide above, hyphae o. 1.7p wide. Hypothecium pale brown, a. 140p deep. Theciun a. 90p high; paraphrase a. 2.3p wide, apices to a. 4.6p, forming a brown epithecium. Asa a. 60 x 23p, inner walls not uniform. Spores with thick, uniform walls, (154.48.8)16.8 x 9.448.5.10.op, apical walls a. 2.7p thick, lateral walls a. 2.1p but lumina becoming occluded, pores a. 1.1p wide when present and septum developing late. Thalline reaotions:- g yellow to red, C:, 12* lemon yellow. Hypotheoium and theoium 1+ blue. Plant siliceous.
Variability' and identification The variation within Rinodina melanocarea is unknown as it has only been possible to examine the type speoimen. There should be little difficulty in identifying the species as the g yellow to red 140. and 14 lemon yellow thalline reactions are unique within the genus. The sub-Pinnate apothecia, without any persistent margin also separates R. melanocarya from all other species examined except R. lecanorina and R. lurijesoens. The species may be readily distinguished from these by the thick, equally thickened walls of the spores which have no connection with the polarilocular type of spore of the latter two speoies. The Chemical reactions and general habit of R. melanocaruit are almost identical with Buellia subdtsoiformis.
Taxonomic remarks The combination of apotheoial morphology and spore type of Rinodina melanocaroa makes it an isolated species. The sub- innate apothecia strongly suggests that the species belongs in the genus Molamvpicilia if further work indicates that the genus should be re-erected.
45) Rincdina muscioola H. Naga. in Acta Hort. Gothoburg., lb 234. (1914.7).
Authentic material
Nyl. 28455 (H) - Lecanora sophadas var. Aaeviaata f. muscioola, Hollola Fennia, Lang, 1871. zusultass
Thallus thin, light or dark grey, continuous, plane to verruciform with granules developing. Thallus indeterminate, without perceptible hypothallue. Apothecia sessile, frequent and often contiguous, to c. 0.54mm. diam. Disc black, plane becoming convex or halfi-globose; margin concolourous with dice or lighter, c. 0.0. wide, entire becoming excluded. Exoipulum thallinmm o. 5011 wide; cortex cellular o. 20p wide, medulla o. 30p. Peripheral hyphae to o. 4.6p wide, medullary hyphae o. 2.8p and Algal cells to c. 1111 diem. Exoipulum proprimm o. 20p wide, laterally expanding to o. 25p above, hyphae 0. 1.7p wide. Bypath:mina a. 7511 deep. Theoium 0. 90p high; paraphyses • c. 1.7p wide, apices to a. 4.714 forming a brown epitheeium. Asei o. 60 x 19p, inner walls uniform. Spores mischoblastiomorph, walls becoming thick and uniform (17.7..22.8)19.5 x 9.2(8.0-10.3)p, apical walls 0. 3.6p thick, lateral walls a. 1.5p, parus a. 2.6p wide when present and septum not well developed. Thalline reactions:- X:, C., I:, P. Hypotheoium 1-, thecium I+ blue. Plant musoicolous.
Variabiliti .and identification Any statement about the variability of the species is precluded by the absence of specimens other than the type. The identification of Rinodina musoicola is made easy by the musoicolous 142.
habit and mischoblastiomorPh spores. Even among the corticolous and lignicolous species there are none with both misohoblastiomarph spores and convex apothecia with excluded margins.
Taxonomic remarks This species has nothing to do with the genus Buellia as
Magnusson (loo. oit.) suggests. The excipulum proprium is well developed but there is also a. well developed excipulum thallinum with algal cells. The spore type indioates some relationship with Rinodina roboris andp. cintrana but the apothooial characters referred to above are quite distinct from these species.
46) Rinodina rbboris Ara. in Flora, Regensburg, as 197 (1881). Lecanora soubpdes var. roborie Duf. ex nyl., in Crouan, Florule de Pinist.: 96 (1867).
EX8100Ati Ansi Lang. 377 (K) -Anciabia metabolica; Arn. 1654 - (inodina ramaicolv Deem. ed. 1, ser. 1, 1594 (K) - ,Lecanora perioleq var. exigua; Johns. 36 (BM, NMW) • Lecanora roborisI Larb. 260 (EM, K) - Leoanora 'collodion Rabenh. 889 (K) • Lecanora =boils; Scbaer. 569 (K) - Lecanora atra var. =Igoe; Welw. 103 (K) - Pannone Imbeds.. 14:3. Authentic material.. Syntypos. Nyl. P.Y. 2822 (11) Lenora sonhodes var. ,roboriss Brest, Crouan, 1864. pigadaktis
Thallus thick or thin, light to greenmmy, continuous or more usually rims., plane or sore usually granular ant verruciforn. Thallue indeterminate, without perceptible hypo- thallus. Apothecia sessile, frequent and often oontigmus, to 1.01-1.58m. diam. Disc dark brown booming bleak, persistently plane; margin concolourous with thallus, 0.11-0.14m6 wide, entire becoming crenulate and persistent. Excipulum thallium 60-130p wide; cortex cellular but partly occluded, to c. 20p wide or absent, medulla 50..110p. wide. Peripheral hyphae not pigmented 5.1-5.7p wide, medullary hyphae 2.84.6p and algal oella to 11-12p diam. Excipulun proprium o. 15p wide laterally, expanding to 15-30p wide above, hyphao 1.1- 1.7p wide. Hypotheciun 95.475p deep. Theciun 55•115p high; paraphysee 1.1-1.7p wide, apices to 4.0-.5.7p wide, forming a brown epitheciun. Asa (50-85) x (18-22)p, inner walls unifam. Spores thick, uniformly walled, (14.3-21.7)18.6 x 10.0(7.7.420)p, apical walls c. 2.8-3.8p thick, lateral walls o. thick, pores not present and septum absent or poorly developed. Thallinc reactions:- Kt yellow C:, Hypotheciun I-, theoium I+ blue. Plant corticolous. le 41111011 ../171,
14110 11".111111111 Rino di na ., 11 roboris IDA) A''.
Arn.
4, 'at ,,D` I4.
NIL ES:
iiih• -- —
0
4
Or
0 P!
..„..„ .,
. . .. v 1 III ,‘,.,,,,I g i 4 , S."—".-'1 .
BOTANICAL SOCIETY OF THE BRITISH ISLES DISTRIBUTION MAPS SCHEME
Figure 16. Distribution of Rirodire robrrir ir. 'ritish 145. VariGhtlAy and identification Rinodina roboris appears to be a relatively constant spades in all respects. The species is characterised by the large, plane apotheoia with Granulate margins, by the KOH yellow thallus reaction and by the thick, uniformly thickened spore walls. This oortioolous species sight therefore be mistaken for some forms of R. cdatrana or R. isidloides. It say be distinguished from the former species by the rougher thallus and 'bluntly ellipsoid spores and from the latter species by the look of isidia and much smaller spores.
74920c EMEM There can be little doubt that Itloodina roboris is very closely related to ,R, ointrana but the relationship with 1. isidioides is less clear. The spore morphology indicates a close relationship but the thallus morphology and spore size differ greatly and although these are relatively unimportant characters, taken together they must be -seen: to indicate a fair degree of divergence.
IV Rinod Sect. Plaoota Vainio in Ark. Bot., (4): 69 (1909) pro parte. ,Berongeria Seat. Plaeothallia Travis., Revista Period. Lavori Acoad. Padova, 265 (1851-52)1 ,DisielaGnst Nora., Nyt Nag. Nature., 2 231 (1853) ; Beltraninia Travis., Revista Period. 146.
Lavcrti Accad. Padova, fe: 66 (1857); Sunamaria Subgen. Dimelikens Boist., Nouv. Flora Lich., 2: 89 (1903).
Description
Apotheoial cortex colourless, hypotheoium colourless or rarely brown, spores 1-septate without lumina. Thallus usually with subsquamuloae tendanoies and waxy.
47) Rinodina orustulata Arne in Flora, Regensburg, 12: 40 (1872). Catoleohia fusca Mass., Rioeroh. Auton. Lich.: 84, fig. 173 (1852). - Rinodina contammts f. orustulata Mass., Sohedul. Critic., 2: 161 (1856). - Lecanora budensis Nyl., Flora, Regensburg, Ai: 529 (1881).
ILmicoati Arn. 536, 1581 (K) Rinodina orustulata; Flag. Alg. 98(K) - Rinodina orustulata; For. 20 (K) -Rindaina orustulata; Krypt. Vind. 1258 (K) -R.inodina orustulata; Lojka Univ. 76 (K) Leoanora budensis; Mass.295 (pro parte) (K) - Rinodina oontroversa; 2w. 935, 973 (K) - Leoanora budensis.
Authentic material Mass. 296 (K, - Rinodina controversa P. orustulata.
147.
Description
Thallus thick or rarely thin, ochraceous and areolate. Areolae 0.72-1.0& m. wide, plane to verruciform and often raised at edges to give subsquamulose appearance, waxy. Thallus determinate or indeterminate, without perceptible hypothallus. Apothecia innate be- coming subsessile, to 0.50-0.901m:6 dies. Disc dark brown, usually becoming black, persistently plane; margin concolourous with thallus, 0.07-0.11mm. wide, entire and persistent. ExciPulum thallirum 75-120p wide; cortex cellular, 15-25p wide, medulla 50-100p. Peripheral hyphae pigmented or not to 440.5.7p wide, medullary hyphae 2.8.5.1p wide and algal cells to 18-23p diam. Excipuldm rroprium 15.40p wide laterally, expanding to 25-45p above, hyphae 1.1.2.8p wide. Trypothecium 90-115p deep. Thecium 75-90p high; paraphyses 1.7-2.3p wide, spices to 4.0-6.3µ wide, forming a brown epithecium. Asci (4(d-50) x (12-15)p, inner wails uniform. Spores with- out lumira, (9.7-17.1)13.0 x 5.5(4.6-5.7)p, sectum prominent. Pycnidia sometimes present, irrate, conidia c. 5 x 1p, bacilliform. Thallire reactions:- X-, C., T., P Hypothecium
thecium I+ blue. Plant calcareous.
Variability and identification
The morphology and anatomy of Riroaira crustulata shows
remarkably little variation and the stecies could orly be confused with R. swackhiana which it resembles in the morphology of the thalius and i8.
spore structure. The latter species, however, has a blue-black epithecium K+ iundLtFt. violet and the spores are broadly ellipsoid. rn R._crustulata the epithecium is brown and the spores are narrowly ellip- soid.
Taxonomic remark!)
The species shows a very close relationship with Rinodina twackhiana as has been indicated above. The structure of the spores and the apothccial cortex particularly indicate that the two species form a distinct taxoromic group within the genus. It has not been possible to find any good reason for main- taining the name Rinodina crustulata but until the type of R. fusee has been examined it is considered adviseable to retain the commonly accepted name.
48) Rinodina rrelanconia Vain. in Ark. Lot., 8 (4): 75 (1909).
Authentic material
Holotype"::- Vain. 8762 (TUR) - Rinodina melanconia, Jinretlen, Vainio. 14.9 •
Dead vlion
Thellus thin, dark brown to blackish, continuous or rimoses plane or verruciform. Phallus indeterminate, without perceptible bypothallus. Apothecia sessile, frequent and scmetimes continuous, to c. 0.36ffm. diem. Disc black, persistently plane; margin concolourous with thallus end disc, c. 0.04mm. wide, entire and persistent. Txcipulum thallinum c. 45p wide, cortex absent. Peripheral hyphae c. 5.714 vide, medullary hyphae c. 2.8p wide and alb el cells to c. 20p diem. Xxcipulum proprium c. lep wide laterally, erpanAirg to c. 301.1 above, hyphae o. 1.1p wide. Hy- ethecium dark brown, c. 851u deep. Thocium c. 85i high; paraphyses c. 1.7p vide, apices to a. 5.714 wide, forming a dark brown epithecium. Asci a. 60 x 14p, inner walla not uniform. Spores witheut luTara, (X.2-16,5)14.8 x 7.9(7.4,-8.5)144 septum well developed. - Cl,-- Thnllire reactinno:- Y-,T- , P. Hypethecium I-, thecium I+ blue. Plant siliceous.
Variability tnd identification
Nothing; is known of the variation within the species as only the tyre spectren has been examined. Within the dark brown, saxicolous smt:cies Rinodinc melanconia is unique in having spores without lumina and in having a dark brown hypethecium. Identification of the species is therefore relatively easy although it shows many other characters in 150.
COMM with R.
Taxonomic remarks
Ritodina melsrconia is a taxonomically iediated species differ- ing from the other species without spore lumina in the dark brown, crustaceous thallus. Although the specimen examined does not appear to be very well developed it is unlikely that the thullus would develop to form a characteristic sub-squamulose thallus typical of R. crustulata and R. zwackhian..
49) Rinodima oreinaMa.ss. in Ricerca.. Auton. Lich.: 16, fig. 24 (1852). Leeatora straminea var. oreina Ach., Licheno Univers.: 433 (181o). - Lecanora mouncotioides Idyl., Bull. Soc. lira. Yormandie, ser. 2, 6; 261 (1872). - Rinodita hueana Vain., Hedwigia, ,mss 38 (1898).
Exeiccati
Anzi It. 218 (v) - Rinodira oreita; Arn. 789 (r) - Dimelaera oreina, 789b Dimelsena mougeotioides; Hepp 209 (K) - Psora °reins.; Koerb. 273 (F) Dimeltens orcina var. simrlex; Vind. 44. (F) Rirodina oreina f.,mom,eotioides; Rabenh. 376 (Y) - Dimeleena orcina; Chcacr. 331 (r) Parmelia amine, 478 (K) Parmelia oreina var. crorsiq. 151.
Authentic material
Ach, H.M. 539 (BM) Lecanora straminea var.,oreina,,
Description
Thellus thick or thin, yellow and ereolate, radiate-plicate at margLn. Areolae 0.50..2.34=6 wide, plane or rarely cracked and waxy. Thallus determinate, without perceptible hypothallus. Apothecia innate ar more usually sub-innate, frequent but rarely con- tiguous, to 0.36-0.76mm. diem. Disc black, persistently plane; margin concolourous with thallus, 0.07-0.111:06 wide, entire and persistent. Exvipulum thallinum 90-110p wide or absent; cortex with structure occluded or rarely cellular, 15-20p ride, medulla 70-50p wide. Peripheral hyphae c. 5.1p vide when not occluded, medullary hyphae 2.18-5.1p wide and algal cells to 11-24p diem. Excipulum proprium 10- 15p wide laterally, expanding to 10-20p above, hyphee 1.7-2.3p wide. PYP0thecium 50-15Up deep. Thecium 50..7511 hiuh; paraphyses 1.7-2.3p vide, apices to forming a dark brown erithecium. Asci (40- 45) x (l3-1014 inner walls uniform or rot. Spores without lumina, (9.1-12.5)10.7 x 6.0(4.6-7.4)p, septum well daveloped. Thalline reactions:- K; yellow or =, 1; blue, C:, P* orange- red or -. Hypothecium and thecitsnr+ blue. Plant siliceous. 152.
Variability and identification
The greatest variability of the species is in the chemical reactions and indicates how careful one must be rot to attach too great a taxonomic importance to these characters. Hale (1952) has con- sidered the chemical reactions of :10aasolia1 in detail and found that the species can be divided into three morphologically identical groups in North America. There are two active moups. P+ C- and Ps. 4, and one inactive phase, P- C-. All specimens with a P+ reaction are also Kf yellow but the latter reaction is often difficult to see as it may be hidden by the yellow colour of the thallus. The species is easy to identify as it is the only yellow Rinodjna in Europe and is further characterised by the lack of spore lumina.
Taxonomicremarks
The spore morphology and radiating thallus indicate that the species is closely related to Rinodina orustulata and R. zwackhiana which are commonly found on calcareous rocks. The spores do, in fact, show some evidence of lumina but these are irregular and have ro Torus.
They are therefore similar to lumina sometimes found in the snores of certain Buellia species, e.g. B. =notate. '153.
50) Rinodina swwekbians.Voerb. in Syst. Lioh. Germakt.: 126 (1855). LecanDra xeackhianalromenp., Flora, Regenaburg,17: 145 (1854). - Lecanora tralsvarspiesNY1., Flora, Regensburg, 98 (1886). - Rinodinainummit6 de Lead., Zell. Soc. Dot. Fr.: 22: 613 (1930).
Exsiceati
MTh 4.53 (10 Rinodina xwackhiaral Erb. 1372 - Rinodina Am.
Authentic material
/sotypes- (V) - Lectanora xwackhiana, Bayern, Krempelhuber.
Description
Thallus thick, light grey to oehraceous or brownish, areolate. Areolae plane, sometimes becoming verruciform, waxy, to 0.90-1.26mm. wide, usually radiate-plicate at margins. Thallus determinate, without per- ceptible hypothallus. Apothecia sessile, not frequent and rarely con- tiguous, to 0.43-0.65mm. diem. Discs black, persistently plane; margin copeolourous with thallus, 0.07-0.11m m. wide, entire and per- sistent. Exoipulum thallinum 85-125p, wide; cortex cellular, 20-30.t 154.
wide, pigmented blue-blade, medulla 65-95F wide. Peripheral hyphae 4.0-8.5u wide, medullary hyphae 3.4-4.61 wide and algal cells to 11- 1711 diem. Excipulum proprium 15-2Cip wide laterally, expanding to 15-75p above, hyphae 1.7-2.5p wide. Rypothecium 110-180p deep. Thecium 85-120? high; peraphyaes 1.7-2.3p wide, apices to 3.4-5.7p, forming a blue-black epithecium. Asoi (60-70) x (17-26)14 inner walls not uniform. Spores without lumina, (12.5-19.4)16.1 x 1.3.1)p, septum well developed. Thalline reactions:- R:, I:, 0:1 P:. FlypotheciuraD., theeium I+ blue, epithecium and N+ violet. Plant calcareous.
