Annual General Meeting 1993: Ripon

HomeEventsAnnual General Meeting 1993: Ripon

18 September 1993 - 19 September 1993

Meeting report


The pleasant campus of University College of Ripon and St John, one of the ‘new’ universities, was the venue for this year’s AGM and paper-reading meeting. Ripon’s reputation as one of the most genteel towns in the north of England certainly seemed to be deserved. It was possibly the first time that the Society has shared a venue with the Mother’s Union, who were there in force. Mike Longman excellent organization ensured that a comfortable and interesting weekend was had by all, including a beautiful woodland for the Sunday excursion. My thanks to all the speakers for giving a range of very interesting talks. The following summaries have been provided by the authors.

Nick Hodgetts

Prof. P.W. Richards (Cambridge): Richard Spruce, the man.

Richard Spruce, whose centenary is being celebrated this year, was by any reckoning one of the greatest botanical explorers of the nineteenth century and he was also a great bryologist. He was a Yorkshireman, born on 10 September, 1817, at Ganthorpe near Castle Howard in the North Riding. After he returned from his South American travels he settled first at Welburn then at Coneysthorpe about a mile from his birthplace. In fact, though he spent a year in France and fifteen years in South America, he spent the greater part of his life in his native county.

His father (who had the same name) was a schoolmaster, first at Ganthorpe and later at Welburn. He had a reputation as a mathematician and is said to have been ‘highly esteemed and efficient’. The maiden name of his mother, who died when he was eleven years old, was Etty and she was related to the famous artist of that name. Richard was mainly educated by his father, but he had lessons in Latin and Greek from a retired schoolmaster who, to judge from Spruce’s proficiency in the classical languages, must have been no mean scholar. Spruce never attended a university but he was awarded a Ph.D. by the Academy of Sciences in Berlin 1864, presumably in recognition of his published work and eminence as a botanist and explorer.

When he was 23 years old he became a mathematical master at the Collegiate School in York. For a while enthusiasm for mathematics supplanted his love of botany, but not for long. Having a regular salary and plenty of time in the school holidays, he explored many parts of Yorkshire and made many botanical discoveries. It was while he was at York that he began to collect and study bryophytes. According to his own testimony, his first ‘advisor’ on mosses was Sam Gibson, a tinman or ‘whitesmith’ of Hebden Bridge, who, like a considerable number of working men at that time, was a keen naturalist. Sam kept a copy of Hooker’s British Flora on his workbench, which in parts had become so begrimed as to be illegible.

In the 1840s Yorkshire was a fine field for bryologists. Spruce was particularly interested in the bogs and fens of the Vale of York which were already beginning to be drained and destroyed by the steam-plough. In some of them Paludella, now extinct in Britain, could still be found and he discovered Helodium blandowii, now also extinct, near Terrington Carr in his home district. On Strensall Moor there were tussocks of Leucobryum glaucum a metre tall. When Spruce took William Wilson, the author of Bryologia Britannica, to see them he first thought them to be sheep and then changed his mind for haycocks; when he could see what they really were he declared that never in his life had he seen such gigantic moss tufts.

Spruce added some forty-eight species to the British moss flora including Myrinia pulvinata from near York (1841) and Platydictya confervoides (formerly Amblystegium sprucei) from near Winch Bridge, Teesdale (1843). He wrote several papers on Yorkshire bryophytes. The most notable of these is perhaps that on the bryophytes of Teesdale: Spruce was one of the first to draw attention to the great floristic interest of this now famous area.

When working at York Spruce began his voluminous correspondence with bryologists and other botanists in several countries, many of whom he came to know personally later on. Among them were Borrer, Bruch, W.J. Hooker, Mitten, Montague and Sullivant. An important contact for his future work was Dr Thomas Taylor who invited him to stay for four weeks in his home at Dunkerron near Kenmare in Ireland. While there Spruce was introduced to the wonderful hepatic flora of Kerry and visited Cromaglan Mountain which he described as ‘a paradise of mosses’ but as the weather was bad and he had a severe cold he could do little field work, though spent many useful hours studying mosses and hepatics in Taylor’s large herbarium. In 1849, after Taylor’s death, Spruce went to London to supervise the sale of his collection by auction. It was bought by a wealthy American and his bryophytes are now in the Farlow Herbarium of Harvard University.

