European Herbertus and the 'Viking Prongwort'

Article he genus Herbertus is widely and Norwegian Herbertus borealis. Because of considered to be taxonomically this Norwegian connection, we chose to include ‘difficult’, largely because the plants all four European species: (1) Herbertus aduncus have remarkably few clear-cut (Dicks.) Gray subsp. hutchinsiae (Gottsche) R.M. characters which can be used to dis- Schust. reported from Scotland, England, Wales, Ttinguish species. In addition, plants are very Ireland and Norway; (2) Herbertus stramineus rarely fertile and no sporophytes have ever been (Dumort.) Trevis. from Scotland, England, found on Herbertus in Europe. Thus almost all Wales, Norway, Faeroe Islands and Iceland; described differences are in leaf features, and (3) Herbertus sendtneri (Nees) Lindb. from the some of these are very hard to quantify clearly, Austrian and German Alps; and (4) Herbertus for example shape and orientation of the lobes. borealis Crundw., described in 1970 from DNA barcoding is a modern technique which it Scotland and Norway (Crundwell, 1970). We, has been claimed is especially useful in character- along with several other bryologists, collected poor groups like Herbertus. At the Royal Botanic Herbertus in Ireland, England, Wales, Scotland Garden, Edinburgh we have been testing this and Norway and very recently Austria, and were approach on British liverworts (Long et al., also given material by other people, including 2007), and for David Bell’s MSc project the genus Herbertus samples from China, the Himalaya and Herbertus was selected as a pilot study, because North America to test links or even synonymy of taxonomic uncertainties around the Scottish which had been suggested previously. European H. hutchinsiae. John Birks Herbertus and the ‘Viking prongwort’ David Bell & David Long review the difficult liverwort genus Herbertus, discussing how DNA barcoding is helping to resolve some of the issues, and has led to the discovery of a fifth European species. A key for identification of the five European species is provided. 2 FieldBryology No106 | Feb12 FieldBryology No106 | Feb12 3 European Herbertus European Herbertus History of the genus Herbertus Raddi (1818), Dumortier (1822) and Corda material now lost, but as later expertly detected did not distribute her Bantry material (probably (1828) were steadily describing new segregates by Proskauer (1962) Dickson definitely included due to insufficient quantity available) but rather 1. Herbertus in Europe of Jungermannia and these gained gradual in his description and illustration sporophyte- specimens from Killarney collected by the In the early 19th century, most leafy liverworts, acceptance. The last two authors were ignorant bearing plants which could only have come English botanist Benjamin Carrington. Not including those now in Herbertus, were placed of Gray’s work and one of Dumortier’s new from Canada (Long, 1979). Proskauer correctly until Evans (1917) was Herberta hutchinsiae in the portmanteau genus Jungermannia, largely genera was Schisma, synonymous with Gray’s fixed the type of Jungermannia adunca with a (Gottsche) A.Evans elevated to species rank, following the concepts of Hooker (1816) in his Herbertus. Schisma gained acceptance and Menzies collection from British Columbia. For though Schuster (1966) preferred to keep it influential British Jungermanniae. His thinking survived for half a century, although it was briefly almost 170 years up to 1962 it had been wrongly as a subspecies of Herberta adunca (subsp. was guided by the Linnaean tradition of replaced by Sendtnera Endl. which also included thought that J. adunca originated from Scotland. hutchinsiae). Ellen Hutchins (1785–1815) lived primarily using sporophyte characters to classify Mastigophora. Carruthers (1865) argued for Not all authorities accepted the name at Ballylickey near Bantry in County Cork and mosses and liverworts, in this case the remarkably resurrection of Gray’s genera (which had Jungermannia adunca, for example Smith & due to her fragile health took up field botany simple, uniform sporophytes of leafy liverworts: nomenclatural priority) and Lindberg (1874, Sowerby (1812) and Hooker (1816) considered and embarked on compiling a catalogue of ‘the plants that form the genus Jungermannia, 1875) was the first to agree, simply changing it to be a synonym of J. juniperina Sw. [now plants of the district (Bevan, 1984; Mitchell, however numerous, cannot be divided into other the gender of Herbertus to Herberta to replace Herbertus juniperoideus (Sw.) Grolle] from the 1999; Hutchins, 2003). She corresponded with genera by means of characters taken merely from Schisma. Lindberg’s spelling Herberta survived West Indies. Soon after 1800, Herbertus was Dawson Turner in England who encouraged the fructification’ (Hooker, 1816). However, for a century until nomenclatural conflict with collected from several new localities, such as Ben her studies and in return she supplied him with Hooker’s concepts became superseded both in a flowering plant genus Herbertia Sweet was Lawers by William Hooker in 1808 (BM) and specimens, including her Herbertus and others Britain and Europe as gametophyte characters pointed out, and following Florschütz & Grolle near Bantry in Ireland in 1810 by Miss Ellen such as Jungermannia (now Jubula) hutchinsiae. were increasingly brought in to define smaller (1975), Gray’s original spelling Herbertus was Hutchins (BM). These were both identified by These eventually came to Turner’s son-in-law genera. Samuel Frederick Gray (1766–1828) in reinstated. Hooker as J. juniperina, although in reality they William Hooker, who doubtless distributed her his Natural Arrangement of British Plants (Gray, represented two different undescribed species. liverwort duplicates to others such as Gottsche. 1821) was the first in Britain to break away, 2. Herbertus in Britain and Ireland Others such as Dumortier (1822) retained the Well into the 20th century, many authors, describing numerous new generic segregates, The first localized and dated collections of name adunca for British plants, though he placed notably Macvicar (1912), continued to recognize including Herbertus. This name, according a Herbertus in Europe were made by the it in his ‘new’ genus Schisma. Nine years later, Herberta adunca as the only species in Britain to Müller (1951–1954), commemorates the Scottish surgeon/botanist Archibald Menzies Dumortier (1831) described a second species and Ireland, although Macvicar segregated some British nobleman Thomas Herbert ca( 1656– (1754–1842) on Ben Nevis and Ben Lomond from Scotland, Schisma stramineum Dumort., mountain plants from Perthshire and Shetland 1733), the 8th Earl of Pembroke and 5th Earl in Scotland in 1778. These were annotated under another vague citation ‘in alpibus Scotiae’. as H. adunca var. alpina, now regarded as a of Montgomery. Herbert was not a botanist, but ‘Jungermannia juniperina’ by Menzies and later This is possibly based on Hooker’s Ben Lawers stunted form of Herbertus stramineus (Damsholt, was one of the patrons of the celebrated early identified as Herbertus hutchinsiae, and are material. Very few botanists accepted this name 2002). Müller (1951–1954) went to the other Italian botanist Pier Antonio Micheli (1679– preserved in Menzies’ herbarium (E, BM). Nine either, and so it was that the epithet straminea extreme and recognized three species from 1737), and one of his plates (Micheli, 1729, tab. years later, Menzies, as surgeon on the Prince remained effectively ‘lost’ until Proskauer (1962) Britain: Herberta hutchinsiae, from Ireland, 27) is dedicated to Herbert. of Wales, explored the northern Pacific and showed that it applied to the plant which many Wales, England and Scotland, H. adunca (syn. Gray’s names, however, were largely ignored collected Herbertus in British Columbia ‘NW had simply called Herberta adunca before that. H. straminea) from Wales and Scotland, and a both in Britain and Europe, partly through America’ in 1787. However, the first description In 1862, Carl Moritz Gottsche, from Altona third species, the North American Herberta Hooker’s influence, the lack of communication of a European Herbertus was not until 1793 by the in Germany, decided that Miss Hutchins’ tenuis A.Evans from Loch Assynt in Scotland of Gray’s work abroad and also because Gray Scottish botanist and nurseryman James Dickson Irish plants represented a taxon different from and Cwm Glas in Wales. Soon after, Jones used the masculine gender for his new genera, (1738–1822) as Jungermannia adunca, described ‘adunca’. He therefore, in Rabenhorst’s Exsiccatae (1958) considered that these plants were not H. rather than the traditional feminine. This was supposedly from the Scottish Highlands ‘in (Hepaticae Europaeae), distributed material tenuis but ‘merely slender forms of H. hutchinsiae’. considered improper – according to Dumortier alpibus Scoticis’, but without locality, date or under a new name Sendtnera adunca (Dicks.) However, Jones also mentioned that ‘an as yet ‘these are the names of men and not of plants’ collector (Dickson, 1793). The Scottish plants Gottsche var. hutchinsiae Gottsche. Although his undescribed species of Herberta has recently been (Evans, 1917). In Europe, bryologists such as may have been those of Menzies or his own new variety commemorated Miss Hutchins, he found on Beinn Eighe in Scotland’. 4 FieldBryology No106 | Feb12 FieldBryology No106 | Feb12 5 European Herbertus European Herbertus History of the genus Herbertus Raddi (1818), Dumortier (1822) and Corda

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