Variability and identificatiort
The thallus morphology of Rinodina zwackhiana is rather variable and the radiate-plicate margin may not be well developed in some cases. Tn other respects the species is constant and well typified by the ochraceous thallus, the spores without lumina and the blue-black epithecium. This pigmentation may not always be very distinct but the KOR+ violet reaction always confirms the presence of the pigment. Rinodina zwackhiana may easily be confused withb_crustulata because of the similarity of thallus morphology and spore structure. Ebwev-r, the narrowly ellipsoid spores of R. crustulata are quite 155. distinct from the broadly ellipsoid spores of R. swackhiam
Taxonomic remarks
There can be no doubt of the very Close taxonomic relation". ahip of Rinodina swaechiana and R. crustulata for the reasons already stated above. There may also be some relationship with R. 9reina which also has a similar thallus and spore type. However, is a yellow species, a Character that has been used to separate groups within other genera (Runemark, 1956). 156.
Discussion
Mount concepts of lichen taxonomy
The dual nature of lichens and. their nomenclature
The classification of lichens adopted. by Zahlbruokner, (1926) Which has alrea4y been desoribed, is based an the assumption that the algal and fungal constituents of lichens exist as symbionts in the association and should therefore be given equal consideration in taxonomic studies. This assumption malt not, however, be entirely justified. Imshaug (1951) has stated that the genus Karschiginms originally described to include species parasitic on lichen thralli but maw of its species have been shown by further study to be lichen-forming (in the genus Duelliq) with algal gymbionts. Tobler (1911) has described E. itcabrosq and K. ditstructans as being paragymbionts at first, then parasites and, finally after the death of the lichen host, continuing their development as saprophytes. The liohenised state therefare seems to be faoultative for these species and there can be no good. reason for maintaining Kamilla as a genus separate fromulVAllia as Nannfeldt (1932) and Butler (1940) have already pointed out. This relation of a lichen genus to a genus in the PatIllipriao,se is not an isolated. occurrence. Both Keissler (1930) and Nannfeldt (loc. cit.) have shown that there are a series of genera with lichen, lichen-parasite and saprophyte counterparts. 157.
It was this type of anomaly that lead to the Inter,- national Code of Botanical Nomenclature being altered in 1950, when it was stipulated that "for nomenclatural purposes names given to lichens Shall be considered as applying to their fungal components" Cuiberson (1961) has made it clear that this rule has profound implications in taxonomy and a strict adherence to it is not possible. In the first place the exact influence of the alga on the lichen thallus cannot possibly be assessed by herbarium observations and when the fungus is isolated in pure oulture, the morphology of the thallus is not reproduced. This night be taken to indicate that the alga has a considerable influence on the morphology of the thallus. On the other hand, Ahmadjian (1958) has pointed out how very little is really known about lichen algae and in view of this it seems inordinate to give as much weight to the algal constituent as in previous classifications. There is, of course, no evidence that each liohen species has its own specific alga and there is a report (Ahmadjian„ 1962) that different species of algae can occur in the same lichen species in geographically isolated areas. The continuing practice (Duncan, 1958) of separating genera such as Peltideek Peltigera, ,Lobarta -Lobarinal Stiota - Stiotina by the presence of bright green or blue-green algae respectively must therefore be condemned on the basis of present evidence. The discovery of Hale (1957) that fungal isolates of BuelAia stillimciane, when grown under culture conditions, produce conidia that are morphologically identical with those of the imperfect fUngus Snoridesmium folliculatum has lead to the speoulation that the sexual stage of some of the Fungi Imperfecti is found in the liohenised state and that the asexual stage only is free living. It should be remembered that the morphological ditilarity of these oonidia cannot be regarded as final proof that they belong to the same species. However, if it were to be shown that this'state existed for more than a small number of lichen and fungal species the nomenclatural problems involved, would be considerable, more espeoially as the starting date for lichens is 1735 and that for fungi 1821. It is evident from tho foregoing that there is a much closer relationship of the fungal component of lichens with the non-liohenised fungi than would generally have been admitted in Zahibrnokner's time. However, the time has not come to completely disregard the algal component in order to conform exactly to the Code, if only because this document was not conceived to dictate lichen taxonomy. Accordingly, throughout this work the greatest weight is attached to the fungal constituent although the alga has not been totally disregarded in as much as its presence is reflected in the morphology of the thallus and of the apotheoium.
Classification of lichens
Zahlbruokner (1926) divided the Ascoliehens into two groups, those with pyrenompetous ascocarps and those with 159. discomycetous asoocarps. Nannfeldt (1932) has shown that amongst the fungi these two types differ fundamentally only in their mode of spare discharge and should be regarded as no more than, biological groups. In the same work he presented a new olassifioation based on oharaoters.of the ascus and hymenium which cut across the old groups. The Ascomycetes are divided into two subclasses, the Ascohymeniales and the Aecolectlares. In the former subclass the aeoi are unitunioate, are formed from a true hymenium and the ascooarp is a perithecium. In the latter subclass there is no hymenium, the asci are bitunioate and are formed in loculi among plectenchymatio tissue Which is later transformed into a network of paraphysoid threads, and the ascooarp is a pseudothecium. Amongst the Asoohmenialea spore dehiscence is by ejaculation; in the Ascolooulares dehisoence is effected. by an endoascus. Hale (1961) states that "it in now generally agreed that Zahibruckner's system is no longer tenable and that the lichens should be classified among the non-lichenised fungi". Santesson (1952) has discussed the application of the new mycological system to lichens but it remained for Hale (loc. oit.) to make the first tentative integration of the lichen families within the fungal orders. The division of lichens into ascoloctlar and ascohymenial groups was hardly begun but the work of Richardson and Morgan-.Tones (1960, as far as it goes, Sub- stantiates Hale's scheme. The genus Rinodina is thus found underss. 16o.
Class Ascomycetes (Ascoliohens) Subclass Ascomycetes (Ascohymeniales) Order Lecanorales Family Buelliaceae The names in bracket's refer to previous equivalent lichen or fungal names.
Taxonomic status of 'chemical species'.
There can be few genera within the Letanoralis that do not possess 'Chemical species'. By 'chemical species' is meant those species that hare been separated from morphologically identical material on the basis of chemical characters. Hylander (1866a and b) was the first to discover that potassium hydroxide and calcium hypoohlorite give colour reactions with some lichens but not with others. He was also the first to use these reactions, whioh we now know to be due to lichen acids, to erect new species. The patient work of Asahina and Shibata (1954) has elucidated the chemical structure of these compounds and produced a means of identifying them by comparatively simple microchemical tests. According to Thompson (1963) over 120 lichen acids are now known of which the chemical structure of about 80 has been discovered. Asahina was also the first to develop the use of paraphenylenediamine as a colour reagent to test for the presence of fumarprotooetraric acid.previcusly known only by the presence of a bitter taste in some 361. liohens. The lichen acids produced by a given species are often constant and correlate well with morphological characters and for this reason chemical tests can often be used in the identi- fication of lichens with great success. The problem for taxonomists arises when a lichen acid maybe present or absent, or is replaced by other lichen made, in morphologically unifarm material. A good example of the latter case is Thannolia vermioularAq in which some material does not produce thamnolic acid but produces instead two other compounds, squamatio and baeomposict acids (Culbersons 1963). The latter type has been called a separate species and given the name T. subulifornis. Many other genera are known with similar examples, the situation being particularly prevalent in the genera Cladolia, Parmelie and Cetrarta (Dahl, 1952). Both Hale (1961) and Thompson (1963) cite instances where ohemical differences in a species have lead to the realisation that the species were not in reality morphologically uniform and many 'chemical species' are known to occupy divergent geographical areas. Many modern lielenologists would agree withOulberson in erecting two species where one set of lichen acids are replaced by another set in morphologically identical material. The case would not however, be argued so strongly if a lichen acid was merely absent in some but not all material. One objection to the recognition such 'chemical species' seems to be that the number of 162. species described is dependent at the umber of lichen acids discern-0d and the number of possible combinations of acids in a given lichen. Bearing in mind that the principal object of taxonomy is to provide a system of olassification, useful and practical to men, the 'held ooneept of tehemical species' seems to be the direct antithesis of these aims. The identification of *chemical species' is a time absorbing laboratory task which results in the reoognition of large numbers of new and artificial species, ehioh in turn do nothing but help to confuse the classi- fication of lichens. This is not to say, however, that lichen acids are not useful tools in the identification of some species or that they are not worthy of further study. Protagonists of the 'chemical species' school argue that the colour of lichen thalli and pigmentation of the hymenium are used extensively as taxonomic characters and reflect nothing more than the chemical composition of the plant. This &albs true but there can be very few cases where these characters are used to separate morphologically identical material. In view of these considerations it has not seemed advisable to erect 'chemical species' in the present work.
The species concept The species is widely accepted as the basic unit of taxonomy but as Davis and Heywood (1963) have recently stated "the concept of a species is a construction of the human mind and cannot ic3. be defined". There are however, two broad categories of species definitions. The first of these is taxonomic in which a species is considered to be an assemblage of in- dividuals with morphological features in common and separated from other such groups by correlated morphological dis- continuities in spacebar of features. The seoond, or biological category, is of resent origin and not yet widely accepted, and can be of no help in lichen taxonomy at the present time because it involves knowledge of breeding mechanisms. The biological species oonoept rests on the contention that there are objective discontinuities in nature which delimit the units that should be recognised as species. These discontinuities are caused by the restriction of gene flow between actual and potential interbreeding populations. The taxonomic concept of species referred to above is, of course, post•Darwinian and although the present revision is based on the species listed in Zahlbruckner (192240), the origin of many of these species is pre-Darwinian. Many lichenologists were slow to accept the concept of the inherent variability of species so that many later species are also the result of typological thipuing. To these workers the type specimen really did delimit the species. Veber (1962) is therefore largely justified blanking the statement that "The (liohenological) literature abounds in taxa which are die- tinguiellarrom each other by infinitesimal differences in the dimensions of spores, hymenium height and cortical thickness. However these distinctions were established not on the basis of a large series of specimens, but often on the basis of a single observation of, a single structural unit on a single individual, and performed by crude hand sectioning techniques. It is clear that up until the present time the concept of biological variation has played an exceedingly minor role in liohenologr. One of .the prissy tasks of this revision is therefore to recognise the extent to which environmental modification occurs in the genus Rinodina. Weber (loc. cit.) considers that environmental modification in the crustose lichens falls into three general classes. 4) Inhibition or stimulation of growth. The most common form of growth inhibition today is that caused by atmospheric pollution. The occurrence of blanket pollution is now so widespread that the effect of pollution :is often observed at considerable distances from its source. A more natural inhibition of growth is the reduction in the number of apothecia which occurs when many lichens grow in shaded places. Shaded orevasses are, of course, usually sheltered fro! atmos..► pherio turbulence and have a high humidity in consequence. In many oases thane same conditions seem to stimulate the growth of soredia and nay also have a narked effect on the colour of the thallus about which more is said. below. 3.65.
The presence or absence of somata and isidia on the the= of otherwise morphologically identical lichens is often used as a basis for speciation. In an attempt to resolve this problem, Thompson (1948) shows that the growth of isidia could be induced in several species of Peltigera by breaking the *art= of the thallus. Du Riots (1924), however, was impressed by the fact that parallel forms with and without isidia or soredia exist in nature side by side without intergradation and therefore did not feel that the presence or absence of these structures could be explained by environmental modification. Stimulation of growth often occurs in nitrogen rich habitats. The most extreme of such habitats are bird cliffs but the effect may be reduced by leaching associated with high rain- fall or frequent washing in salt spray. Similarly roadsides where nitrogen accumulates in blown dust and occasionally desert rooks where nitrogen accumulation exceeds the very slow rektos of leaching show nitrophilons tendencies. 2) Destruction of accomplished growth. This may take place in at lei two ways. In regions subject to high winds accompanied by blown sand, mechanical erosion may produce profound alterations to lichen thalli. The second type of modification is due to the rapid erosion of the substratum itself so that the lichen is never able to produce its normal growth form. This is often the case with friable sand- stones and certain conifers that shed their bark rapidly. 166.
3) Accumulation of metabolites. Lichens often displgy wide variations in oortieal pigmentation. Orange species eontaining the pigment parietin may often appear yellow or even green due to the lank of the pigment. These forms can usually be oorrelated, with a low level of incident sunlight. Pr uinose foram of species have often attained specific rook. This phenomenon involves the precipitation, by evaporation, of metabolites on the arras* of the thallus or apotheola. In oases that have been analysed the pruina is comprised of calcium oxalate crystals and the presence of pruina can be correlated with the calcium carbonate content of the substratum. It follows that the speciation of pruinose forms is more a description of the sUbstratun than of the lichen. A similar situation exists When iron oxide is deposited in the cortex giving en oohracsous calour to the thallms. This form is, of course, most common on iron rich rooks. The various aberrant morphological forms that have been described above he's all lead to a multitude of incorrect speoiee and should therefore be treated with caution in any tax/made revision. Generally speaking internal characters of the apothecium are less variable and therefore mare reliable oharaaters to use in any such work. They are of course subject to inherent variability and the extent of the variability of any one character 167. differs from species to' species.
Subspecific taxa There has, in the past, been little agreement amongst taxonomists and lichenologists in particular about the definitions of the three oonmon subspecifio units of subspecies, varieties and forms. The comparatively stable use of these terms in recent years stems from the work of Du Riets (1930). Re recognised that some species showed regional differentiation and these races were described as subspecies, usually showing a certain amount of morphological intergradation between themselves. Varietiesiere defined as being local races of species, morphologically distinct and occupying a restricted geographical area. Davis and. Heywood (1963) remark that the variety is often used "for variations whose precise nature is not understood, a treatment often necessary in the pioneer phase of taxonomy", a. statement that would seem to be very pertinent to modern lichenology. The forams regarded. by Du Riots as a sporadic variant distinguished by a single or a few linked characters, without a distinct distribution. The form concept must be rejected as the present state of lichenological knowledge does not allow such fine differentiations to be made, nor is it widely accepted in modern angiosperm taxonomy (Ahti, 1961). The subspecies has rarely been utilised in lichen 1(8. taxonomy, the emphasis here is on geographical rather than morphological isolation and, to quote Davis and Heywood (boo. alt.), "subspeoies may differ from one another in fewer and less well marked characters than varieties or even forms within the same species". It therefore seems preferable to reject the subspecies in this work because of the limited knowledge of the distribution of the species at the moment. Ahti (loc. cit.) refers to Hillman and Grumman (1958) who consider the subspecies superfluous in lichens because of their simple structure. loth Lamb (1951b) and Ahti, however, hats been disposed to use this taxon. 169.
The ontogenir and range of aeotheoial structure within the Liohenised Fungi
Nomenclature of apothecial tissues.
The wide variation of the structure of lichen apotheoia has only recently► been realised. It is necessary therefore, before making any attempt to describe the apotheoia of the Buelliaceae and Phyeciaceae, to review the development and structure of the apotheoia generally. In gathering data for this discussion it was found that there was great confusion in the terminology of the various apothecial tissues and a standard nomenclature had first to be derived. Degelius (1954.) summarised the terminology to that date and arrived at a more or lass satisfactory system of nomenclature. However, a number of new and revived terms have since been used and these have been summarised in Table i. The terms used in the present work are found at the head of the table and it will be noticed that the word thecium and its derivatives has been used in preference to hymenium. The reason for this preference is to maintain a uniformity of terminology and to avoid the necessity of phrasing a new term 'suprahymeniumt to replace the word epithecium. Epitheaiun Theoium .tral_.tiecium Excipulun Exoipullaa .prouriumthallinun
Santeasan Epithecium Hymenium Hypotheciun Escipulum (1952)
Excipulua Excipulum '1952) proprlun thallinum
Degelius EPihymenium Hymenitm Stbhymenium Eacipulun Excipulun • (1954) PriVrilla 01•131.nimi • Gelting Epithecium Papa. Excipulun (1954)
/mahaug Epitheaium Hypotheaiun Exaiple • Anphithecium (1957)
Hens sen iymenium Hypothecium Excipulum Thallus (1963) Margin
Table 1. Summary' of Recent Terminology of Apotheoial Tissues 171.
The ontogeny of the liohen apotheoium
The pairing of nuclei to form a dicaryotic nucleus has never been observed in a lichenised fungus. Pycnidia are found in most lichen genera. They exude miorooaddia which earlier lichenologista (a. f. Smith, 1921) called spermatial assuming they had a sexual funotion. The female reproductive body is the asoogonium which consists of a coil of hyphae and a triohogyne extending from the coil to the upper cortex. Microconidia have actually been observed (Stevens, 1941) attached to the apex of the trichoeyne, evidence not so much of copulation but of the simultaneous occurrence of male and female organs. It remains to be shown that the nucleus of the microconidium enters and migrates down the triohogyne to pair with a female nucleus in the coil. Hale (1961) suggests that the sexual process may have degenerated in many lichens and that the apotheoium develops after the pairing of two female nuclei in the ascogonium. After the degeneration of the triohogyne the medullary hyphae adjacent to the asoogonia aggregate, according to Hensaen (1963a), to form a 'creative layer' from which the excipulum proprium and paraphyees are later derived. The thecium is therefore of dual origin constituting both ascogenous tissue and tissue from the 'creative layer'. The hypothecium is also a 172. tissue of dual origin, comprising the 'creative layer' and the 'hypotheciuml of mycologists (Corner, 1929), itself made up of asoogenous hyphae. These two tissues are however, rarely distinct (see Gelting, 1954). A number of other tissues are sometimes developed in connection with the apotheoia: A stipe is often developed below the exoipulum and may appear to be more or less continuous with the bypotheoiue. This tissue is medullary in origin and its elongation is responsible for the sessile nature of mature apotheoia. This emergence of the apothecium fram the thallus is not to be confused with the increasing convexity of the apothecial disc which often occurs with age and which is duo to enlargement of the hypotheoium. In the case of lecanorine apotheoia an excipulum thallinum is developed outside the excipulum proprium. The relationship of these two tissues is dealt with at length in the following section. Another tissue of medullary origin, the pseudoexoipulue, has been described Hanssen (loo. cit.). It is sitelltvr to the stipe but thick walled, pigmented hyphae proliferate from it to fill the excipulum thAlinum, thus giving it the appearance of an exoipulum proprium.