In the summer of 1844 the Collegiate School at York closed and Spruce found himself without a job. He was determined to find employment as a botanist, if possible abroad. His botanical friends who were well aware of his great abilities, made various suggestions. Sir William Hooker proposed that he should go on a collecting expedition to Spain, but enquiries indicated that the country was so disturbed that travelling might be dangerous and there would be difficulties in sending collections home. In the end Spruce, attracted by their reputation as a good area for mosses, decided to go to the French Pyrenees. He left England in May 1845 and returned in April of the following year. He had collected over 300 species of higher plants and numerous bryophytes, of which seventeen were new to science as well as many not previously recorded from the Pyrenees. The proceeds of selling exsiccatae were more than enough to repay a loan from William Borrer. Moreover, a year of working mainly in the open air had much improved his health and convinced him that his physical stamina was sufficient for an arduous collecting expedition abroad.

He now set his face against returning to the teaching profession (or entering the church, as one of his friends had suggested) and began to consider seriously the possibility of botanical exploration of the Amazon. He was encouraged to think this feasible by Sir William Hooker, then Director of Kew and by the zoologists Bates and Wallace, both of whom set out on collecting expeditions in South America in 1848. It was late in that year that Spruce finally decided to follow them. During a few months making preparations at Kew and the British Museum he met Robert Brown whose plant descriptions he regarded as models. He left England in June 1849 and arrived at Pará (now Belém) in July.

At Kew George Bentham had undertaken to sort the collections and make up sets of exsiccatae on condition that Kew kept the first set. He also promised to name all the previously described species and take a share in the work on the others. When the first consignment of specimens arrived Bentham and his assistant Professor Daniel Oliver were delighted both with their quality and their great scientific interest. Oliver wrote to Wallace:

‘Mr Spruce’s specimens were most carefully collected, dried and packed, extraordinarily so, considering the difficulties of all kinds with which he has had to contend; and what was of special value, they were accompanied by beautifully legible labels giving precisely the information as to locality, habitat, habit etc., required to supplement the dried specimens.

‘The collections were specially rich in arborescent species, the obtaining of which must often have been of considerable difficulty.’ Few of those who have collected plants in the tropics in recent times have been able to equal the quality of Spruce’s Amazon collections.’

After arriving at Belém Spruce spent some months working in the neighbourhood. Fortunately some forests in the area have been preserved and many of the species he collected can still be seen growing perhaps in the exact localities where he found them nearly 150 years ago. In 1849 Spruce went further afield and mapped and collected in the basin of the previously unexplored Rio Trombetas which joins the Amazon from the north near Obidos. He spent July and August of the next year at Obidos and sailed up to Manaus in October. In 1853-54 he explored and mapped the Rio Negro and some of its tributaries which proved to be botanically one of the most interesting parts of Amazonia, and went via the Casiquiare to the upper Orinoco.

In March-June 1855 he took ship from Manaus to Tarapoto in Peru. He then explored some of the Amazon headwaters, the Huallaga, Pastasa and Bombonasa rivers which were particularly difficult and dangerous to navigate. He arrived at Quito in the Andes of Ecuador in 1857.

In 1860 the Government of India asked Spruce to collect seeds and living plants of the Peruvian ‘Red Bark’ (Cinchona) because they were concerned about the supply of quinine, which was essential for safeguarding the health of the Indian army. Spruce was able to collect these in the rain forest below the volcano Chimborazo. He took great trouble in packing and despatching the material. It arrived safely and plantations were established in the Nilgiri Hills and elsewhere in southern India. Unfortunately as the plant is adapted to ever-wet rather than seasonally dry conditions, these were not permanently successful and were later replaced by plantations in Java.

On 24 April, 1860, while at Ambato, Ecuador, Spruce had a stroke and awoke to find himself partly paralysed in the neck, back and legs. ‘From that day forth I was never more able to sit straight up or walk about without great pain and discomfort. For a while he struggled on with his collecting, but the following year he had another disaster: owing to the failure of a business firm in Guayaquil in which they were deposited he lost most of his savings. This left him almost destitute and he was obliged to sell some of his books. After two more years on the coast of Ecuador and a further period in Peru he found it impossible to work and decided to return to England.

So ended his years of travel. It had been a heroic achievement. He had survived all kinds of dangers, illnesses and privations; for long periods he had worked alone except for Indian assistants. As well as collecting over 7000 species of vascular plants and large numbers of bryophytes, lichens and fungi, he had added enormously to scientific knowledge of Amazonian botany. The high quality of the specimens he sent home and the scrupulous care with which they were labelled and annotated was maintained to the end. In Bentham’s opinion it was the greatest contribution to tropical botany since the work of Humboldt half a century earlier. Spruce’s contribution involved not only botany: he added much to the ethnography of the Anierindians and his accurate mapping of little known parts of Amazonia was recognized by his election as Honorary Member of the Royal Geographical Society in 1866.

Spruce arrived in England in May 1864 almost penniless and in very poor health — the effects of his stroke were not his only trouble. Luckily he did not lack friends and some of them were influential. Thanks mainly to the efforts of Clements Markham, secretary of the Royal Geographical Society, he was awarded a civil list pension of £50 a year in 1864 and in 1877 this was supplemented by a further £50 from the Government of India in recognition of his work on Cinchona. It was not until four years after his return that the bowel trouble which had plagued him was correctly diagnosed and after this his health somewhat improved, but until the end of his life he had many afflictions to contend with. At one point he decided that he would have to give up microscope work, but fortunately he was able to return to it. Much of his writing had to be done in an easy chair with a large book on his knees as a table — for this reason many of his letters are written in pencil.

In order to concentrate on his large collection of hepatics Spruce decided to entrust the working out of his mosses to William Mitten, instead of dealing with them himself, as he would have preferred to do. Soon after arriving in England he stayed with Mitten at Hurstpierpoint in Sussex sorting out the moss collection and making up sets of his Musci Amazonicae et Andinae for distribution (see Spruce 1860).

After 1864 Spruce published some twenty-seven papers, but not all were on bryophytes. His most important non-bryological work was his classical account of the palms of the Amazon (1870), but his greatest work, which in the opinion of J.D. Hooker is his ‘crowning one and will ever live’ is the Hepaticae Amazonicae et Andinae (1884-1885). This was written under great difficulties. It appeared in the Transactions of the Botanical Society of Edinburgh in two parts. As the reader is informed inside the cover of Part 1, Spruce had intended to include a short, mainly geographical introduction, but this was never written. Plates I-VIII were drawn by his friend Robert Braithwaite (author of The British Moss-Flora 1887-1905) and the rest by George Massee, the mycologist who was also a friend of Spruce. This was probably because Spruce, though a good draughtsman himself felt that after his stroke he was no longer able to make sufficiently accurate drawings (see Richards, in preparation).

Spruce’s views on the classification of the Hepaticae had been earlier set out in his papers on Anomoclada (1876) and Cephalozia (1882). He had long since been sympathetic to Darwin’s views on evolution. After revising his South American Plagiochilas he wrote to Stabler (1871), ‘The result has been to make me more Darwinian than ever’. He went on to say that if we had all the forms of a genus which had ever existed, as well as those now existing, we could not define a single species and ‘could trace the unbroken pedigree of every form’.

After returning to Yorkshire he lived in lodgings at Welburn from 1867 to 1876 when he moved to the cottage in Coneysthorpe which now bears a plaque in his memory. He never married and was cared for by a devoted housekeeper and a girl attendant who acted as messenger. Stabler gives a good idea of his personality. He was no narrow specialist. He loved music and literature. He carried the works of Shakespeare with him on his travels. He took a lively interest in life around him. When the Duke of Argyll visited him at Coneysthorpe, they chatted for two hours on Spanish and Russian politics as well as on natural history and the undulatory theory of light. Even when ill and in pain he enjoyed a good joke and made puns. He was very methodical and numbered his notes so that he could instantly turn up a given topic. According to Wallace who had spent some time with him in Amazonia, he was as orderly in his work in the forest as in his cottage in Yorkshire. Above all he had a great capacity for friendship, took much trouble to help colleagues, and was always kind and sympathetic. He died of influenza in December 1893 and is buried beside his parents in the churchyard at Terrington close to where he was born.

This sketch of Richard Spruce’s life and work is largely based on Alfred Russell Wallace’s ‘Biographical introduction’ to Spruce’s Notes of a botanist on the Amazon and Andes (1908) and Stabler’s obituary (1894). There are several other obituaries, but Stabler’s is particularly valuable because he was a lifelong friend, had been at school with Spruce at Glanthorpe and was also a bryologist. It is strange that there is no full-length biography of such a remarkable man.


Braithwaite R (1887-1905). The British Moss-Flora. 3 vols. London: Reeve.

Richards, P.W. (in preparation). Two unpublished letters from Spruce to Braithwaite about the illustrations to Hepaticae Amazonicae et Andinae. Spruce Conference, York. Sept. 1993. Linnean Society.

Spruce, R (1850). The mosses and hepaticae of the Pyrenees. Trans. Proc. Bot. Soc. Edinb. 3: 103-216.

Spruce, R. (1860 [’1861’]). Mosses of the Amazon and Andes. J. Linn. Soc. (Bot.) 5: 45-51.

Spruce, R (1876). On Anomoclada, a new genus of Hepaticae, and its allied genera Odontoschisma and Adelanthus. J. Bot. 14: 129-136, 161-170, 193-203, 230-235.