Range of Apotheoisl structure The early liohenologists (Sehaerer, 1850) utilised the 173. apothecial type in classifying the higher groups of Disoolichens* but only the external aspect of the apothecia was considered. Those species with apotheoiel margins similar in colour to the thallus were called leoanorine; and those with the margin concolouroua with the disci were termed lecideine. It was only later realised that the leoanorine margin (exoipulum thellihum) had a similar structure to the thallus, i.e. it contained algae and overlay a reduced leoideine margin (exoipulum proprium). The oommon concept of a lecanorine apothecium today is, simply, that it contains algae in the margin. Waal* (1890) in his treatise on the Brazilian lichens states that the margin is formed either from the exoipulum thallinum* the exoipulum proprium or from both. He therefore overlooked the fact that the excipulum proprium is necessarily present in both the leoanorine and lecideine spothecial typea. Hue (1906) concluded that the presence or absence of algae in the margin did not necessarily allow one to confirm the presence or absence of an exoipulum thallinum. He demonstrated that algal cells can become trapped in the exoipulum proprium of lecideine apotheoia or converaley that they can be absent from the margins of leoanorine spothecia. He concluded that all leoanorine apothecia possess within the exoipulum thallinum a cortex continuous with that of the thallus, More recently Prey (1936) has added a third class o1 spot/m.1a* the super- lecideine typo, in Which the oxoipulum thmllinum is composed 174.
entirely of a simple thalline cortex. The recent work of Dughi (4952) on the origin and structure of leoanorine apotheois has edema to our knowledge and ahowe that there are a number of intermediate types separating the lecIdeine and lesanorine types. His conclusions are based on the discovery that in the ease of the lsomerine apothecium, the hyphse of the excipulum propriun proliferate and return to their original vegetative state. The emerging spothecium has a pmudothalline margin and the algae of this margin Qome into association with the proliferating vegetative hyphae to form the excipulum thallinta. In the ease of the lecideine spothealum the proliferating hyphas of the excipulum proprium are incapable of returning to the vegetative state and of supporting the algae of the pseudothalline margin. On the basis of these new facts Dughi crested the following six classes of apothecia which are illustrated in Pig. 17.
Locideine apothecia - The amipulum proprium is composed of thick walled* more or less strongly coloured hyphas with reduced cellular lumina, incapable of returning to the vegetative state, and is the only envelope of the apotheolum. Fteudolocanorine apothecia - The excipulum proprium is the same as in the lecideine apothecium but has a clear 1755,
Lecideine Pseudoleconorine
thec
hypo
excip prop
tort -t,1;;Aft:Irifkl• .69:;•14•111.-.1, r algae
Superlecideine Mycolecanorine
thec
hypo excip thall
excip prop
tort
algae
Cryptolecanorine Eulecanorine
thec
hypo tort
algae
excip prop
excip thall
Figure 17. Diewanatio representation of apothecsium types (after Dughi, 1952). 3.76.
colour (biatorine) and contains in an indefinite manner some algae or groups of algae. Superleoideine apotheoia- The excipulum proprita is the same as in the lecideine apothecitm but the extremities of the hyphae return partially to the vegetative state by the direct formation, without passing through ninon-differentiated vegetative state, of an external layer similar to the thalline cortex, with which it is continuous. Wooleoanorine apothecia The exoipulumproprium is weakly coloured or hyaline, of proso- or pare- pleohtenchyma. It gives oft normal vegetative hyphae which form an additional envelops, the excipulua thalliauo, composed of meihillary and oortioal regions but without algae. Bulecanorine apotheoia - The exaipulum proprium is weakly coloured or hyaline, of prose- or para-pleehtenohyma. It gives off normal vegetative hyphae which form an additional envelope which has the anatomy of the thallus. The excipulum thallinum therefore includes, wherever the thallus itself does, a medulla, some algae and an ordinary oortex similar to the upper thalline cortex with Whit& it is continuous. Cryptoleoanorine apothecia - The exoipulum proprium is faintly coloured or hyaline and is proso- or para-pleohtenohymatous. 177.
It is capable of giving off vegetative hyphae but the hyphae become immersed in the thallus mar merely sustain a radial increase of growth. They mingle with the hyphae of the thallus, without forming anything other than a reduced emoipulum thallinum.
These classes form a transition from the undifferentiated leoideine type of apotheeium to the highly specialised eulecanorine type. If the function of the apothecial margin is to protect the ascocerp, then the eryptoleoanorine type without any raised margin might appear, on first considerations, to be a retrograde step. However, it is probibly afforded a similar degree of protection by its immersed position in the thallus and can therefore be regarded as being equally as successful as the other apotheoial types. This might be taken as an indication that the crypto- lecanorine type of apotheaium is following a fundamentally different line of evolution. 178. tuzpholont in Anatol&
Thallus Morphology The morphology of the thalli of most crustaceous lichen species is extremely variable and the species of the genusjtinodisle are no exception. The genus mscr, however, be subdivided into two broad emotions, those species with &pantilese tendencies and those which are strictly crustaceous. The species which exhibit squamulose tendencies may, in turn, be divided into two groups, the first of which has a Characteristic radiate-plioate margin (See Figure 18). lingiag ,oastamomelsA la. crustulata, R. nimboses m. °reins and R. swaokhiant belong to this groups This form of thallus is not very well developed in R. cruatulatta and the squamulose nature is often limited to the central areloae being raised at their margins. The remaining coup of species, P. oonfinis, R. oonfrascsa, R. orassescens, and R. isidAeideso probably should not be termed squamulose at all because their thalli, except that of R. isidioides, consist of well developed, verrucae which possess lobate margins. ln the latter species the squamules develop by a number of isidia merging to fora a palmate, squamulose structure. The species of both these groups have been termed subsquamulose as true squamules are generally considered to be fruticose, scale like tha1l4ne structures and not radiating structures. In the genus Buellia (Sheard, 1964) it was possible to define two groups within the crustaceous species, distinguished by thallus sereat BESMILMItabh ostler s 180.
the areolation of the thallusi Such a division does not 000er in the present genus although it should be mentioned that three of the species previously discussed, R. orustulatg, R. drains and swaokhidinso have regular areolae in the central portions of their thalli. The crustaceous speoies,of the genus therefore possess continuous or risme thalli, or thalli consisting of more or.lesa discrete verrucae, all these forms intargraing with each other. The thallus may be plane, granular or verruoiforso or sorediate or isidiate. The first three types intorgrade although Rinodina eolebin o R. oonfinie, ji&jiedabas R. roboriaamd R. teiohoohila stand out as being characteristically granular. The thallus of R. isidioides, illustrated in Figure 19, is Composed entirely of coralloid isidia end isidia also develop in R. =Gramme. g. furfuracea has a granular thallus, the granules tending to break up into farinose soredia. The surface of the thallus has a matt appearance in the majority of Rinodina species studied but Rs orustulata, R. oreina and R, swaokhiana have thft114 with a wag, appearance. The terricolous and musoicolous arctic-alpine speoies, R. mniarea, R. niabgsa and R. turfacea may develop pruina on the surface of the thallus. It has been assumed that this condition is due to the deposition of calcium oxylate crystals on the surface of the thallus. The characteristic colour of the thallus may be light to dark grey, ochraceous or brown as shown in Table 2, or yellow in j3 *Int 1:1244 awry rlin • a earkinti Awrie X 4/T4/IOTA/Olt"; -itepv).5;.,1 182.
Table 2. CHARACTERISTIC THALLUS COLOUR OF TO
SPECIES OF RpODINA (=RACE= BROWN
R. &equate& A. arenaria R. archasa H. albana R. bisohoffii R. badiella R. atrooinerea. R. eonsooians R. oalolgena R. bilooulata R. controversa oastanossola R. oaloarea R. orassesoens R. hyperbarea R. ointrana R. orustulata R. luridesoens R. oolobina R. occults R. aelancenla R. oonfinis R. masts R. allivina R. coutragosa R. swaakhiana R. nuorenatUla R. oonradi R. sophodee R. eartioola R. turfaoea R. disslaills R. eruapens R. exerescens YELLOW R. exigua R. furturacea R. oreina R. glebulosa ' R. ieidioides R. narltima R. nelsnocarpa R. aniarea R. niibosa Ai polyspora R. pyrina R. rOborls R. saline. R. telohephils R. trivets/an& 183.
oreina only. These colours intergrads and difficulty is often experienced in deciding whether or not an individual specimen belongs to the grey or oehraceous groups or to the ochraceous or brown groups. The grey and brown species are,
however, usually distinct. A further difficulty is often encountered when a grey species is found growing on an iron rich substratum giving the material an oxidised or ochraceous appearance. 'Shea this occurs with a normally oohreseous species it may give rise to a rich red-brown thallus as in the type material of oxvdata. The thickness of the thallus is probably the most variable of all thalline characters and no attempt has been made to measure it because of the difficulties involved. In the most poorly developed specimens the thallus may be reduced to a mosaic of dispersed verrucae resting on the substratum. This condition has been termed evanescent. The thallus is said to be in- determinate when it is poorly delimited around the circumference, as in the latter type. A small number of speoies have characteristically determinate thalli, the following species
representing this group:- Rinodina atrocinerea, R, castaniomela, R. 0in-trona, R. luridesceu, R. nimbosa, R. orei9a, R. souhodee and R. wackhiansi. A well developed, dark hypothallus as in Figure 20 is rarely found among the European Rinodinie. The following species Figure 20. Determinate thallus of Rinodina atroeinerea shoving the dark, limiting hypothallus. Magnification x 7. 185.
TABLE 3. TBALLINE REACTIONS OF RINODINA SPECIES REAGENT SPECIES X - R. atrootnarea + y or - + +or + r R. beadle. R. oiatrana R. oceragpaa R. oortioola R. orassesoens R. exigua R. hyperborea b R. isidioides •+ 7 R. lurideeoens •+ r R. maritime + y + R. malanooarpa + y .0> r + R. ocoulta + y or. + ++ R. oreina + 7. Or um + or - .1. 7 or = + - R. oxydata +y Or an 0. R. polyepora + y or - R. roboris •+ y 186.
do, however, show the character fairly consistently although it may also be developed occasionally in other speciosa.. R. 4Aroolnereal R. 12/Aiello:, Lajakes, R. luridescont, As stialdis and R. occults{. Rummark (1956) has suggested that in the genus Rhisocarcon the development of the hypothallus is influenced by the exposure to sunlight and by humidity. Personal observations suggest that this nay also be true in the genus liastang. Thus in R. occult4 plants found in open habitats rarely develop a hypothallus but when feund, in damp, shady places where the humidity can be expected to be highs well developed black hypothsllus is formed, giving a dark overall aspect to the plant. Chemical motions are often of great help in identifying species and Table 3 gives a summary of the positive reactions observed in this genus. It should be noted that for many of the species listed the reaction with RCil is variable and therefore unreliable for identification purposes. The lichen substances responsible for these reactions have not been analysed but Hale (1952) states that the acids in Table 4 are present inRinodina ardas.
Table 4. Distribution of lichen acids in the three okmical forms of Rinodina °reins
Reaction Lichen acids present
P+, C- protecetraric (?), usnio P-, C+ gyrophorio, usnio P-, usnic 187.
The internal anatagy of the thallus has not been studied in detail because it soon became apparent that it was identical in structure to the exoipulum thalLtmum of the apothooium. Reference should therefore be made to the section dealing with the anatomyr of the apotheoium. The algal constituent of all the speoies described belongs to the genus =mai and is remarkably uniform in structure.
A few species are, however, characterised by possessing algal cells with well developed and readily distinguishable pyrenoids, a feature unrecorded in the majority of species studied.
Apotbsois - morphology Perhaps the most obvious characters of the apothecial morphology are the convexity of the disc and the persistence of the margin. In many species the disc is plane with a prominent thalline margin but in a number of species the disc quickly becomes convex and the margin excluded. The following list of species typically possess convex discs with the margin excluded at maturitys. Alkodina arenaria, 4. bischoffilo R. calcieena,
t graaleacenn, R. dissimilis, R. erumnens, R. furfuraces, luKidenoens,Ruggleman, R. mniarea, R. polvepora and R, trevisimns. It should be remembered that the discs of moat species become somewhat convex with age and may, in exceptional cdrommtances, approach the above species in the degree of convexity.
The margin of the apothecium in those species where it is persistent is usually entire but in some species it may beoome crenulate and this most commonly occurs in Rinodiracastanomele, R. excrescensi R. isidioides, R. maritime., R. roboris and R. turfacea. The size of the apothecia varies widely both between species and within species and the apothecial dimensions of any given species are therefore likely to overlap with many others. Accordingly, this character is made virtually useless both from the taxonomic point of view and for purposei of identification. It was later thought that the thickness of the apothecial margin in relation to the diameter of the disc might provide a diagnostic character for some species but reference to the ratio (TA/GI) in Table 5 shows that this is not the case either. In the majority of the species studied the apothecia are sessile on the thallus. The remainder of the species have sub-innate apothecia, so called because they are not truly sunken into the thallus as in the genus Aepicille or in the Buellia aethalea group of species but are nevertheless very widely attached to the thallus. The apothecia of the sessile species are, on the other hand, rather narrowly attached to the thallus although they lie directly on it. These three types of apothecia are illustrated in Figure 21. The follawing species have been classified as possessing sub-irnate arothecia but it should be recognised that under adverse environmental conditions many other species develop this character:- Rinodina caleigena, R, controversa, R. evresoens, R. plebulosa, R. luridescens, R. melanocarpa, and Re teichoPhila. 189.
TABLE 5. .APOTRECIUM DDENSIONS OF RINODINA SPECIES
SPECIES DIAMETER MARGIN DISC. m/d (a) (a) R. osquatuls 0.36-0.54 0.04-0.07 0.28-0.47 0.08-0.25 R. *Maas 0.50-0.65 0.07 0.36-0.50 0.144.20 R. orohsso 046442 0.04-0.11 0.22-0.51 0.11-0.32 R. arenaria 0.43-0.50 0.044.11 0.21-0.42 0.09-0.52 R. atrocincres 0.43-1.26 0.074.11 0.29-0.61 0.12-0.18 R. Wiens 0.40-0.90 0.07444 0.32-0.62 0.22 R. biloculata 0.29 0.04 0.21 0.19 R. bisohoffii 0.43-0.58 0.07-0.11 0.29-0.43 0.17-0.37 R. calcoroo 0.65 0.07 0.51 0.14 R. caleigons 0.72 0.72 IMP R. oastsaccola 0.90-1.01 0.11 0.684.79 0.14-0.16 R, cintrona 0.61-1.01 0.07-0.11 0.47-0.79 0.1i -0.16 R. colobina 0.37-0.47 0.04-0.07 0.29-0.36 0.10-0.25 R. oonfinis 0.504.79 0.07-0.11 0.32-0.65 0.11-0.33 R. coatroom's 0.58-0.86 0.07-0.11 0.4.34.64 0.124.17 conradi 0.29-0.83 0.04.-0.07 0.15-0.69 0.07-0.49 R. oonaooians 0.434.72 0.07 0.29-0.58 0.12-0.24 R. eontroverso 0.47-0.58 0.07 0.32.o.44 0.16-0.22 R. oorticola 0.504.83 0.04-0.07 0.36-0.68 0.08-0.19 R. arassesoino 0.90 0.07 0.76 0.09 R. oruetulata 0.50-0.90 0.07-0.11 0.29.0.68 0.144.37 R. disolailio 0.4.3 0.04. 0.35 0.11 R. crampons 0.36-0.47 0.04-0.07 0.29-0.33 0.12-0.21 R. exoreacens 1.26 0.18 0.90 0.20 R. exiguo 0.294.58 0.044.07 0.21-0.50 0.08-0.25 R. glebulcsa 044-0.83 0.04-0.11 0.46-0.61 0.09-0.18 R. furturscos 0.61-0.79 0.074.11 0.47-0.57 0.15-0.19 R. byperboreo 0.47-0.58 0.04-0.11 0.36-0.44 0.09-0.30 R. isidioides 0.90-1.12 0.11-0.14 0.68-0.90 0.12-0.18 R. lurideocono 0.79-1.44 0.04-0.11 0.65-1.30 0.04-0.11 190.
TABLE 5 foontinvoit).
R. maritima 0.72-1.26 0.07-0.18 0.58.4.04 0.1-0.20 R. nelanocarps. 0.43 0.04 0.35 0.11 R. seta noonia 0.36 0.04 0.28 0.% R. ailiana 0.58-0.86 0.074.11 0.43-0.65 0.12-0.19 R. =dares 0.72-1.04 0.04-0.11 0.58-0.83 0.054.17 R. auoronatula 0.4.7 0.07 0.33 0.21 R. musoioola 0.51,. 0.04 0.46 0.09 R. :limbos& 0.61-1.12 0.07-0.14 0.54-0.86 0.12-0.19 R. **cults 0.25..0.36 0.04 0.10-0.28 0.14-0.23 R. *raft :0.36-0.76 0.074.11 042-0.54 0.18-0.33 R.°Wats. 0.29-0.4,7 0.04-0.11 0.14-0.32 0.14.0.75 S. polyspora 0.25-0.54. 0.04. 0•1&0.47 0.08-0.20 R. pyrine. 0.36-0.50 060404.07 0.22-0.36 0.10-0.32 R. roboris 1.01-1.58 0.11.0.14. 0.79-1.23 0.08-0.14 R. salina 0.36-0.61 0.04.047 0.29.4.47 0.09-0.25 R. sophodes 0.43.4.15 0.07-0.14. 0.29-0.87 0.16-0.27 R. teidhophila 0.68.4.26 0.07-0.11 0.54.1.04 0.10444 R. trevisiana 0.43-0.61 0.04. 0.35-0.53 0.07-0.11 R. turfaosa 0.90-1.55 0.07.0.14. 0.76-1.33 0.08-0.17 R. swaokhiana 0.434.65 0.07-0.11 0.29-0.51 0.14.0.34 a)
a)
Figure 21. Insertion of the apothecia on the thallus. :eirsilc .dothecia of Rinodina maritimrt; b) eub-inntte apothecia of R. luerleficium, magnifications x 20, and c) apothecia of Buellia.verrucaosa, mr!:;nifloation x 45. 192.