Spruce, R. (1882). On Cephalozia, its subgenera and allied genera. Privately printed, Malton, Yorks.

Spruce, R (1884-85). Hepaticae Amazonicae et Andinae. Trans. Proc. Bot. Soc. Edinb. 15: i-xi, 1-590, pls. l-XXII.

Spruce, R (1908). Notes of a Botanist on the Amazon and Andes (ed. and condensed by Alfred Russell Wallace, with a biographical introduction). 2 vols. London: Macmillan.

Stabler, G. (1894). Obituary notice of Richard Spruce, Ph.D. Trans. Proc. Bot. Soc. Edinb., 20, Session LVIII, Feb.1894.

Sources of further information

Desmond, R(1977). Dictionary of British and Irish Botanists and Horticulturists. London: Taylor & Francis.

Gillispie, C.C. (ed) (1975). Dictionary of Scientific Biography. New York: Scribners.

Seaward, M.RD. (1980). Two letters of bryological interest from Richard Spruce to David Moore. Naturalist 105: 29-33.

Dr S.R. Edwards (University of Manchester): Spruce in Manchester.

The substantial holdings of Richard Spruce (1817-1893) material, at Manchester Museum Herbarium (MANCH), seem to have been one of the best kept secrets about this remarkable Yorkshireman who died one hundred years ago. Not only do we have Spruce’s own large personal herbarium, plus (and including) sets of his Hepaticae Spruceanae: Amazonicae et Andinae, his Musci Amazonici et Andini, and substantial lichen collections, but we also have a collection of his letters and maps and other documents which may interest Spruceologists more than bryologists. These all total about 1 6,500 items.

The Spruce material came to Manchester mostly in 1919, nearly 26 years after his death. Matthew Slater was Spruce’s botanical executor and he had inherited Spruce’s massive personal herbarium; when Slater died, it was W.H. Pearson who ultimately effected the transfer to Manchester Museum.

The large Spruce collections at Manchester Museum number over 16,500 items, consisting of:

  • his own personal herbarium of liverworts: 8,264
  • additional liverworts such as distributed sets:~ 700
  • his own personal herbarium of mosses:5,000
  • additional mosses in distributed set:289
  • his own personal herbarium of lichens:2,000
  • documentation such as letters and maps:~ 300

The liverworts, which form the bulk of the Spruce material, are largely from Spruce’s fifteen years in South America (June 1849 to June 1864), but also from his year in the Pyrenees (April 1845 to April 1846) and from elsewhere, and also include specimens collected by others. The mosses are more or less equally divided between British and non-British collections, and the lichens are mostly from the Pyrenees. There are also a few flowering plants and ferns. The letters to Slater in effect form a diary of Spruce’s last thirteen years; Slater was Spruce’s friend, factotum and confidant, and the letters make fascinating reading, both from a social and historical standpoint, and also for any bryologist interested in perceptive and detailed observations by one of the world’s greatest hepaticologists. The maps include hand-drawn examples by Spruce, including a finely detailed map of the River Trombetas, with compass bearings and lines of latitude and longitude. It appears to have been drawn by Spruce from first principles. The caption explains how five points were fixed by astronomical observation and the remainder by compass bearings, and how he ascended the river in 1849.

The following data give an indication of the significance of the Spruce liverwort collections at Manchester Museum:

  • 2,000,000 estimate of total plant collections;
  • 34,346 total liverworts;
  • 8,971 all Spruce liverworts (including personal herbarium and Hepaticae Spruceanae);
  • 924 Spruce liverworts designated TYPE (including holotypes, isotypes, lectotypes, isotypes, n. sp., sp. n., etc.); about 200 further packets have been designated TYPE, etc. since data were input to the database, although there may be some overlap;
  • 8,264 Spruce’s personal herbarium accessed in 1919.

We know these figures because comprehensive data from all of our liverwort collections (as well as from our Foreign Flowering Plants, etc.) are on a computer database. The database had only been available for editing and manipulation for about three weeks before the B.B.S. Autumn meeting at Ripon in September 1993; improved search criteria applied since then have revealed over double the number of Spruce specimens mentioned at that time.

This report of the Ripon lecture has been substantially updated in light of subsequent work on the Manchester material. A comprehensive account is given in the chapter: Spruce in Manchester: Manchester Museum Herbarium (with an Appendix on Manchester City Library by Professor Brian W. Fox), in the forthcoming volume Richard Spruce (1817 – 1893), Botanist and Explorer, to be edited by Prof. M.R.D. Seaward and published jointly by the Royal Botanic Gardens, Kew and the Linnean Society.

Mr A.R. Perry (National Museum of Wales): The early embryology of the British Bryological Society.

Before the foundation of the Moss Exchange Club in 1896, several important bryological books were published in Britain. Notable among these were Dillenius’ Historia Muscorum of 1741, Turner’s Muscologiae Hibernicae Spicilegium of 1804, Hooker’s British Jungermanniae (1816), and Hooker & Taylor’s Muscologia Britannica of 1818, 2nd edition 1827 with Wilson’s revision of 1855. In addition there were several, more popular, publications which gave impetus to the study of mosses, for example Stark’s A Popular History of British Mosses (1854), Berkeley’s Handbook of British Mosses (1863, 2nd ed. 1895), Tripp’s British Mosses… (1865, 2nd ed. 1888), Hobkirk’s A Synopsis of the British Mosses (1873) and Fry’s British Mosses (1892). The liverworts fared rather less well:

Cooke’s Easy Guide to the Study of British Hepaticae… (1865), Carrington’s British Hepaticae... (1874-75) and Cooke’s Handbook of British Hepaticae ( ‘1894’) were all that were easily available to the aspiring hepaticologist in the latter half of the last century, and Carrington’s work, the best of the lot, was never completed because of illness. Besides these, a few specimen books were available during the century: for the mosses Flintoft’s Specimens of British Mosses in the English Lake District (Ca. 1830), George Gardner’s Musci Britannici, or Pocket Herbarium of British Mosses (1836) and William Gardiner’s Twenty Lessons on British Mosses (1844, 1846), second series (1849); for the hepatics Mclvor’s Hepaticae Britannicae, or Pocket Herbarium of British Hepaticae… (1848). But as the century drew to a close a stimulus was required to get the ailing subject back to good health.

The Revd C.H. Waddell, Rector of Saintfield and (later) Grey Abbey, Co. Down, realizing the lack of progress in bryology, placed an advertisement in the Journal of Botany in 1896. Headed ‘EXCHANGE CLUB FOR MOSSES AND HEPATICAE’ it starts: ‘While much has been done for the study of phanerogams by means of clubs for exchanging, recording, and naming specimens, I believe the want of such a society for mosses hinders the advance of bryology. If one could be established … it would prove of great assistance to beginners…’ Offers of support from a number of friends had already been received, and he thought that at least thirty active members would be required to make such a society successful. There was an immediate response from H.N. Dixon (whose The Student’s Handbook of British Mosses was published the same year) by letter to the Journal of Botany. Dixon thought an Exchange Club would serve a very useful purpose, but was concerned that it might tend towards the extermination of our rare species. In reply, Waddell said the following rule had been drawn up for the Moss Exchange Club: ‘Great care should be taken not to injure or exterminate any rare or local species. If a plant only occurs sparingly, not more than one or two specimens should be taken … Localities near towns or where there is any danger of a rare species being exterminated should not be too definitely published’ — a rule that we still observe.

Twenty-three members enrolled in 1896 having paid the annual subscription of one shilling, and Waddell acted as Secretary, Treasurer and Distributor of specimens that were submitted. Some of the new members who joined are well-known names: besides Waddell and Dixon there were W.E. Nicholson, S.M. Macvicar, W. lngham and Miss E. Armitage all with considerable biyological knowledge and eventually leaving their undoubted mark on British bryology.

Waddell published his Annual Reports in the Moss Exchange Club Annual Reports (1896-1922). In his Report for 1897, one year after the Club’s formation, he wrote:

‘During the past Winter and Spring many enquiries have been received as to the work of the Club & copies of Rules sent out which has involved much correspondence, and a large expenditure on postage. … Other enquirers have not seen their way to join us for two reasons 1st the majority being beginners & not having any stock for exchange or considering that the Club would not be of much help to them in naming their finds prefer to wait till they have made further advance in the study. 2nd Some residing outside the British Isles find that the Society offers no opportunity for the exchange of foreign or continental plants.’

Waddell suggested the formation of three lists of names, 1) those who wished to exchange British for non-British mosses, 2) a list of helpers who would assist beginners, and 3) a list of those wishing to receive help in naming their plants. Later in the same Notebook he reported that Dixon had sent him some ‘Notes on Mosses’ that had been sent to him in the Exchange and had made a suggestion (which Waddell proposed to carry out) that these notes should be circulated to all members who were to be invited to add notes and criticisms and to ask questions either on the plants sent them or generally on any matter of real interest to the Club. Waddell continued: ‘Of course each note must be signed; and I hope the experiment of the “Note Book” may prove a success. It will go on a short circuit of about 6 and then back to me; first to the largest and most important contributors also taking locality into consideration.’ Thus were born the Circulating Notebooks in which members could read comments about the specimens that had been submitted for exchange.