The colour of the thallinc merrin is typically the same as the thallus in species which have eulecanorine apothecia although rarely the margin may be somewhat lirhter or darker than the thallus.
/n four of the species studied, namely Rinodinabiloculata, R. luridescens,
R. coculta and R. oxvdata, the margin is the same colour as the disc as shown by Figure 22. This is a very constant character although a few specimens of R. oxvdatal have been observed in 7;hich the margin was somewhat lighter than the disc.
Pruina have only been observed on the discs of four species.
This condition is typical of Rizodina cintrana and R. niMbosa and of sporadic occurence in R. *Urea and R. turfaceo.. It should be noted that pruina were also found on the thallus of the last three species. It is exceptional to find pruina developing on corticolcus species and one wonders if this can be due to the accumulation of calcium oxylate crystals.
Apotheciaatatomy
.xcipulum Thalilnum In eulecanorive apothecia, characteristic of the genus Rirodina, the excipulum thallinum consists of a cortex and medulla, the latter including the algae -which are uniformly distributed throughout. The excipulum thallinum is therefore very similar in structure to the thallus the only differance being that the algae of the thailus are Figure 22. Pigmentation it mothemium margin. a) Apothooium margin of fet conoolourous with thallus and b) n of P., occult/4. Magnification x 30. 194.
present in a layer below the cortex and are rot uniformly distributed throughout the nedul] a. The cortex is a pseudocortex (c.f. Degelius, 1954) and differs from a eucortex in being poorly delimited from the medulla and con- sisting of more than one layer of cells. In many species the cortex cannot be delimited as a separate tissue and is present only as a peripheral layer cf cells. The cortex is said to be absent in this instance and Table 6 gives lists of species found to possess a cortex or not,
The cortex is not a uniform ctrueture throughout all the species studied and varies in two ways. The upper and lateral part of the cortex may be a primitive eseudecortex which has a n-re or less cellulsr structure, the outer cells often being expanded end pigmented peripherally, or it may appear to be structureless. These two types of cortex sometimes appear in one species and in the latter instance observations have indicated that the cellular nature of the cortex has been occluded by the presence of artifacts, possibly calcium oxylate crystals. This view is substantiated by the fact that those species which arc known to develop pruina, nay also T.essese this typo of cortex.
However, there are merry other species with on occluded cortex that never develop pruina. The distribution of these sertlerl types morgst the species studied is eresented in Table 7. The thickness of the cortex is given for each species in
Table 8 and it will be noticed that in a small number of species the 195.
TABLE 6. PRESENCE a ABSENCE OP CORTEX IN =IMMIX THALLINUM OP Hawn SPECIES
MUM ALWAYS PRESENT CORTEX ALWAYS ABSENT CORTEX pREssnT tit ANEW R. &sambas S. erumpens R. bisdhoffii R. almost R. furfurapea R. oolobina R. ardhaea R. glebulosa R. oonfinis R. areaaria R. luridesoens R. exigua R. atrooiaerea R. melanoaarpa R. hype:tares R. badieila R. melanaonia R. isidioides R. bilooulata R. polyspora R. miliviaa R. (salaam R. pyrina R. molars& R. Wagons R. nisbosa R. eastanomela R. ()milts R. antra= R. roboris R. oonfragosa R. maims R. oonradi R. teidhophila R. oonsoaians R. trevisiaaa R. oantroversa R. turfacea R. oortioola R. arassesoens R. oruatulata R. dissimilis R. exaresoeas R. maritiaa R. eneronatilla R. oreina R. oxydata R. sophodes R. zwaokhiana 196.
TARTm 7. TYPE OF CORTEX WITRIA TI =KUM THALLINUE OF MAMMA, SFECIES,
PRIMITIVE OCCLUDED LUBER TYPE PSEUDOCORTEX CORM PRESENT UR ABSENT R. asquatula R. atrcoinerea R. oonfinis R. &Dana R. aintrant R. oonradi R. ardhaea R. oonfragosa R. exigua R. arenaria R. corticola R. byporborea R. badiella R. crassesoens R. niabosa R. bilooulata R. isidioides R. °mita R. bisoboffii R. maritime R. roboris R. caloarea R. mitres. R. oreina R. oaloigena R. anrdata R. salina R. castanomela R. trevisisma R. colobina R. turfacea R. consociant R. oontroversa R. crustulata R. diestroilis R. excrescent; R. ailivina R. muoronatula R. sopbodes R. teicbcpbilo R. swaokhiana 197.
TABLE 8. DIMENSIONS OF THE TISSSES OP THE APOTHECIUM MARGIN IN RINODINA SPECIES
SPECIES EXCIPULUN THALLIVUE DXCIPULUM PROPRIUM lat. crop. dull. CORTEX MEDULLA TCTALMTH LAT. PEWS. lat. low. R. aoquatula 15.45/4 40 /4 25-65/.50-80i., 1005fr 15-30/A 0.12-0.20 R. albana - 15 4545 60-95 15-20 15.40 0.13-0.33 R. arohaea - 20 15-40 35.75 40-95 10-20 15.40 0.10-0.36 R. arenaria - 15 30.40 40-45 10-15 10-40 0.25-0.33 R. atrocineraa 15-25 30 40000 50-120 10-20 15-40 0.11-0.25 R. banana 10-15 45.55 60-65 15-20 25.40 0.26-0.31 R. biloculata (10-15) 5-35 20-45 10 10-15 0.22-0.50 R. I:deo:Wen - 25 20-70 40-90 15-30 30-65 0.15-0.75 R. oa2oarea 20 85 105 15 25 0.14. R. gulag.= 15 60 75 i5 50 0.20 R. oastancaela 20-40 85-90 105-130 15-20 25-40 0.14-0.15 R. cintrana 10-15 45-60 60-70 15 15-25 0.18-0.22 R. colobina - 10 65-95 65-95 10-15 15-20 0.08.0.18 R. =finis - 15 55-110 70.110 10 20-40 0.06-0.14. R. coafragosa 15-20 70-100 85-115 10-15 10-25 0.05-0.29 R. conradi - 15 35-75 45-90 10-20 15-50 0.12-0.33 R. cansoolans 15 .40-60 55-75 10 20-25 0.14.0.18 R. controvaraa 10-30 40-85 65-115 15-20 20-30 0.11-0.27 R. cortical& 10-.50 40-90 30-65 70-115 10-15 15 0.05-0.20 R. era:sae:scene 65 65 130 20 25 0.15 R. crustulata 15-25 50-100 75-120 15-40 25-40 0.10-0.33 R. diss4milin 10 40 50 15 25 0.30 R. crumpens 40 40.45 40-45 15 20-25 0.33 R. exoreecena 15 90 105 15 40 0.11 R. exigua - 15 25-110 40-110 10-15 10-30 0.06-0.22 R. glebulosa 65-115 65-115 15 15.30 0.11-0.15 R. Purfuracea 50-75 50-75 15 15-30 0.20-0.30 R. hyperborea - 15 40-70 40-70 10-15 15-25 0.17-0327 1 98 .
TABLE 8 (continued).
R. isidioides - 25 40-60 30-.90 55-90 10-20 25-40 0.07-0.36 R. luridoseens - - - 45-90 45-90 MP R. maritime, 10..30 60..75 70-95 10.40 15-25 0.07-0.21 R. melanooarpa - 80 80 5 10 0.06 R. melanoonia - 45 45 10 30 0.22 R. milivina - 15 50-90 65-100 15 25-30 0.12-0.23 R. mniarea - 25 70-90 75-95 15-40 30-50 0.14-0.53 R. muoronatula 25 50 75 15 15 0.20 R. nusoioola 20 30 50 20 25 0.40 R. ninbosa - 30 404.155 55-155 10-15 10-25 0.04-0.27 R. 000ulta (10-30) 25-55 40-65 10-15 1040 0.23-0.27 R. orsina 15.20 30 70-90 90-110 10-15 10-20 0.06-0.13 R. oxydata (1045)+10-20 10-60 30-75 10-15 10-25 0.08-0.30 R. polyspora - 25-65 25-65 10-15 15-40 0.10-0.60 R. pyrina - 30-85 30-85 10-15 15-30 0.08-0.50 R. roborie - 20 50-110 60-130 15 15-30 0.10-0.25 R. sauna - 15 30-75 35-85 15-20 30-45 0.15-067 R. sophodes 15-20 25-30 65.115 75-130 10-20 20-30 0.10-0.17 R. teichophila - 25 40-85 60-95 15-20 20-50 0.15-0.25 R. trevisiana - 15 5065 50-75 10 15-25 0.15-0.20 R. turfaooa - 20 40-70 50-85 65-100 10-20 20.45 0.08-0.20 R. swackhiana 20-30 65-95 85-125 15-20 15-75 0.13-0.18
Zigures in braokets refer to a oortex with an occluded structure due to pigmentation. The measurements may therefore inolude part o! the medullary tissue. 199.
lairs cortex becomes =carded. Amongst these species the following possess a more organised pseudocortex has been termed colic nrsr because of the arrangement of the kaphae:- Rinodipa archaea, R. cortiocis, R. crassescens,,R. isicitoides, R. sophodes and,Ra turracea. This type of cortex consists of closely packed, thick walled hyphen running perpendicularly to the surface of the excipulum thallinum. The columnar cortex is typically hyaline and. han a very Pint I+ blue re- action. In R. arouses} the character is only developed in large, mature apotheoia. The thickness of the medulla is very variable and Probably related to the age of the apothecium. The mcdu3la is a eutivplectenoka- mataus tissue (c.f. Degelius, 1954), made up of loose hyphee and usually with algal cells uniformly distributed throughout it. Rarely and then usually in depauperate specimens, algae are missing from the excipulum thallinum. The dimensions of the by phal and algal cells of the excipulum thellinum are presented in Table 9. The measurements for the cortex refer to the peripheral cells only and these are usually marginally larger than the inner cortical cells. The very wide variatation of these measurements will be apparent but a real difference between the width of the cortical cells and the narrow medullary hyphae is indicated. The peripheral cortical cells may or :Ay not be pigmented at their apices, the tresenco of pigmentation is some cases at least being TABLE 9. CELLULAR VITEli STOWS OF EXCIPULUM TRALLTEUM RIFODINA SPECIES
CORTICAL IMULLARY ALGAL SPECIES HYPHAE MBAS CELLS R. acquatula 4.6-6.3 t. 2.8-5.1 , 14-19 fr. R. albana 4.0.4.6 2.8-4.6 11-23 ' R.'archaea 4.6-6.8 2.3-4.6 12-20 RE 'arenaria 2.8-4.0 2.8-3.4. 13-14 R. atrooinerea _ -5.7, 2,8-5.1 .9-15 R. bidiel1a 4.6-5.7 2.8-5.1 12-23 R. biloculata 4.6- 1.7-2.3 9 R. bischoffii 5.7-8.0 2.8-5.7 10-19 ,R.'caloarea - 5.7 5.7 23 R. caloigona 5.7 2.3 15 R. oastaromela 4.6-5.7 2.8-4.0 14.-16
4.: -R. ' cintrima 203400 11-15 R. colobina 5.7-7.4 2.6-4.0 12-18 R. confinis 3.4-4.6 2.8-4.6 10-13 R. corfragosa - 2.8-5.7 12-18 R..conradi 2.8-5.7 2.3-3.4 11-20 R. consocians , 3.4-4.6. 2. 8-4. 0 114 R. controversa 4.6-6.8 2.8-4.0 24-20 R. corticola .: 2.8-4.6 9-24 R.' orasSesoens - 2.8 11 R. crustulata 4.0-5.7, 2.8-5.1 18-23 R. dIssimilis 4.0 2.8 , 9 R. erumperia - 2.8 2.3-2.8 13-15 R. exoressoens 5.-7 4.0 ' u+- R. cxigua 2.8-5.1 2.3-4.6 11-18 R.' furfuracea 4.0 2.8-3.4 12-18 R. glebulosa 4.0-4.6 2..3- az 13-19 201. otd.) 7.:1311; 9,
R. hyperboroa - 3+4:6+3 2.8-4.0 15-23 R. isidiaidea 2.3-4.6 13-15 R. lurideecers ( 3.4-4.6 ' 9-15 ) R. maritime. , 32:43:.46 9-11 R. rr:elx rocarpa 6.3 2.8 13 R. melarconia 5.7 2,8 - 20 R. miliNrira 4.6.6.8 2.8-4.6 R. =Urea - 3.4m.8.5 2.35.7 1:::: R mucrovatula •4.6 3.4 16 P. misdeals 4.6 2.8 ' 11' R. niriboss +b. 4,0-5,7 15-22' R. moults: - 4.0-5.7 11-17 R. croira ... 5.1. ' 2.., 783• 415 ".106 11F-21+- R. oxydats 4.0-6.3 9i-17 R. polyspora 2.8-5. 7 ::33:44f) 10-23 R. pyriva 3.'....,3 2.8-4.6 '8-24 R. roboris 5,2...'5+ 7 2.6444,6 11-12 R. saliva - 4.0-6.3 2.3-4.6 13-19 Re sophedes 3.4-5.7 3.1!-6.3 13-19 R. toichaphila 4.0-6.3 3.5-5,1 10-14 R. trovisiara 3.4-5+7 2.8-3.4 14-15 R. turfy cea 5.1.'74 2.841,6 12-16 R. zwackhiana. 4.0-8.5 3.4-4.6 11-17
Dashes (-) refer to species which characteristics:IV possess an occluded cortex, 2026 influenced by exposure to sunlight.
The apothecium margin of Rinodira luridescens appears to be unique in the genus in that it rarely possesses a medullary layer in the early stages of its development and sometimes never does. In Table e measurements for the excipulum thallinua of the species have tot been given because of the difficulties of interpreting the anatomy. Hoe- ever, it is evident that the excipulth proprium posseso the facility of becoming vegetative and it is surgested that the species illustrates a stage between the true lecideine and eulecarorire types of apothecium.
Sections of the margin of those species in which it is pig- mented show that the asothecia are of the typical claecancrire type.
The cortex is, of course, pigmented either in total or in pert. The pigmentation is the same colour as the opithecium and -peripheral parts of the excipulum proprium snd this seems to su:gest that the cortical tissue might be reverting back to the nonvegetative state from which it orteirated. The structure of the excipulum thellinum in identical to the structure of the thallus and must therefore be regarded as truly euleeanorine. However, some relationship with the mycolecanorine type of apothecium in indicated and differs from the latter type mainly in the presence of algal cells.
Reproductive tissues
The parts are enclosed by the excipulum thallinum and limited by the excipulum proprium. The excipulum proprium is a much reduced tissue in the genus Rinodina as it is in most of the 203.
genera possessing euleoanorine apothecia. it is commonly a hyaline,
prosoplectenclAymatous tissue, that is to say the hyphae are colourless
and cellular and the hyphal cells are elonate. The tissue is
remarkably uniform in structure throughout most of the species of the
genus.
Table 8 gives the lateral and peripheral dimensions of the excipulum proprium and it will be seen that the tissue characteristically expands in its upper parts although this does rot always occur.
Riredina-biloculata, R. bisohoffii, R. mniarea, R. rolyspora and R. reg.ina are unique in that the development of the excipulum proprium is
such that it may be over half the width of the excipulum thallinum. The peripheral portion of the tissue is pigmented and forms the lateral parts of the epitheciumi the pigmentation being identical with the apices of the paraphyses and illustrating the common origin of the two
tissues. The excipulum proprium merges with the hypothecium below and
therefore does rot appear to form a complete envelope z,rourd the
'reproductive parts as it does in lecideine apothecia. In four of the species examined, Rirodina castanomela, R. erumaens, R. luridescenq and R. turfacea the excipulum proprium has a light brown colour. In all other respects, except in the case of
R. luridescens, the tissue has an identical anatomy to the other species of the genus. It has already been stated that the excipulum proprium of R, luridescens has many lecideine characters and this may again be recognised in the wide, thick walled, semi-carbonaceous hyphae of the
tissue. Table 10 shows that the width of the hyphae of the excipulum propriUm is somewhat greater in this species than in the rest of the
genus. The hypothecium is- generally a colourless tissue in the spedies.
of Rinodina but in R. castanomela, R. controversa, R. dissirilis, . R. luridesceps and R. melanocaTra it is brownish and in R. melanconie
and R. biloculata it is dark brawn. The upper part of the hypothecium - generally reacts V* blue but in sixteen of the species studied the whole of the hypotheciuM gives a positive reaction. These species are: R. albanal.R. archaea, R. badiella, R. confinks, R. confragosa, R. •
dissimilisi R. erumpens, R. exerescens, R. exigua, R. maritime, R. melanocarpa, R. milivina, R. mein% R. pyrina, R. saliva andR. sophodes. The-dual nature of the hypothecium has already been discussed (see page In and it seems that the iodine reaction would be an effective method of
identifying the ascogenous tissue if the distribution of isolichenin (I+ blue) can be shown to be limited to this tissue. Only rarely is it possible to distinguish the ascogenuous tissue from the araphy- spgenous tissue anatomically. The hypothecium is a euthplectenchymatous tissue and it is therefore possible to distinguish it from the excipulum proprium in its lateral Parts.
Six species, Rinodine badiella, R. bischoffii. R. calotzena,
R. luridescens, R. melacocarpa and R. nimbosa have been recorded as
205.