Evidence for the immediate success of the Club is that in 1897 Waddell reported that 24 members had contributed 2163 Mosses and 104 Hepatics … as well as 28 plants sent in to be named, 2295 in all.

It is interesting to examine the list of helpers that emerged from Waddell’s request. There were seven; six of them, including Dixon and Nicholson, said they would be willing to help with ‘mosses only’. Miss Armitage, the seventh, said ‘mosses & hepatics, not critical’. It is clear from these responses that hepatics had been little studied and were consequently very scantily known — a inevitable reflection of the inadequate literature that had been published up to that date. Pearson’s two volume work The Hepaticae of the British Isles (1899-1902), which should have stimulated the study of hepatics was produced at such an exorbitant price ( £11.2s.6d with coloured plates; £7. l0s.0d uncoloured) that it was put beyond the pockets of most people (the equivalent in today’s prices for 11 gold sovereigns is something in the region of £750!); and it was not until Macvicar’s timely and brilliant The Student’s Handbook of British Hepatics was published in 1912 at 18s.6d. that hepaticology in the British Isles took off.

Meanwhile the circulating notebooks became a platform from which members could air their views on various bryological topics. Waddell, always helpful, advised strongly and kindly on the quality of the specimens sent in, and there followed a lively debate on labelling, folding of packets, and the formation of a personal herbarium, with many correspondents sending in descriptions of the way they prepared packets, labelled and stored them. Revd S. Gasking finally drew the arguments together in a wry comment ‘We all have our different ideas … & we will stick to our own methods notwithstanding this controversy.’

In 1897 E.C. Horrell, though not yet a member, wrote to Waddell telling him that he had started compiling ‘as exhaustive, a list as I can of the existing lists of the Mosses found in the 112 [ vice-counties’, and inviting help. This led to the eventual production of the first editions of the Census Catalogues of Hepatics in 1905 and Mosses in 1907 of the whole of the British Isles, oddly with the hepatic catalogue preceding the one for mosses. Horrell also became involved with helping beginners by setting up a Junior Section. Waddell reported in the 1900 Notebook that this was ‘doing well & Mr Horrell has enrolled over 30 members’. This Section came to have its own Annual Report, but on a smaller scale than its parent.

The Notebook for 1901 was 87 pages long and the various controversies and discussions continued; but not for long. The circulating Notebooks ended after 1903 when Waddell decided that the Club was flourishing well enough and he needed a rest from the arduous circulation that he had nobly instigated and the onerous duties of Secretary, Treasurer and Distributor. The Annual Reports carried on, however, and continued to have succinct versions of the notes on specimens submitted but hardly any other material; they were circulated to all members from 1896 to 1922.

The circulating Notebooks had been an important mouthpiece for those developing interests in bryology, an outlet where ideas and problems had been put forward, mulled over, spat out and reconstituted. Their demise heralded the formalization of British Bryology which was now on a firm basis, with the Moss Exchange Club having among its membership many members rapidly gaining confidence in bryology. The Club and its Junior Section continued to flourish until 1923 when the two amalgamated to form the British Bryological Society that we know today.

Mr. R. Stevenson (Kings Lynn): An amateur in the tropics.

A selection of holiday slides was used to illustrate the basic concept that, for many amateurs, the best opportunities for getting to, and collecting in, the tropics are on package holidays arranged by so called ‘Adventure Holiday’ companies (e.g. Explore; Exodus; Guerba). Through these companies one can travel to places which are relatively off the beaten track, and which are certainly likely to be bryologically underworked.

Most of these holidays involve rather a lot of travelling, often using a variety of means of transport. Space is often at a premium, so luggage needs to be minimized, in order to maximize space for collections. Collecting also has to be done swiftly and efficiently, since suitable opportunities are often rather brief, e.g. lunch stops by roadsides.

In order to cut costs many of these trips involve group members in helping out with various tasks, such as food preparation, luggage loading, shopping, etc. Some thought needs to be given to which tasks to volunteer for: gathering firewood, for instance, offers opportunities for bryologizing which scraping potatoes does not.

Basic equipment used is the same as at home, though collecting packets need to be strong where there is any great danger of heavy rain or persistent damp. A string bag is useful for suspending specimens in, to dry out. A major problem is always locating oneself: maps should be bought beforehand if possible (e.g. from Stanford’s). (A useful tip is to take a colour photocopy of those bits of the map which are going to be most used, and then laminating them with plastic, for use in the field.) A watch with a built-in altimeter is a useful tool in places where maps are inadequate.