TOLE 10 LT11TSICES OF RITT;OTTC:11.1/E arrssirFs IN R1VODTFA SF YCTES
SPECIES PAR:T117.5ES itPEX FILAMIT f1/4 FJC C17 TILT.r :7(113I T
ca) fib) R. ae quatula 5.1-7.4? 1.7-2.5p 0.27-0.45 1.7-2.3fr R. albara 3.4-4.6 1.7-2.3 C.37-0.51 1.1-1.7 R. arch:: as 3.0-6.8 1.7-2.3 0.37-0.50 1. 1-1:7 R. ar cnari 3.4-3.5 1.1-1.7 0.31-0.50 1.17-2.3 R. atrocinerea 2.374.7 I.1-2.3 0.27-i.57 1.1-1.7 R. I:MI:liana 3.45.7 1.7-2.3 0.30-0.68 R. biloculata 2.341.6 1.1 24-c•ite 1.1 bicchoffii 14. 0-6.6 1.7 (.26-u.43 1.1-2.3 R. calcarea 4.6 3.4 0.74 1.7 R. calciE,ena 4.6 2.3 0.50 1.1 R. castr-noxela 4.0-4.6 1.7 37-CP.43 1. 1-1. 7 R. cirtrana 3.4-4.6 1.1 0.24-0.33 1. 1 R. colobina 4.6-5.7 1.1-2.3 0. 50 1.1.-1.7 R. confinis 2.8-4.6 1.7-2.3 27-0.67 1.1 R. confrajosa 3.5-3.1 1.1-2.3 '...31-0.44 1.1-1.7 corradi .3.4.-5.7 1.7-2.3 C. 33-C. 50 1.1-1.7 R. consoc Inns 4.0-5.1 1.7 0.33-0.43 1.1-1.7 R. ccntroversa 4.0-5.7 1.7-2.8 0.37-0.71 1.1-2.3 corticrin 4.0-5.1 1.1-1.7 0.22-0.43 1.1-1.7 R. eras sescens 3.4. 1.7 0.50 1.7 R. crustu1ata 4.0-6.3 1.7-2.3 0.27-0.50 1.1-2_.8 R. dissi i±11s 4.0 1.1 0.27 2.8 R. erump en 13 2.3-3.4. 1.1-1.7 0. 1.1-1.7 R. excrescens 4.6 2.3 0.5(4 3.7 R. exicua 3.4-5.7 1;1-2.3 0.24-0.51 1.1-2.3 206.
TABLE 10 ctd.)
R. furfuracct 4.0 1.7-2.3 1.7-2.3 R. glebulosa 4.0-4.6 1.7-2.3 0.37-0.57 2.3 R. byperborea 3.4-5.7 1.7 0.30-0.50 1.1-1.7 R. isidicides 4.0-5.7 1.1-2,3 0.24-0.50 1.1-1.7 R. luridescens 3.4-5.1 0.27-0.50 1.7-3.4 R. maritima 1.7' 0.21-0.33 1.1-1.7 R. molanocerpa 4.6 2.3 0.50 •1.7 R. malanconia 5.7 1.7 0.30 1.1 R. milivina 1.7 0.33.0.43 1.3. mnlarea 4.0.5.1 1.7-2.8 0.33.6.62 1.7-2.3 mucronatula 5.1 2.3 •0.45 1.1 R. muecicola 4.7 1.7 0.37 1.7. nimbosa 1.7-2.8 C.33-0.62 R. occulta 4.0-5.7 1.1-2.3 0.24-0.50 1.1-2.3 R. oreina 5.1-5.7 1.7-2.3 0,30-0.44 1,7-2.3 R. oxydata 3.4-5.7 1.1-2.8 6.27-0.61 1.1-2.8 R. polyarcra 3.4-4.6 1.1-1.7 1.1-1.7 R. pyrina 4.0-6.3 1.1-2.3 :427-6.51 . 1.1-2.3 rol,cris 400-5.7 1.1-1.7 0.22-0.42 1.1-1.7 R. salina 3.4.6.3 1.1-1.7 0.27-0.37 1.1-1,7 8 R. sophcden 2. -5.7 1.7-2.3 0.33-0.61 1.1-1.7 R. teichophi1a 3.4-5.7 1.1-1.7 _424-0.50 1.1-1.7 R. trevisinra 1.1-1.7 0.27-0.37 1.1-1.7 R. turfacea 3.4-6.3 1.1-2.3 0,27.-0.44 R. zwackhiana 1.7-2.3 0.33-(_.50 1.7-2.3 207.
possessive a stipe below the hypothecium and of these species the stires of R. lurideseens and R. melinocarra are brownish. In all the • species therefore the stipe is concolourous with the hypothecium and the tissues can only be distinguished by the arrangement of the hyphae. The stipe may always be recognised as a prosorlectenchymatcus tissue with vertically arranged, loose hyphae. The dimensions of the thecium and hypothecium are given in
Table 11. The first thing to be noticed is that in general the hypothecium in -.ore variable in sine than the thocium. This is un- doubtedly due to the fact that the hpothecium continues to enlarge after maturity and gives rise to the characteristic convex discs of old apothecia. Nevertheless the ratio of the thecium to hypothecium appears to be a useful character in the identification of certain species. 'The species of the genus have been classified in Table 12 according to whether the thecium is larger or smaller than the hypothecium. Tho paraphyses are very uniform structures as indicated in Table 10 and nave a characteristically expanded apices. The capitate
apices are pipmented brown but the colour of the epithecium may be influenced by another pigment or pigments which are dispersed in the
gelatine of the thecium. This additional pigment gives the epithecium a red-brown w blue-bled< colour. The distribution of the various
epithecium colours crongst the species is given in Table 13. The
epithecia of the two species with blue-black pigmentation are X+ and }+
violet. 20$.
TABLE 11. =mum OF THE THE= AFD BYPCTEBTDRA IN RIFODTNA SPECIES
SPECIES ThECIUM BYPOTHECIUM t/h. (t) ( h) R. ae quatula 75-115 , 110-115 , 0.67-1.00 r I- R. aibana 75-130 50-95 1.00-2.25 R. archaea 75-110 20-85 1.13-3.75 R. arenaria 75-110 60-100 0.75-1.83 R. atrocinerea 75-140 45-140 1.00-1.67 R. badiella 90-110 110-130 0.69-1.00 R. biloculata 40-70 45-60 0.89-1.17 R. bischoffii 85-115 75-100 0.78-1.25 R. calcarea 100 150 0.67 R. calcigena 115 150 0.77 R. castanomela 90 75-85 1.06-1.17 R. cintrana 75-130 90-140 0.83-1.20 R. colobina 60-95 70-115 0.70-1.00 R. confinis 115-]40 45-90 1.44-3.14 R. confragosa 90-100 60-135 0.67-1.78 R. conradi 90-130 25-75 1.73,4.0o R. consocians 70-90 40-6o 1.17-2.33 R. controversa 75-115 65-90 1.15-1.29 R. corticola 90-115 40-95 0.93-2.25 R. crassescenn 100 130 0.67 R. crustulata 75-90 90-115 0.80-1.00 R. dissimilis 95 5o , 1.90 R. erumpens 85-90 25-40 2.12-3.50 R. excrescens 90 130 0.69 R. exigua 7o-110 45-115 0.89-2.00 R. glebulosa 75-85 55-60 1.25-1.54 R. furfuracea 75-90 75-95 0.79-1.20 209. etd.) Table 11
R. hyperborea 60-85 40-65 1.20-1.62 R. isiBioides 130-160 30-50 2.50-4404 R. luridescens 75-115 70-155 0.48-1.50 R. maritima 95-115 110-140 0.75-1.00 R. melanocarpa 90 140 0.64 R. melarconia 85 85 1.00 R. milivina 75-110 100-140 1.00-1.67 R. mniarea 100-140 75.-130 0.80-1.65 R. mucronatula 90 15 6.00 R. muscicola 90 75 1.20 R. nimbosa 95-130 50-140 0.78-2.50 R. occults 55-85 10-55 1.36-5.50 R. oreina 50-75 90-190 0,37-0.71 R. oxydata 70-115 20-90 1.00-4.25 Rd polyspora 70-100 50-90 0.93-1.37 R. pyrina 70-90 25-60 1.17-3.60 R. roboris )5-115 95-175 0.54-1.20 R. sauna 70-95 25-95 1.00-3.75 R. sophodes 85-130 60-130 0.78-2444 R. toichophila 80-145 55-100 0.85-2.15 R. trevisiana 70-90 20-90 1.00-3.67 R. turfacea 115-150 25-50 1.43-4.60 R. zwackhiana 85-120 110-1c0 0.50-1.80
210
TABLE 12. RELATIVE DIMENSIONS OF, Timm AED HYPOZECIUN IN RINODINA SPECIES
THSCIUM LARGER TIIECIUM LARGER c& SEALLIdl THECTUVI =ALM THAN ITYPOTHECIDII THAN EXPOTTZGIDX THAN Ir/POISEGITAI R. albana R. arenaria R. aequatula R. arches& R. biloonlata R. badiella R. atrooinerea bisehoffii R. et/cares B. oastanomela R. ointrana R. oaloiona R. confinia R. oonfragesa R. colobina A. eonradi cortioola R. orassescens R. oonsocians R. exisua R. crustulata R. oontroversa R. furfuraeoa R. exer8806US R. dissimilis R. lurideseens R. maritima eruppens R. melanomas. R. pelanooarpa R.-glebulosa R. nniarea R. °mina R. hyperborea R. nipbosa Ito sweekbiana R. isidioides R. pcayspora R. pilivine R. roboris R. nazaronatula R. sophodes R. musoloola R. teiehophila R. (=vita R. oxydata R. pyrina R. salina R. trevisiana R. turfalsea 211 .
TABLE 13. DISTRIBMON OF EFITHECTRICOLOURS
DARK BROWN LIGHT BROWN RED-DOWN BLUE DICK R. asqustuls R. stroolneres R. albana R. colobina R. biloculata R. badlillIs R. archaism R. wasoftlimmi R. exlgus R. bisithoffii R. arenaria R. melanoonla R. caloarea R. (=MIA R. oreina R. caloigena R. oortioola R. oastanomela R. enlivens R. ointrans R. exoresoens R. oonfragosa R. slibulosa R. oonradi R. isidioldes R. oonsocians R. maritime R. oontroverca R. =Urea R. orassesosns R. ninhosa R. orustulata R. trevisians R. disslaillis R. turfaces R. fUrftraosa R. hyperbores R. luridesoens R. nelanoosrpa R. nilivina R. munronatula R. occults R. oxidata R. polyapora R. pyritic. R. roboris R. saline. R. sophodes R. teiohophils 212.
The asci are uniformly elevate, the dimensions being given in Table 14. The inner wall of the ascus in most of the species studied is uniformly rounded at the apex but in a number of soecies it may be cone shaped as in the majority of,Buellia species (Sheard, 1964). There are indications, however, that this is a variable character and further studies are needed to elucidate the structure. The asci contain 8 spores, bi-serially arranged except in Rinodina rolvsrora which has 8 - 16 spores in the ascus.
The range of score structure in the genus is remarkable. The snores of the genus Rinodira are usually referred to as being 1-septate although the rrescnt work has shown that a septum, or dividing wall, need not necessarily develop. It would be more accurate to say that spores possess two lumina although in cne From) of the truly septate species these do not occur either. Four typos of spars occur within the genus, the sentate type without lumina referred to above, the polarilocular, mischoblastiomorph and placcdiomorph types. These are illustrated in Figure 23 and their distribution amongst the snecies indicated in Table 15. The septum is usually well developed in the polarilocular group and poorly developed in the mischoblastiomorph and placcdiomorph groups althourh this is not alwrys the case. The dis- tinction between the polarilocular and mischoblastiomorph groups is further confused by the fact that the snores of acme species of both groups tend to develop rounded lumina with age. Fevertheless it is 213.
TABLE 14. ASCITS "IF.Al.11ROF.ICTS2
SPECIES VT= BRY.i;"1:71.11 bil. (b) R. aequatula 45-60 t 13-15 /...... 22-0.29,. R. albana 65-E5 15-19 0.18-0.26 R. ardhaea 40-70 15-26 0.22-0.42 R. arenaria 65-70 19-20 0.27-0.31 R. atrocirerea 50-90 19-23 0.21-0.38 R. badiella 50-65 15-19 0.30-0.31 R. bilcculata 45 19 0.4.2 R. bischoffli 50-70 18-22 0.25-0.40 R. calcorea 70 26 0.37 R. calcigena 60 23 0.38 R. castarcmela 50-65 19 9,30-0.38 R. cintrana 60-70 14-18 0.22-0.31 R. colobina 50-70 13-19 9.22-0.36 R. confinis 85-90 26-32 0.29-0.38 R. confregosa 60-75 16-19 0.22-0.29 R. corradi 65-95 23-28 0.29-0.40 R. corsooiars 60-65 1'5-19 0.25-0.30 R. coritroversa 40-50 19-22 0.37-0.47 R. corticola 7C-85 15-22 0.29-0.50 R. cressecens 85 19 0.22 R. arustulata 40-50 12-15 0.22-0.33 R. dissimilis 60 19 0.32 R. erumpens 65-75 22-25 0.29-0.40 R. excrescens 60 13 0.21 R. exigua 45-65 13-20 0.22-0.33 R. glebulosa 45-50 15-19 0.30-0.43 R. furfuracea 50-60 13-17 0.26-0.28 R. hyoerborea 60-70 17-18 0.24-0.29 R. isidioldes 75-110 22-35 0.20- 0.41 214.4 etd.) TABLE Me* AWIS 11,EAstr.7ETITTS spEct LEYGTH BREADTH b/1. (1) (b) R. lurideacers 50-65 18-23 0.30-0.36 R. maritime. 65-95 15-20 0.19-0.32 R. melanooarpa 60 23 0.38 R. melanconia 60 14. 0.23 R. milivina 50-60 19-20 0.34-0.40 R. mniarea 70-95 19-26 0.27-0.36 R. muoronatula 85 19 0.22 R. muscioola 60 19 0.31 R. nimbosa 50-70 15-19 0.22-0.35 R. occulta 45-50 13-20 0.24-0.44 R. oreina 40-4.5 13-14 0.28-0.37 R. oxydatt 60-90 18-26 0.25-0.40 R. polyspora 50-65 17-19 0.29-0.32 Re pyrina 4.5-65 13-19 0.21-0.37 R. roboris 50-85 18-22 0.26-0.36 R. saliva 50-70 15-21 0.22-0.38 R. sophodes 50-70 15-20 0.22-0.35 R. teiohophila 75-90 20-32 0.27-0.40 R. trevisiara 50-70 14,23 0.20-0.38 R. turfecea 75-100 26-29 0,29-0.35 R. mwackhiana 60-70 17-26 0.26-0.40 215.
A B
C
Figure 23. Spore typos of the genus Rino4-13/11. a) Spores with unthlokened wallso p. orustulpta; placodiomorph spores of R. bileoulate; o) polarilooular spores, R. archaea and a) misohoblastiomorph spores of R. oxydata. Magnification x 4,000. 216.
TAME 15. DISTRIBUTION' OF SPCRE TYPES WITHIN THE RIMDINA
1-SEP TATE POLARILCCULAIR XISCHODLASTIOMORPH PLACCDICMORPH
R. crustulata R. aequatula R. arenaria R. biloculata R. melanconia R. albana R. atrocinerea R. archaea R. creina R. badiella R. cintrana R. bischoffii R. zwackhiara R. calcarea R. confinis R. calcilera R. castancmela R. corradi R. colobina R. confragosa R. dissimilts R. consocians R. controversa R. isidioides R. corticola R. crassescers R. marittma R. erumpens R. excrescens R. melanocarpa R. exigua R. furfuracea R. oxydata R. glebulosa R. hyperborea R. roboris R. luridescens R. milivina R. teichophila R. mniarea R. mucronatula R. rimbosa R. occulter R. polyspora R. pyrina R. sauna R. sophodes R. trevisiara R. turfacea 217. always possible to classify the spores of a given species by examining the structure of the yeunz, immature spores. The dimensions of the spores ore given in Table 16 and it may again be seen that there is a considerable vcriation of size within the species. However, the spore sizes do serve to characterise some species and the shape of the spores as defined by the ratio of the breadth to length (b/1) is another useful character in this respect. Pour groups of species may be distinguished within the polar- iloculai group of species and these are identified by the thickness of the spore wall, the spore length, the shape of the lumina and the septation and size of the spores. The groups of species are listed in Table 17 but it must be stated that they tend to intergrade and should only be regarded as a preliminary grouping. These store tyres are illustrated in Figure 24. It appears that the misehoblastiomorph group of species may also be divided into two sections as only four of them characterist- ically retain the mischdblastiomorph shape of their lumina. These species are Rinodina arenariq, R. dissimilis, R. =data and R. teichooila, the remaining eight species beloncjing to the other section.
Cf the latter species the spores of R. corradi have 1f. or more lumina, the lumina often dividing longitudinally. R. isidioides although it has predominantly 2 lumina also has tendancies in this direction.
These spore types are illustrated in Figure 25. It has already been indicated that the term septum' has 218
T. 16 EFORE UPIFINSICVS CP arcriirl, SPECIES
SPECIES LAMTH (1) EREADTH (b) EJETIZM AVERE.T. EXTREM MEW= R. sequatula 11.4-17.1fr, 14.2/. 6.2 f, 0.41-0.46 R. albana. 16.5-25.6 20.2 e.0-11.4 9.7 0.44-0.51 R. archaea 13.1-25.6 22.8 6.8-11.4 9.7 0.41-0.49 R. arevaria - 17.1-23.4 19.5 8.0-12.0 9.7 0.47-0.52 R. atrocinerea :16.0-21.1 18.1 8.5-13.7 11.2 0.57-0.63 R. badiella 13.1-20.5 15.8 6.3-11.4 8.8 0.51-0.60 R. biloculata 15.747.1 15.6 6.8-8.5 7.6 . 0.49 R. bischcffii 14.3-19.9 16.8- 9.9-12.5 11.3 0.65-0.70 R. calcarea 20.5-25.1 22.7 11.4-14.2 12.8 0.56 R. caloilena 19.4-22.8 21.7 12.5-14,8 13.4 0.62 R. casteromela 14.2-19.9 17.0 6.8-10.3 8.6 0.43-0.57 R. cintrara 14.C-23.4 18.5 8.0-10.8 9.3 0.48-0.52 R. colobira 12.5-23.4 17.1 5.1-11.4 8.2 0.39-0.52 R. corfinis 20.5-35.3 27.0 12..-18.8 14.0 0,51-0.52 R. corfregosa 16.0-22.0 19.7 7.7-10.3 9.0 0.45-0.46 R. corredi 18.2-30.2. 25.4 2.5-14.2 11.7 0.37-0.53 R. consocians 17.1-21.7 19.3 8.5-10.3 9.1 0.46-0.49 R. controversy 12.0-16.0 13.0 7.9-10.3 8.5 0.64-0.67 R. corticola 17.1-24.5 20.9 8.0-12.0 10.0 0.45-0.53 R. cressescens 19.4-24.5 21.7 9.1-12.5 10.5 0.48 R. crustulata 9.7-17.1 13.0 4.6-5.7 5.5 0.41-0.43 R. dissimilis 15.4-22.2 18.3 6.3-9.1 8.3 0.45 N. erumiens 18.2-24.5 22.3 8.5-12.0 10.1 0.44-0,49 R. excrescens 13.7-17.1 15.3 6.3-8.0 6.9 0.45 R. exigua 11.4-22.8 16.5 5.7-9.7 7.5 0.42-0.49 R. furfuracea 13.7-19.9 15.6 6.8-9.7 7.5 0.48 R. glebulosa 13.1-17.7 14.6 6.C-9.1 8.0 0.54-0.56 219: eta.) TATILE 16.
R. Iriperborea 14.2-19.9 17.7 7.4-10.3 8.8 8.494-0.50 R. lel/Asides • 20.9E-31.3. 26.7 11.4-17.1 14.2 0.52-0.55 R. luridesoeM 14.8-22.8 17.8 7.7-12.0 9.1 0.49.0.54 R. , maritime 16.5-22.8 '18.7 8.5-]4.2. 10.7 •0.57.. R. • melanocarpe. 154-18.8 16.8 : 8.55-10.8 9.4 . 0:56 R. stelsmonia 14.2-16.5 14.8 7.4-8.5 . 7.9 . 0.53.. R.• milivine. . 12.1-26.8 20.5 7. 7-12. 5 11.4 0.5.59 R. suliarea ' 18.8-31.3 25.4. 9.7-16.5 12.2 0.47-0.49 R. mucronatu1a 23.9-27.4 25.9. 9.1-11.4. 9.9 •0.38
R. inuseicola 17.7-22.8 19.5 8.0-10.3 . 9.2 0.47 . R. Ambma 17.1-22.8 19.7 8.C-10.8' 9.2 0.46-0.48 R. onculta 12.1-18.2. 14.3 5.7-8.5 7.4 0.47-0.56 R. oreina 9.1-12.5 10.7 4.6-7.4 6.0 0.52-0.63 R. ovfeiata 16.5-25.3 20.7 .8.5-16.5 11.5 0.52-0.63 R. rolyspore. 11.4-19.8 16.3 6.6-11.4 7.4 0.43-0.49 R. pyrina 9.7-17.7 13.6 5.1-8.5 6.3 0.43-0.51 R. re.-,cris 14.3-21.7 18.6 7. 7-12. 0 10.0 0.52-0.56 R. st)lina 11.0-19.8 15.1 6. 8-11.0 8.5 0.54-0.62 R. sophodes 1.3.2-19.8 15.7 6. 6-9.1 . 7.9 0.47.-0.54 R. teichophila 20.5-31.9 26.3 11.1-19.8 14.8 0.55-0.64 R. • treY-Isiara 13.1-23.4 17.9 • 6.8-10.3 8.8 0.47-0.50 R. • turfaces. 21.4.36.3 28.4 10.3-18.2 13.2 0.39-0.55 R. m7ackhiarE 12.5-19.4. 16.1 7.4-13.1 9.5 0.55..0.60 220.
TABLE 17. DISTRIBUTION CF FOUR POLARILOCULAR SPORE TYPES WIT11N TUE GaUS
THICK WALLED TERN wALLED. over c. 20u long under c. iCu long Persist.polariloc Lumira. round Lunina irregular Lumina round with septum with/Jut septum with or without, with aeotum septum R..archaea R. albena R. bidiella R. secipatula, R.:confragosa R. corm:4am R. colobira R. bischoffil R. erumpens R. corticola R. exigua •R. calcarea R. exdrescers R. crassescens R. furfUracea R. calcigena. R. mnierea R. castaromela R. glebulcsa R. controverea R. mucroratula R. hyperberca R. milivlra R. mimbesa R. luridescens R. polyspora. R. .trevisiana R. occults. R. pyrira . R. .turfacea R. snlina R. scphodes 221.
A B
C D
Figure 24.. Types of polarilaoular spore. a) Spore of plasdina jasi iga itth angular lumina and prominent septum; b) spore of R.__elptissLa vdthout prominent septa's; a) spore of R. ex/ vzith samonhat irregular lumina and (1) spore of R. soabodtg 17ith rounded lumina. Ilagnification x 4,000.
C A
Figure 25. Types of mischobIustiomorph spore* a) Typical mischoblastiomorph spore of Rinoaina oxydati; b) spore of R. maritime. with rounded lumina and a)
4Aeolied spore of R. /wadi.. Liatsnifiestion s 14000. 223.
been misused in the genus Rinodina. The true structure of the spore is elucidated by four of the polarilocular species, R. bisohoffifs R. eastancmela, R. colobina and R. controversa, in which the spore wall can be seen to be a dual structure. Figure 26a), of the spore of R. bischoffii, shows that the wall consists of an.outer and inner part and it may be inferred by examining squashed spores that the outer wall is rigid and the irner wall is of a gelatinous nature. The latter is deMonstrated by the fact that the lumina retain their shape after the outer wall is broken and. by the lumina of many species becoming dilated and rounded at maturity. It may also be inferred that the lumina are limited by a semi-rigid membrane., The true nature of the septum is also illustrated in Figure 26a), the old 'septum' often being described as the gelatinous part of the viull between the lumina and around the porus. The dimensions of the lumina and porus are also given in Table 18 for the species studied. Two other spore characters must also be mentioned here. These are the thin apical walls found in Rinodina consocians, R. con- fraerosa and R. mucronatula which are very similar to thin apical walls found in the Buellia disciformis group of species and the meridionally dilated srores of Rinodira cintrara, R. consocians, R. hveerborea„ R. oxydata and R. teichophila. These spare structures ere illustrated in Figure 26b) and is) respectively. 224.
Pieurc 26, pores of a) chr,,,Arp :?lml structure cf the vll; b) r, railororntx2% with thin a7ical end o) IZt teichorbilq showirm lateral thi.c;•-nnirg, of srcre necnifieation x 4,0C;,„ 225
TAME 18. mamas OF span Lunn IN RINODINA SPECIES
SPXIIES LENGTH(1) 1/ext. BREADTH(b) b/ext. KBD'S
R. aequatula i elm NMI MOW R. albana 11.4.17. 0.64.0.70/4 p 0.67/4 2.3-2.8 R. arohaea 8.5.18.2 0.64,0.77 4.0-8.5 0.65.4.73 1.1-2.8 R. arenaria 8.0.1448 0.51.0.59 4.6-6.8 0.57-0.62 1.8-5.7 R. atrootaerea 9.1-14.8 048+040 4.1-6.8 0.40-0.50 2.7 R. Wiens 7.4-16.0 0.70.0.72 2.8-8.0 0.59-0.66 R. bilooelata 11.4.16.0 0.85 3.4-4.6 0.54 1.1 R. bisdhoffii 12.1.17.1 0.86-0.92 74+9.7 0.75-0.77 2.7-3.0 R. oaloarea 13.7-18.2 0.70 6.8-9.7 0.69 R. odlagena 16.5-21.0 0.88 967.114 0.77 5.7 R. mammal& MINIM WIMP MM. =MD fl. cintrana 6.5.-16.0 0.57-0.61 4.6-6.8 0.61 -0.64 1.3-1.7 R. colobina 8.5-12.5 0.53 4.0.547 0.51 1.2 R. confinia 12.0-22.8 0.60-0.63 5.1.10.2 0.51-0.52 11=Ir R. oontragosa 9.1-16.0 0.62-0.63 5.1-6.8 0.62-0.68 2.0-2.2 R. oonradi 11.4-25.6 0.72-0.78 2.8-8.5 0.44-0.56 R. oonsooians 10.3-16.5 0.64.0.68 5.7-8.0 0.67-0.71 1.7-2.6 R. controversa MOW One OM, IMO =NOM R. corticola 114.16.0 0.64,0.65 5.1-8.5 0.65-0.70 1.9 R. crassesoons 12.5-16.5 0.67 6.8-8.5 0.74 3.1 R. orustulata OM. WWI OOMP WINO R. dissimilis 8.5-13.1 0.61 3.4-5.7 0.61 1.4 R. @reopens 11.4-16.0 0.62-0.65 6.3-8.5 0.69-0.71 1.4-2.1 R. excresoens 6.9-10.3 0.55 2.9.4.6 0.51 1.7 R. exigua 7.4-14.2 0.60-0.77 3.4-6.8 0.61-0.73 0.6-1.8 R. glebulosa 8.0-11.4. 0.60.0.64. 0.54-0.55 1.1 R. furfuracea 9.7-14.2 0.73 446.6.3' 1.7 R. hyperborea 8.5-14.2 0.63-0.68 0.61-0.67 1.4 R. isidiodee 12.5-21.1 0.59.0.67 5.5+9.7 0.50 1.1-2.8 R. luridescens 9.9-17.1 0.70.0.78 4.0-7.4 0.64-0.65 1.4 226.
TABLE 18 (oontinmod).
R. maritima 8.5•46.0 0.49 4.64.3 0.53 2.8 R. melanooarps 10.8-12.0 0.68 5.1 0.54. 1.1 R. salanoonia .110115 OMNI IMO R. milivina 9.9-19.9 0.77-0.81 5.5-8.5 0.644.70 2.04.4 R. mniarW 12.5-25.6 0.68-0.73 7.4-14.2 0.72-0.87 2.9-3.3 R. anoronatuls 17.1 0.66 5.7 0.57 2.8 R. ausoioola 11.4-17.1 0.66 5.1-6.8 0.68 2.6 R. nimboia 12.0-17.1 0.72 5.7-8.5 0.74 R. occults 6.3-13.1 0.57-0.72 4.0-5.1 0.57-0.67 1.1-2.0 2. oreina OMB ammo R. oxydata 74.13.2 0.47-0.56 5.7-9.1 0.56-0.71 1.5-3.2 8. PolYsPora 6.3-8.5 0.68 3.44.6 0.61 1.7 R. pyrina 8.5-11.4 0.68-0.70 3.4-6.3 0.65-0.68 1.1 R. roboris 9.7-13.7 0.60-0.66 4.0-6.6 0.50-0.53 A. salina 7.4-14.2 0.66-0.71 4.4-6.3 0.60-0.68 1.i.2.0 R. sophodes 9.1-17.6 0.73-0.84 0.58-0.71 1.4 R. teiohophila 9.7-18.7 0.51-0.65 6.8-11.0 0.54-0.65 2.6-41 R. trevisiana 10.3.16.0 0.71-0.77 4.0-7.4 0.58-0.69 1.1-1.7 R. turfacea 14.2-28.5 0.69-0.77 7.4-1I .4 0.68-0.71 3.0-4.0 R. swackhlana •Nwi 011116 OMPO 41111IND
Dashes refer to those speoies that do not possess lumina and to some additional species which have rounded lumina at maturity and hence lank a pores. 227.
Pvenidia
Pyonidia have only been observed in two species, Rinodina ktrogigprea and R. 'crustulata, where they are more or less innate in the thallus. The conidia are bacilliform and approximately (4. .. 6) x lu in size.
The taxonomy of the genus Ritodina
The species and taxa cf lower rank
The species is the fundamental unit of plant taxonomy and must therefore be attributed the greatest importance. The species concept in relation'to the lichen has already been discussed at length on page 162. The present work has ahem a variability of thalline morphology and anatomical dimensions which has not been suspected by previous workers.
The use of mechanical sectioning techniques iind more advanced optical
systems has also led to a more complete understanding of the anatomy of the species. Thus it is necessary to accept a wider species conception and this in turn has inevitably led to the 'lumping' of species.' Mai= is regarded as the father of the taxonomy of the genus Rirodina and although his works (1895, 1902) only deal with a relatively small number of species they show a remarkably wide•Species conception for the period. Unfortunately the /meat majority of Rim:lima species were not treated by :Tuline and many of them have been described as a result of much narrower species concepts by a great number of authors. 228e
Magnusson (1947) is the only author to have attempted a morographical
'treatment of the genus and this wes limited to the nonmaxicolous species.
-It is difficult to understand Magnusson's view of the species concept. Although he. often points out the inherent variability of a structure he then proceeds to use it as a diaerostie species character. Thus, regarding the spores of the genus he states that "The species with apical incrassation (thickening) of the spore have been divided into two groups: those whore the increseation protrudes inwards and is convex, and those with less thickened and therefore concave inner score wall. Although,qeite distinctive in most species we find others with vacillant (variable) characters either individual or specific". This is'not only true of his treatmert of the genus Rinodina but also of,Aseicilia (1939) where his attitude may be sumearised in his own words:. "The number of species, 114, may seem too high for such a small ;croup, within the genus, and the new species, 51, created by me, too numerous, especially as there are about 50 (with different authors) based upon , only one specimen or specimens from one locality. But modern taxoncmy - works with small units and does not believe in the great variability of species supposed by older authors", and later "I have myself seen so many astonishing examples of new finds of species, previously known in only one or a few specimens that I have lost much of my fear of describing new species upon.one specimen, vie. 7,hen there are apparently reliable characters". The remarks of Heber (1962) regarding
Magnusson's treatment of the genus Acarosrora (1923) seem to sub—
stantiate the views expressed above.
In the taxonomic arrangement of Rirodira species, which he
admits is not entirely satisfactory, he uses the apical incrassation of 229. the spore wall, the constriction of the base of the arothecium, the colour of the thallus and the turgidity of the spores in NCH to delimit the groups of seecies. All these characters are now 'crown, in general, not to be species specific and illustrate the type of character he used when erecting and describing species. Another Character which Magrusson used, and which has rroved to be taxonomically useful in some instances, was the nature of the cortex. There is no doubt, however, that this character lead him astray in the Rinodipa archaee complex of species where it is variable, the very group to which he payed particular attention. For all these limitations of Meerusson's work one cannot help admiring his powers of observation, particularly if one takes into account the techniques and equipment that were at his disposal. The species treated in the present won: are based on a species conception that broadly speaking requires a difference of more than one character in separating species from their nearest relatives. Rarely, the spore character alone has been accepted as sufficient evidence for the separation of species and there cantle no doubt at all that the spore structure and size have proved the most useful of all characters in speciation. There is little purpose in listing the various characters' and numerous combinations of characters that have been used to delimit the species in this work as this information can be found from the species descriptions and taxonomic remarks on the species. 230.
With regard to the taxa of lower rank than species it has
already been stated (see page 167) that the only taxa which the author considers suitable for use in the present state of microlichen tadonamy is the variety. Furthermore a statement made by. Degelius (1990 that "The closer one studies a species, tho smaller will be the wish to distinguish taxa of lower rank", became increasingly true as the work 'proceeded. The writer is now firmly of the opinion that without additional and detailed field studies, which time has not permitted, it is imposnible to establish subspecific taxa on the basis of herbarium material alone. In a number of cases, however, where species are thought to be particularly closely related, the possibility of uniting them as varieties has been mentioned and will be studied more closely uhen the opportunity arises. A complete list of the species studied is given below. basil 96 species have been critically examined and this number includes 5 species formerly classified under the genus Buellia. 47 species have been reduced to synonory and of these 8 have been transferred to Buellias 7 species under the name of b. aethalea. The renaming 50 species include 1 species new to science and 2 raw combinations.
List of Rinodina species studied
species Revised species
Rinodina aeguatula Rthcdina aeppatula(Uyl.) D. de Lead. 231.
R. alba R. atrocinerea (Dicks.) Koerb. R. albana R. albana (Mass.) Mass. Buellia alocisoides R. saline Degel. Rinodina archaea R. arohaea (Ach.) Arm R. archaeoides R. ardhaea (Ach.) Arm R. arenaria R. arenaria (Arn.) Th. Fr. R. atrocirerea R. atrooinerea (picks.) Koerb. R. atrocirerella Buellia aethale,a (Ach.) Th. Fr. R. atropallidula B. aethalea (Ach.) Th. Fr. R. badiella Rinodina badiella (ryl.) Th. Fr. Buellia bilcculata R. biloculata (ryl.) nov.'comb. Rinodina bischoffii R. bischoffii (Travis.) Vass. R. budensis R. crustulata (Mass.) Arr. R. buellioides Buellia aethalea (Job.) Th. Fr. R. calcarea Rinodina calcarea (Sapp) Arn. R. calcigera R. calcigena (Th. Fr.) Lyrge R. cans R. arenaria (Arn.) Th. Fr. R. canella Buellia aethalea (Ach.) Th. Fr. It. oastanomela Rinodina castarcmela Cryl.) Arr.
R. cintrana R. cintrara (Samr.) Sam.
R. colobina R. colobira (Ach.) Th. Fr. R. confinis R. confinis amp.
Rinodina confragosa Rinodina corfragosa (Ach.) ✓cerb. R. coniopta R. luridescers (Anzi) Arr. R. conradi R. conradi Kcerb. 232.