The tropics encompass more than just rain forests, or montane areas, and it was suggested that collecting in the less obviously exciting dry, or urban and suburban areas, might be where amateurs are likely to make the most useful contributions.

Dr Harold Whitehouse (University of Cambridge): A presentation of stereoscopic bryophyte photographs.
Prof. B. Crandall-Stotler (Southern Illinois University, Carbondale): Apical organization, gametophyte ontogeny and phylogenetic implications in the moss Fissidens.

Fissidens differs from the vast majority of mosses in possessing uniquely constructed 3-parted leaves that are, at least in part, vertically inserted upon the stem in a distichous arrangement. Early anatomical studies of Lorentz (1864) and Leitgeb (1874) demonstrated that these characters are correlated with a distinctive type of apical organization that centres around a lenticular apical cell with two segmenting surfaces. These authors also both described the early stages of leaf development, but their interpretations of ontogeny differed. Various authors since have provided contrasting views regarding the phylogenetic significance of these characters, but without clarifying the morphogenetic patterns involved. Using a combination of serial paraffin sectioning and SEM techniques, this reinvestigation was consequently undertaken.

In protonemally formed buds and young branches of F. taxifolius, the apical cell is initially obovoidal and spirally segmenting, just as it is in other mosses. Divergence angles between the first formed derivatives are 137°, but this angle is quickly increased so that after two complete spirals of segmentation, a 1800 divergence angle and lenticular apical cell geometry are established. In contrast to other mosses, the division wall that separates the derivative from the apical cell is deposited parallel to the segmenting wall of the apical cell, rather than at an oblique angle to it. As a consequence, the free surface of the derivative is symmetrical in outline and the derivative lacks the anodiclcathodic polarization that characterizes the asymmetric derivatives of other taxa.

As in other mosses, two divisions, the first periclinal and the second anticlinal, generate a single leaf initial from each derivative. Two sequential oblique divisions in this initial then form the leaf apical cell with two segmenting surfaces. In most species of Fissidens, the first of these divisions extends towards the dorsal surface of the transversely inserted initial so that the first formed basal segment is dorsal in position. In mosses with spirally segmenting apical cells, the same cell is always anodic so that it is dorsal on one side of the stem and ventral on the other (Berthier, 1973).

After the 3-celled stage, the mitotic spindle rotates towards the ventral side of the stem, causing the next dorsally positioned cell to be slightly displaced from the transverse plane. This gradual rotation of the spindle continues through the next three division cycles of the leaf apical cell so that by the 7-celled stage the leaf apical cell is reoriented at a 90° angle and is segmenting cells in a horizontal rather than a dorsi-ventral plane. The vaginant lamina is produced from the original basal cells, with the dorsal lesser lamina of the vaginant lamina emanating from divisions in the first formed basal cell. The vertically oriented superior and inferior laminae develop from the reoriented segments. These observations are in complete accord with the interpretations of Leitgeb (1874) and suggest that the unique form of the Fissidens leaf is controlled by modified patterns of spindle orientation. These patterns can be altered by the addition of hydroxy-1-proline which, when supplied at 10-6M concentration, inhibits the production of the superior and inferior laminae, and kinetin, which at 10-5M concentration, increases the size of the lesser vaginant lamina.

Studies currently in progress on F. asplenioides, a species which forms the lesser vaginant lamina dorsally on one side of the stem and ventrally on the other, suggest that primitive species of Fissidens may possess apical organizations that are intermediate between those of

typical mosses and the lenticular system of most Fissidens species, and support the hypothesis that Fissidens is an ontogenetically more complex, phylogenetically derived, taxon.


Berthier, J. (1973). Recherches sur Ia structure et le développement de l’apex du gamétophytes feuillé des mousses. Rev. Bryol. Lichénol. 38: 421-551.

Leitgeb, H. (1874). Zur Kenntnis des Wachshums von Fissidens. Denkschr. Kaiser!. Alcad. Wiss., Wien. Math. -Naturwiss. Kl. 69: 47-69.

Lorentz, P.G. (1864). Studien fiber Bau und Entwicklungsgeschichte der Laubmoose, pp. 1-36 + 4 p1. In PG. Lorentz (ed), Moosstudien. Leipzig: Engelmann.

Dr Philip E. Stanley (Cambridge): The cumulative index to BBS publications.

The British Bryological Society and its predecessor the Moss Exchange Club have recorded their activities in the Moss Exchange Club Reports (1896-1922), Reports of the British Bryological Society (1923-1945), Transactions of the British Bryological Society (1947-1971), Journal of Bryology (1972-present) and the Bulletin of the British Bryological Society (1963-present). There have also been other occasional publications including volumes containing the proceedings from meetings held in conjunction with other societies.

Although indexes have appeared for each volume of the Transactions and the Journal, there has not been a cumulative index embracing all of the Society’s publications and some years ago I agreed to prepare such an index. A short note was published (Stanley, PB. (1992). Cumulative index to BBS. publications. Bull. brit. Bryol. Soc. 59: 3 1-32) recently which outlined the status of the project. Since then I have reworked the entries to reflect suggested revisions both in style and items for inclusion. Further, the proceedings of meetings held in conjunction with other societies have now been included.

A draft of the index was exhibited at the conversazione held in conjunction with the AGM in 1992 and members saw that its format is similar to that used in the Journal, namely short phrases or at least several words. Thus it differs from some other indexes in which entries consist of a single or at most a few words. The index includes group headings such as Floras and Checklists, Keys to Genera and Species, Species New to the British Isles, Reports of Meetings (separate lists by date, place and kind, e.g. AGM), Membership Lists. It is hoped these will assist the user in locating useful information. In the main, no attempt has been made to cross reference taxonomic synonyms and if a paper refers to Hypnum cuspidatum it will be cited under that name and not under Acrocladium cuspidatum or Calliergon cuspidatum or Calliergonella cuspidata.

The inclusion of citations is now finished and they are now in a pseudo-alphabetic sequence as far as the computer is concerned. There remains the not inconsiderable task of editing these entries to produce a formal index. This will be published from camera-ready material which will avoid the need for a second proof-reading. Each member will receive a copy which will have a page size similar to recent issues of the Journal.

Field excursion to Hackfall Wood, 19 September 1993

The field excursion on Sunday 19 September was to Hackfall near the village of Grewelthorpe about 6 miles NW of Ripon. Hackfall is a semi-natural deciduous wood on the steep banks of the R. Ure. In recent years, however, it has suffered extensive loss of elm trees and parts of it are therefore more open than presumably they were previously. The wood is owned by the Woodland Trust, to whom we are indebted for information and access. The diversity of habitats within the wood and on the river banks ensured a rich flora and over 140 species were recorded on the day.

The underlying rock is Millstone Grit and there are low crags in the higher parts of the wood. The more acidic of these produced small quantities of Bazzania trilobata, Barbilophozia attenuata and Cephalozia lunulifolia but in other places there was evidence of slight base enrichment, with such species as Eucladium verticillatum and Leiocolea turbinata. Grit boulders and stones occurred in various places at lower levels throughout the wood and these produced Jungermannia pumila, Hygrobiella laxiflora, Scapania umbrosa, S. nemorea, Blindia acuta and Heterocladium heteropterum (some of these only in very small quantity).

The gully formed by the stream flowing through the centre of the wood was obstructed by logs and much overgrown but produced some additional species, including Metzgeria conjugata and Fissidens crassipes. There were fine patches of Hookeria lucens c.spor. in various places on the woodland floor and Nowellia curvifolia on rotten wood. Plagiothecium laetum and Plagiochila britannica were also reported.

A notable feature of the wood is the presence of a number of calcareous springs, often with abundant Cratoneuron commutatum. Some of these have extensive masses of tufa, on which the Eucladium and Leiocolea were plentiful, along with Jungermannia atrovirens and a little Tortella tortuosa. Rhynchostegiella teesdalei was found on wet grit rocks in a runnel where the water was probably calcareous.

The epiphytic flora was not very rich, although there were some fine tufts of Dicranum montanum on sycamore and other tree boles. Metzgeria fruticulosa, M. temperata, Zygodon conoideus and Orthotrichum pulchellum were also recorded.

The R. Ure has a marked flood zone and its banks had well developed communities of riparian bryophytes. Dichodontium flavescens, Fissidens rufulus, Barbula nicholsonii, B. spadicea, Oxystegus sinuosus and Schistidium alpicola were mainly confined to boulders, while Porella cordaeana, Radula complanata and Leskea polycarpa occurred about tree bases. Other species, such as Homalia trichomanoides, were indifferent to substrate. Also on riverside rocks, and presumably benefiting from the calcareous river water, were Anomodon viticulosus, Rhynchostegiella teesdalei and Jungermannia atrovirens. Willow bushes on a small island in the river, and therefore well illuminated, produced Orthotrichum rivulare and fittingly, on the anniversary of the great man’s death, a little O. sprucei. The latter was also reported from alder.

Tom Blockeel