R. consocisns R. consocians H. Magn. R. contribuers R. oxydata (Travis.) Mass. R. controversa R. controversa Mass. R. convexula R. trevasinii (Mass.) Koerb. R. corticola R. corticola (Arn.) R. crassescens R. crassesoens R. crustulata R. crustulata (Mass.) Arm R. demiasa R. saliva Decal. R. diplinthia R. corradi Xoerb. R. discolor R. oxydata (Travis.) ?lass. R. disAmilis R. dissimilis Arzi. R. erumpens R. erum ens H. Magn. R. excrescent' R. excrescens Vain. R. exigua R. exigua (Ach.) 13. Gray R. eurfuracea R. furfuracea H. !lags. R. clabulosa R. glebulesa (Arm) Arr. R. Erenulans R. srchees (ich.) Arn. R. griseofusca 2, oxydato (Travis.) ',lass. R. hyperborea R. hyperborea H, 'Sage. R. immersata Buellia aethalea (Ach.) Th. Fr. R. intermedia Rirodirs ccnraii 1:cerb. Buellia interaolata R., occulta(':oerb.)rov. comb. Rincdina intuta R.'oxydeta (Travis.) Atsr. R. lsidioides R. isidioides (Borr.) Oliv. 233.
Rinodina leoidotropa Rinodina oxydata (Travis.) Mass. R. ligraria R. turfacea(Uahlebb.) "Foerb. pro parte R. archaea (Ach.) Arr. pro parte R. luridescens R. luridescens (Ansi) Arm. R. lusitanica R. corradi Yoerb. R. maouliformia R. exigua (Ach.) S. Gray
R. melanconia R. -relancoria Vain. R. meleroctarpa R. melanocarpa Mull. Arg. R. milivina R. milivira (Wahlerb.) Th. Fr. R. mniarea R. mniarea (Ach.) Th. Fr. R. mniareciza R. mniarea (Ach.) Th. Fr. R. muororatula R. mucronatula H. Vagn. R. murorum R..zwackhiana (Y ramp.) Koerb.
R. muscicola R. muscicola H. 11444 R. nimbosa R. nimbosa (Fr.) Th. Fr. R. ocellulata Buellia aethalea (Ach.) Th. Fr. Euellia occulta Rinodina occults (Koerb.) nov. comb. Rincdina olea R. exigua (Ach.) S. Gray. R. oreina R. oreina (Ach.) Yass•
R. oxydata R. oxydata (Travis.) !lass. R. plumbella R. atrocirerea (Dicks.) Yoerb. Buellia polospera R. biloculata (Vyl.) nov. comb. Rincdina polyspera R. nclyspera Th. Fr. 234.
R. pseudopetraea Buellia sn. (c.f. B. conicps)
R. Pyrina. Rinodina pyrite (Ach.) Arm R. rnmulicola R. exipua (Ach.) S. Gray R. rhododendri R. pyrina (Ach.) Arn. R. roboris R. roboris (Dui'.) Arm Rinedina saliva Rinodina sauna Degel. R. sciodes R. luridescens (Anzi) Arm R. sibirica R. archaut (Aeh.) Arn.. R. sophodes R. sophodes (Ach.) ?'ass. R. sophodiodes R. archaea (Ach.) Arr. R. sorediata R. furfuracea H. Magni R. subarenaria R. oxydata (Travis.) Mass. R. sUberumpens R. teichophila (LA.) .Arn. R. subexigua R. sauna Dego'. R. sUbfusca R. archaea (Ach.) Arm R. subobscura R. archaea (Ach.) Arm R. tegulicola R. occulta (Foerb.) non. comb. R. teichophila R. teichophi3a (1:y1.) Arn. R. trarsaylvanica R. zwackhiana (Tramp.) "verb. R. trevisanii R. trevisanii(Mmss.) koerb. R. turfacea R. turfacea (WahleVh.)Yocrb. R. verrucarioidea R. occulta (Yoerb.) nov. comb. R. umbrirofirusca Buellia sethalea (Ach..) Th. Fr. R. w6ackhiana Rinodina zaackhiana (Yremp.) Ycerb.
New srecies Rinodina maritima nom. sn. 2,35. Subdivision of the genus
The division of the gems Ilinedina into four sections by ZahIbruckner (1926) fellows the scheme proposed by, !,lalme (1902). These sections are discussed in detail below. peat. Creularia This section is characterised by its plecodicmorph spores; and dark hypotheciush Of the species studied in the present work only Ri9cdina bp,seculate was found to ix to the section. This species differs from Tai Insperatt, the species for which 7aIme originally described the group, in having a dark epothedial margin which is concolourous with the disc. 7falme considered that the position of the section in RArodina was umbiF,uous and believed that it could be placed in Ruollia with equal justification. Such a class- ification woul:11 hcwever, seem to Eive tco 'uch weight to the colour of the hypothel-Aum and en there can be no doubt that the excipulum thallirum ccntains algae the section is retained in the genus lirodine. Sect. Eurirodine This aecticn was divided into trio subsections by ZahSruckner. SUbsects. PechYziperia end Corradis. The lntter subsection contained those species with 4-celled spores. In the rresent work only RIX:aline conradi was found to belong to the latter group and it is considered that the spore structure of the species is more closely related to the mischoblanticmerph chore than to the pclarilocular type. SUbs,.,ct. 236.
Conradia is therefore transferred to Sect. MisehoblaAtia. The species with uniformly thickened spore walls of Magnusson's Series Roborina are also transferred to Sect. Mio.leblastia in the belief that their spore structure is a developyent of the mischoblastiomorph type and not of the polarilocular spore. Subseot. Paohvarcria is therefore left with only those species that possess a truely polar- ilocular spore. The problem of finding a natural grouping of species within the section now presents itself. !legnusson found it impossible to propose an arrangement of species entirely to his own sati.sCactiom His arrangement of the Subsect. Pachysporia into three series, (see page 12 ) was probably quite sound although it has been found necessary to remove Series Roborina into another section. His grouping within Series Archaeina, however, cannot be accepted because it is based on the apical thickening of the wall which tends to be a variable character. Thus, the tvAD meet characteristic species of the group Rirodina archaea and R. turfacea have spore walls that are commonly convex inwards but numerous examples have been noted whore the irrer spore walls were concave. It has already been observed that 7n:Trusson's groupings of lower crdors were also based on characters row !:nown to be unreliable at the species level. It is not imperative to subdivide such a section although it may be large but this undoubtedly helps to indicate relationships 237. between species if a ratural arrangement presents itself. A provisional grouping of species within the secticn is therefore proposed based on spore characters and following the arrangement of species in Table 17. This grouping is again, not entirely satis- factory and probably suffers from being too intricate. It is hoped, however, that it will serve as the basis of a mere satisfactory arrangement when more information becomes available. Sect. Misehoblastia This section of the genus was originally erected by Malmo to include those species with truely misohoblastiomorph spores but is now extended to include those species with thick, uniform spore wails. The present studies have shown that the lumina of the mischobiastiomorph spores tend to dilate with age and become rounded which indicates the close relationship of the two types. Two groups of species therefore present themselves within the section, the first being represented by 74agnusson's Series Roborina and Subsect. Conradia in which the spore lumina are uniformly rounded end the secord by the mischoblastiomorph species characterised by Rinodina oxydate. Sect. Placothallie In this section Zahlbruckner included those species such as }lima** niMbosa and R. oreiri which possess scuamulose thalli. However, the spere structure of R. nimboas irdicates suite clearly that the species more correctly belongs to Sect. ,Eurirodire despite the development of its thallus. R. oreinal cn the other hand, has a spore structure not found in any other section. It has already been 238. stated that the spores of this species are 1-septate and without lumina and that this type of spore is also found in three other species. Three of these four species have squamulose tendencies and and further united in having a waxy thallus surface, unique in the genus. The fourth species, R. melarconia, does not possess these characters and appears to be something of a misfit in the section. In other genera, such as Rhizoearron(Runemark, 1954) and Acarosrora (Magrussor, 1923), the yellow species have been given sectional rank. The yellow species of Riredina, represented in Sect.
Plecothallia by R. °rein& and in Sect. 7urirodirla by other species not studied, do not appear to form a natural Ercup of their own united by characters other than thallus colour and they are rot therefore given any status In the genus Rirodira. The arrangement of the species in the following list is believed to show the natural Ercupirgs within the genus as indicated by the rreoent work.
Taxonomic arrangement of the cpecies
of the genus Rinedira S. Gray
Sect. I Orcultrips.
1) Rirodina biloculata (Ey1.) Sheard
Sect. II Eurirodint Stiz. Ser. 1 Archaelna 239.
2) Rinodina arohaea (Ach.) Arn. 3) R. confragosa (Ach.) ScoTb..
4) R. erumpena H. Magn. 5) 1. excrescent; Vain. 6) mniarea (Aoh.) Koerb. 7) R. mucronatula H. 11411271. 8)R. nimboaa (Fr.) Th. Fr. 9) R. trevia'srii (Maas.) oerb. 10)R. turfacea (Wahlerb.) Yoerb. Ser. 2 1 1barina 11)Rinodina albana("ass.) Hass. 12)R. consocians H. Vagn. 13)R. corticola (Arn.) Arn. 14)R. crassescens Arn. Ser. 3 Exiguima 15)R. badiella (1y1.) Th. Fr. 16)R. colobina (Ach.) Th. Fr. 17)R. oxigua (Ach.) S. GM" 18)R. furfuracea H. !iagn. 19)R. glebulosa (Arm) Arr. 20)R. hyperborea H. Maim. 21)Rinodina luridescens (fInzi) Arm 22)R. occults (roerb.) card 240.
Ser. 3 (cont.) 23)R. salinaDegele Ser. 4 Sophodina 24)Rirodina aeopatula(Vyl.) B. de Lead. 25)R. bisehoffii (Travis.) Mass. 26)R. calcaren (Pepp) Arn. 27)R. calcigona (Th. Fr.) Lynge 28)R. csstaromela (ryl.) Arn 29)R. controversa Mass. 30)R. milivira(Wahlorb.) Th. Fr. 31)R. polyspora Th. Fr. 32)R. pyrina (loch.) Arn. 33)R. sophodes (Ach.) nass. Sect. III VashoblastiaTialme
Ser. 1 Czydatina 34) Rinodina arenaria (Arr.) Th. Fr. 35)R. dissimilis Anzi 36)R. oxydata (Travis.) 741-s's• 37)R. teichophila (ryl.) Arn. Ser. 2 Robcrira 38)Rinodira atrocinerez- (Dicks.) l'oerb. 39)R. cintrana (Samp.) Sanp. 40)R. confiris S& p. 43.) R. corradi Foerb. 241. Ser. 2 (cont.) 42) R. isidicides (Born) Cliv. 14.3) R. maritime Theard 44) Riredina melanccarnanall. Arg. 45) R. muscicola H. Meter. 46) R. roboris(Duf.) Nyl.
Sect. IV Placothallig Vain.
47) Rinodina crustulata (!-rass.) Arn• 48) R. melancenia Vain. 49) R. oreira (Ach.) '.lass. 50) R. zwackhiara (Eremp.) hoerb. 242.
Relationships with other genera
There can be little doubt of the close relationship of the genus Rinodina with Buellia and it has already been demonstrated that the tater/end° proximity of the two genera has led to confusion between them. This has largely been due to a misunderstanding of the nature of the apothecial margin. The work of Dughi (1952) has been discussed above (see page 171+ ) and provides the basis for the clearer understanding of the range of apothecial types in the two genera. Using DUghiss classification it is evident that only the eulecanorine tyre of apothecium is fcurd in Rinodina. The apothecial Structure of R. luridescens'is, however, something of an enigma and might be considered by some workers as falling midway between the lecideine and euleoanorine types and by others as being truely lecideine. The position within Bu_e~,lip is more complex as indicated by Sheard (l961.). Within this genus the B. aethalea group of species have innate apothecia and more critical examination has subsequently shown that these apothecia have a characteristic cryptolecanorine structure. The regularly areolate thallus, broadly ellipsoid spores and epithecium colour demonstrate that in other respects also, these species form a natural group, distinct from the remainder which possess lecideine apothecia. 243.
The germs Melsnasoicilia (Vainio, 1909) is of great interest here. . This genus was described to accomodate those species with innate apotbmcia that appear to fall midway between the genera Rinodina ardBuellig. Three sections were established based on the colour of the hypothecium. Five of the species, distributed.in two of the three sections, have been studied in the course cf the present work. Three of the species, ienanora atrocinerella, L. atronallidula and L. umbrinoftnusca, have been shorn to be synonyms of Buellia aethalea and demonstrate the variable colour of the hypothecim of this species. Of the remaining two species, Lecanora intuta is a poorly developed specimen of Rinodinekoxvdata and Lecanora.conioota is a synonym of Rinodina luridescens, neither have truly innate apothecia although the apothecia of R. luridescens are widely attached.
It aprears therefore that the genus Melarainiciliq should include the Buellig aethalea group although it is strange that none of the species
B. aethalea. B. stellulata or B. verruculoqa, some or all of Which
Vainio must have known, are listed.
. All the available evidence suggests that the Buellia aethalea
group is a real and natural group of species but the question of the taatmomio status of the group remains. It is the author's opinion (see pige 177) that the crypotlecanorine apothocium is as widely differentiated from the lecideire as the eulecanorine types cad that the group should therefore be afforded sectional rank within the genus Auell%p, if not reinstated as a separate genus Melenesniciliso The fleal decision of such a course cammot, hoaeVer, be taken, until an exhaustive study of the:whole of the gerus Suellka has beenundertskerwp
It is possible, however, to take an entirely different view - of the classification of the genera within the Duelliaceae• Time, if early the Nurepean ppecies of the fa rAly are corsidersd it is true to say that the spores of the great majority of the species of Nellie and Ainoding are simply 1-septate and polarilocularres7ectively'and Should there not therefore be a strict reorganisation of species within the genera defined by the spore types? Cutaide Europe, how.- ever, and particularly in the Southern Hemisphere and the 7ropies it is possible to finimmy Duellist species with polarilocular spores (Malme, 1927 and Imehaugh, 1955a). It is unfortunate that pt the present time there is not the evidence availe:ae to make such a classification even if further research proved the practicability of the Belem°. Tt is interesting to rote that the apeoies with irrate apothecia ray also possess nolarilceular sprxes as the author has collected two such species, evidentliprevicusly uezrowr, on Jan'. eyen inland.
In placirg the genus Rhieocamon in the Thyrelacese 7Tatton
(1928) suggested a close relationship with,fluellia, and this is further illustrated 1.:y his arrangement or the sections within the two genera, based primarily on the character of nrcre septations Section Catocaroork 245. of Buellia, with 1-septate spores, was transferred to Rhisocarpon by Sheerd (1964) following the perk of Zabibruckrer (1926), Lamb (1940) and Runemerk (1956). Section Diplotomma placed in Rhizocaroon by Watson, has 3iseptate or pseudomuriform spores and a spurious thallins margin to the apothecium, is regarded.as belonging to Buellia by Zahlbruckner and this classification is believed to be correct by the author (Sheard, 1564). With this rearrangement of the sections it is more difficult to maintain a close taxonomic relationship bet weer/ Buellka and Rhisocarpon. The genus Rhizocarpon differs fund- amentally from Buellia in haVing a more elevated, gelatinous thecium, branched and often anast-cmosing paraphyses and large spores usually with a demonstrable hyaline epispore, and for these reasons the genus should le placed in the Lecideacese as Zahlbruckner has previously done. The Buelliaceae anlYhysciaceae are traditionally separated, by their crustaceous and foliose thalli respectively. However, both Buellia and Rinodina, have squamulose species and there seems to "beno logical reason why these species are retaired in the Buolliaealt rather, than the :Physeinceae. These squamuloso species have previously been placed in their own section within the respective genus. Sheard (1964) transferred Buellie eeigen and B. pulchella from Sect. Diraoicia and Sect. Placothallia of Rirodina has now been redefined so that its status does not primarily depend on the squamulose habit of the thallus. Thus RI/Iodine nimbosa has been transferred to Sect. •246;
Eurirodine. These transfers have been made within the Eerera.beeause the species show a closer relationship in other respects with- the crustaceous species of the Buelliaceee than with the foliose species of the Physolseecte. However, one species, Buellia carescens of Sect. DiploicAp does not readily lend itself to classification within the Buelliaceae. Buellisa canescene is unique among the Buellta species of the British Isles in having polarilocular spores. This presents no problems in itself because other Buellia species are known to possess polarilocular spores but further investigations showed that the species has a well developed thalline cortex. ' Like the remainder of the Buellia species and the species of Rirtdina studied the cortex is a pseudocortex: but unlike these species it is made up of closely packed columnar hyphee. This type of cortex is very similar to the columnar cortex that develops in the lower parts of tile excipulum thallinum of certain Rinodina species. The thalline cortex of all other ,Buellia and Rinodina species studied is a primitive pseudocortex, Moro or less, Cellular in structure but never With columnar hyphac. It is evident that the cortex of Buellia canescens is more closely related' to the tyre of cortex found in the Physciaceae which personal obserVations have shown to have similar tendencies. The possibility of transferring Buellia caneseens to the Phveciagene as the monospecifie genus Diploicie must therefore be seriously considered. Such a separation based on cortical structure 247.
would not be entirely without precedent. The genera lentoeiura and
Colleme are separated by possessing a feueurtex and pseudocortex
respectively (Degelius, 1954). The yellcw species of Rhiaocarter, given sectional rank by Runemark (1956),. differ from the remainder of
the genus in having a rseudocortex rather than a eurortex. It would be unwise for anyone who has not specialised in the taxonomy of both the ,Buelliaceae and Physeiaceae to make such a fundamental change of classification tut attention snould be drawn to the possibility. The relationship of the genera within tho Fnvsciaceae is relatively well krown and documented. Irshauah' s suggestion (1957)
that Sect. Dirinaria of Pyxin.e, should be regarded as a serarate genus seems to be.well fcunded in view of the fact that it differs from
both Pyxine- and Physcia in having an aeruginose epithecium, from Prxine
in possessing an eulecanorine margin and from Physcia by having a dark hypothecium. The work of. /mshaugh further showed that the apothecial margin was not lecideine in nature but mycoleceronine ( see page 176)
and that all stages existed between the typical mycolecanorine state and the eulecanorire type. The relationship between the genera Pyxire and Physcia therefore appears to be closer than that between Iuellie and
Rinodina. Unlike the Buelliaceae all the genera of the Physciaceae are united in having polarilocular snores. Roth P,rxine and Piriscie
are subdivided into two sections by the septation of the spores, Sects. Fyxire and Physcia having 1-septate spores *nd Sects. Phragmopyxine and. Hyperphyscia, 3-septate spores. a8.
A taxonomic arrangement of the genera as indicated by the rather narrow limits of the present work is given below. The views expressed on the genera Melarasnicilia and Dioloicia are only tentative
and further work should be carried out to substantiate the status of these taxa. The shore charactera used to define the sections within the genera are also given in the list but it Should be remeMered that in
the Buel iaceae these are not the only characters used to define the sections. In the Phesciaceae additional or more suitable criteria
should be sought for the separation of sections or the sections should be dropped.
Phylog,eny within the Buelliaceae and Physcianeae
The arrangement of species within the genus Rinedira has
already been discussed (see page 235) and was arrived at by attaching a greater iMportance to the structure of the spores than to any other character. This procecaure is known as the weighting of characters
and is the basis of lichen taxonomy at the present time. It has been Shown (see page 227) that mmry of the characters formerly used for the separation of species are subject to wide degree of crefiromental mod- ification and are therefore rejected in the presentwork. This type of reighting.is known as a rriori weighting; as distinct from correlation wei ghting. Correlation weighting is one of the babes of ' .'good1 or 'reliable' characters, i.e. those charooters that' are constantly 249. •
Taxonomic arrangement of the genera related to;the genus Rinodina.
Buelliaoeae Spore type Buellia Sect. Buellia 1 -3-4eptate Diplotomma pseudomuriform
Melanaspicilia 1 -septati
Rirodina 1-3-Septate
Sect. Orcularia placodiomorph
Rinodina polyriloculer Mischoblastia Mischoblastiomorph Placothallia simply 1-septate
Physciaceae polerilocular Diploicia' 1-septate Pyxine Sect. Pyxine 1-septate Phrajmopyxine 3-septate
Diriraria • 1-seotate
Physcia • Sect. Physcia 1-septate Hyperphyscia 3-aeptete
Araptychla 250.
associated. Unfortunately there are so few characters available to the lichen taxon:y:1st that Is is not possible to work cut these corre- lations statistically. No doubt some useful characters have also been overlooked intheprescnt work but with the increased knowledge of the ontogeny of the anothecial and thalline structures the number of useful characters has been increased.
At the genus and family level the of characters useful
for indicating relationships is further-reduced. For the trio families
under consideration the spore structure, thallus habit as distinct
from the morphology, apothecialtype and cortical structure appear to be
.host suitable for this nurpose. 'Tout modern classifications have
claimed to be rhylo:enetic but as in the cane of the Anaiosperms- the phylogemy of the lichens has to be inferred because of the lack of fossil
data to act as a guide., Nevertheless there are some characters that may
be accepted as being more primitive or less specialised than others. Tn'the present case crustaceous species may betaken to be
less'specialised than the squamulose. ortoliose species; the lecideine type of 'apothecium more primitive than the myColecancrine, eulecanorite or cryptolecanorime apotheoiumv the cellular cortex more primitive' than- the columnar cortex; the dark hypothecium 'less apecialised than the light hypothecium; , the.simplY.1-septate spore more primitive than
the placodiomorph, polarilocular or.mischoblcstiomorph spore; and the
1-septate spore less spedialised than the 3-sept6te cr pseudomuriform spore. The spore characters, however, appear to be more useful for classification within genera thaM:betweemH 251. genera. It must finally be appreciated that advanced characters may be closely correlated with more primitive characters and this come plioates the construction of any phylogenetic tree. A derdrogram has been constructed below (Table 19) to sheer the inferred phylogenatio relationship of the genera.
The dondrogram indicates all stages of the most primitive genus, with its cellular cortex, erustaceous thallus, locidcine apotheoia and. dark hypothocium to the most advanced genera, Phyacia and Apaptvehia, with their columnar or horizontal cortical hyphae, foliose thallus, eulecanorire apothecia and light coloured hypothocium. The relations ship of the cortical structure of the latter two gerera is fioubtfel, it seems possible that neither should be regarded as being .mare spec- ialised than the other. Some' workers might-ergue that the morophy- letic origin of the genera shown by_ the derdrceram is false on the groubds that not enough weight has been attached to the spere structure,
If new evidence were to be put foruard rhich irdicated that a greater importance should be'attached to the (mores it would then have to be admitted that the genera of the two familien were of polyphyletic origin and were therefore less closely related than has been suggested. here. 252.
TABLE 19. DENUROGRAU mama PHYLOGEDETIC RELATIONSHIPS BET ZEN THE GENERA OF TUE BUELLIACEAE AID PHYSCIACEAE.
Buellia Nolanaspicilia Rinodina Diploicia Pyxino Dirinaria Physcia Anaptychia I 1 I I I LJ I Hypothecium Dark Dark Light Light Dark Dark Dark Light Light 1 1 I I I I Apothecium type Lecidoine Cryptolecanorine Lecanorine Lecideine gycolecanarine Lecenorine Lecanorine I I I I Thallus habit Crustaceous ± Squamulose Squamulose Foliose Foliose 1 Cortical hyphae Cellular Columnar Horizontal
Common Ancestor
(A hypothetical crustaceous lichen, with cellular cortex
and lecideine apothecia) 253.
Summary
The arrangement of the species of Rinodina into sections was first fully carried out by Malme (1902) and this system was later used by Zahlbruokner (1926). Magnusson (1947) undertook a taxonomic: revision of the nan-saxicolous species in Europe but apart from this no monographic treatment of the genus has ever been undertaken. There was therefore an urgent need for a comprehensive revision of the genus Rinodina, more particularly because the advent of mechanical sectioning techniques, refined optical equipment and a clearer understanding of the limits of cnvironmental modification (17eber, 1962) would undoubtedly provide adaitional information on which to base such a taxonomic rev= ision.. The present thesis gives the results of the first two years work of these studies and deals with over half the published species of the genus in Europe. Of the 96 species studied 41 have been reduced to synonomy. The remaining 51 species include 1 new to science and 2 new combinations. The work has also involved revising the limits of the genus and 5 species are now included in Rinodina which were formerly in the genus Buellia. Conversely 8 species of Rinodina have been transferred to Buellia, 7 of them under the name B. sethalea.
Yore important than these changes of nomenclature, however, are the changes in status of taxa of higher rank within the genus. 254.
Subsea. Conradia of Sect.- kurinodina fcrmerly distinguished from
SUbsect. Pachysporia by its 4-6-celled spores has been transferred to
Sect. Mischoblastia.because of the similyrity of spore type. The
species of Magnusson's Ser. Roborinao of Subseet. Paclarsrorie. have
Also been transferred to Seat. Visehoblastia for the same reason. Finally Sect. Placothallia has been redefined to include thoae species with spores lacking lumina although it still retairs same of the squamulose speCies.
The revision of the limits of the eerus Rirodina.has brought to light some interesting facts-regardirg the relationship of the genus with the other genera of the Buelliacene and PhY_siaceae. The work of
Dug hi (1954) and the author (Sheard, 1900 suggesta that the genus Melaraspieilia Vainio should be reinstated and should include the
Buellia aethnlea group of sPeciee. The apothecia of these species are cryptelecancrine and are therefore distinct from the lea deice arothecia of and the eulecanorire arethecia of Tirodinsw Tt is also suogested that nuellia, careseera of Sect. DiPloicia be regarded as a morospecific genus and transferred to the Physciaceme on the i!rounds that the columnnr cortex of its thallus has a closer counterpart in this family than in the Ruelliecee. The other British species of Sect. Diploicia have been traneferred to Sect. Eubuellia by the author (loc. cit.). The relationships of all the genera of there two families is nummarieed in Table 19 (page 252). Ack.nowledrements
I am very greatly indebted to,my-Supervisor, Dr. Eershamr, for his guidance and encouragement durirg my-studies. For the loan of herbarium material I am very grateful to the curators of the herbaria listed on page 16 and also for the private collections loaned to me • by various individuals. ...The help of the Nature Conservancy is also _acknowledged for the -6roparation of the distribution maps of ,he British sp6cies and of Mr. T. Homo for the prciaration of the photo— graphs. 256.
Bibliozraohr
Aeharius, E. (1798) Lichenographiae Suecicae prodromus, Linkoping. Acharius, E. (1803) Methodus Lichens, Stockholm.
Marius, E. (1810) Licherographioauniversalis, Gettingen. Ahmadjian, V. (1958) A guide to the identification of algae occurring
as lichen symbionts. Bet. Votiser, (4): 632. Ahti, T. (1961) Taxonomic studies on Reindeer lichens (Cladonia sUbgenusCladina). Suomal. Blain-ja kasvit. Seur. (1)* Asahina, Y. and Shibata, S. (1954) Chemistry of lichen substances, Tokyo. Butler, E.T. (1940) Studies in the Patellariaceoe. Mycoloria,Ip 791. Corner, E.J.H. (1929) Studies in the morphology of Discomycetes : 1 the marginal growth of apothecia. Trans. Brit. ”ycol. Scc., „al 263. Culberson, W.L. (1961) Proposed changes in the International Code
governing the nomenclature cf lichens. Taxen, 10: 161. Culberson, W.L. (1963) The lichen genus Tharnolia. Brittonia,11:
141. Dahl, E. (1952) On the use of lichen chemistry in lichen systematics.
Rev. Bryol. Lichen., 21: 119. 257. Davis, P.H. and Heywood, V.B. (1963) Principles of Angiosperm tax- onomy, Edinburgh and London. Degelius, G. (1954) The lichen genus Collema in Europe. Symb. Bet. Upsaliens., (2). Dughi, R. (1952) Tin problem de lichenologie non resolu : L'origine et la signification de l'apothecie lecanorine. Arn. Fac. Sci. Marseille, Ser. 2, AL 218. Du Rieta, G.E. (1924) Die soredien and isidien der flechten. Svensk Bot. Tidekr., 18: (3) 371. Du Riots, (1930) The fundamental units of biolcgidal taxonomy. Svensk Bot. Tidakr., ze 333. Duncan, B.F. (1959) A guide to the study of lichens, Arbroath. Fee, A.L.A. (182k) Essae sur lee crrptogames des ecorces exotiques officinales, Paris. Flbrke, H.G. (1815-19) Deutsche liohenen, Berlin. Frey, E. (1936) Vorarbeiten zu einer monographic der Umbilicarinceen. Her. Schweiz. Dot. Cos., b4 198. Frey, Elias. (1831) Licherot raphica Europese reformats, Lund. Gelting, P. (1954) The Rhizocarpon species with peltate areoles occurring in Europe and N. America. Svensk. Bet. Tidskr., 511 71. Gray, S.F. (1821) A natural arrangement of British plants, London. Grumman, 17. (1963) Catalogue lichenum Germanise, Stuttgart. 258.
Hale, M.E. (1952) Studies on the lichen Rinodina °rein° in N. America
Bull. Torrey Bot. Cl., 22: 251.
Hale, LE. (1961) Lichen Handbook. A guide to the lichens of
Eastern north Amerada. Smithsonian Institution Pubi. 43341
Washington.
Hale, M.E. (1957) Conidial "stage of the lichen fungus ,Buellia
stillirpiana and its relation to Speredesmium folliculatum.
Mycologia, A2: 417:
Hemmer, A. ( 1963) The north American species of I,Tassalongia and generic relationships. Carad, J. Dot., as 1331. Hillman, J. and GrArnma.n, V. (1958) i:ryptogamerflora der mark Brandenburg
and Angrozerder, Berlin.
Hue, 1..7. (1906) Lichens morphologico et anatomic° dispositi.
• Arch. Mus. Hist. Nat. Paris, Ser. '4, 8: 237.
Imshaug; H.A. (1951) The lichen forming species of the genus Buellia'
occurring; in the United States and Canada. University
7licrofilms Inc., Ann Arbor.
Imshaug, Ii.'A. (1955) Lichen genus 13unllia in the West Indies. Farlowin, k 473. Imshaug, H.A. (1957) The lichen genus Pvxine in North and ?.Middle Arzrica.
Trans, Amer. Mier. Soc., 21: 246.
7eissler, K. von (1930) Die flechtenparasitcn in Raberhorst,
Irrypteramen flora, 8. Foeber, G.G. (1839) De gonidiie licherrurn, 255
roerber, G.C. (1855) Systems lichenum Germanise, Breslau.
Lamb, T.M. (1940) Liehenological rotes from the British Museum Herbarium. IV. Miaow/mon sect. Catocarrion in the British Isles. J. Bet., Land., jk 12:. Lamb, I.% (1951) On the morphology, phylogeny and taxonomy of the lichen genus Stereocaulcn. Cared. J. Dat.,22: 552. Lamb, I.M. (1963) Index nominum lichenum, New York. Linnaeus, C. (1753) Species pluntmrum, Stockholm. Lynge, B. (1915) Index specierum et varietatum lichenum ciao collectionibus 'Lichenes Exsiccati' distributae mint, Kristiania. Lyrge, B. (1924) On S. American Anaptychiae and PhysciAt. Skr. Videnskselsk., Christ., (16). Magnusson, A.H. (1929) A monograph of the genus Acarospora...K. Svenska VetenskAocid. Handl.,2 (4). Magnuson, A.H. (1939) Studies in the species of Lecaroras mainly the Aspic ilia gibbose group. K. Sverska VetenskAcad. Handl., .112 (5). Magnusn-on, A.B. (1947) Studies in non-saxicolous species of?irodina. Acta Mort. Gothoburg., .0 11. Malmo, Gust. C.M.V (1895) Rinodinaanphodes ooh RinotAna exiKua. Svenska VetenskAcad. Harald, A (11): Maine, G.O. Aal (1902) Die flechten der estenReemellschen7xpedition. II. Die gattung Rinodina. Bih. K. Svenska VetenskAca0. Handl., 28 (3): 26°.
Maim, Gust. O. A:N. (1927) Buelliae itineris ReErelliami prime. Irk. Hot., 21 (14), Nissalongo, A.B. (1852) Ricerche sull'autonsmia del licheni crustosi, Verona. Mudd, W. (1861) A manual of British lichens, Darlington. Nannfeldt, J.A. (1932) Studien Ober die morpholoEie and systematic der nicht-lichenisierten inoperculaten Discomyceten. Nova Acta Soc. 201. Upsal., Ser. 8 (2). Notaris, G. de (184.6) Framerti licheno;:refici de un lavora inedlto. G. Bot. Ital., 2: 174, 299. Vylander, W. (18%) Essae d'une nouvelle classification des Lichenes. Mems. Soc. Sci. Nat. Cherbourg, 2: 5. Nylander, W. (1866a) Circa rovum in studio lichenum criterion chemica n. Flora, ReLonsburg, Al: 198. NYlarder, W. (1866b) fuaedans addenda ad nova criteria chemica in studio licherum. Flora, Rerensburg,.42: 235. Reinke, J. (1894..96) AbhandlurgenT3er Flechton. I-V. PrirEsheim, Jb. Vass. Bot., 26-29. Richardson, D.N.S. and Morgan Jones, G. (1964) Studies on lichen asci. I The bitunicate tyre. Lichenolorist, 2: 205. nunemark, H. (1956) Studies in Rhisocarron I. Taxororly of the yellow species in Europe. Op. Bot. Lund, 2 (1): 261.
Su Lesson, R. (1952) Foliicolous lichens. 4mb. sot. UpErdicrE., JO 1. Schserer, L.E. (1650) Ennumeratio critics lichenum '12ropteorum, Berme. Schwerdener, S. (1867) Lieber den bau des flechtenthallus. Verh. Pchweiz. Yeturf. Ges. :88. Scott, C.D. (1959) Stable Pd solution, Lichenologist, 1: 88.
.heard, J.';. (1964.) The genus Buellia de rotaris it the British Isles. Lichenologist, 2: 225. Smith A.L. (1521) Lichens, Cambridge. Stevens, R.P. (1941) Yorphology and ontogery of Pormatccurrrn
aquaticurn. Amer. J. Dot., 28: 59. Swirscow, T.i;.V. (1962) Pyrerocarpous Lichens : 3. The petusrorinn in the British Isles. Lichenologist, 2: 6. Thom;pson, (1548) Experiments upon the reEeromticr of certain
species of .reltigcra cnd their relationship to the taxorcmy of
. the genus. Bull. Torrey Dot., 486. Thompson, 0.17. (1963) Modern species concepts: Lichens. 1=r;:ologist, 66: 94. Tobler, F. (1911) Zur biologie von flechten and flechtenpilzen. T. Jb. VAss. Bot., ha: 389. Vairio, (1890) Etude sur la classificstior rwturelle et la • Morphologic des lichens du Eresil, Helsirgfors. (1909) Lichenes trope ritlekai collect/. Ark. Pct., 8 (4): 175. 262.
Viallroth, F.V. (1825) Vaturgeschite der flechten, I, Prrnkfurt am
Yttson, W. (1920 The classification cf rehens. Tow Phytol.,
1, 85. Matson, V.. (1953) Census Catelctue cf the Pritish Lichens. 7'ritish Mycological :;ociety, London. Vieber, W.I. (1962) Trvirormentcl modification cnd the trotoromy cf
the crustose lichens. Swinsk Bot. Tidsr., Ihrialk (2): 293. Z&hlbruckner, A. (1926) In Enrler, A End Prartl, Latbrlichen pflanzenfamilien, 8, ed 2, Leipzig,. Zahlbruckner, A. (1922-40) Catalorls lichenum universals, 1-2. Leipzit.