Grimmiology

GRIMMIA Hedw. (1801)

Introduction
Named originally by Ehrhart in honour of J.F.K. Grimm, physi­cian and botanist in Gotha, Germany († 1821), based on forms of Bryum apocarpum L. As under this taxon nowadays is under­stood Schistidium apocarpum (Hedw.) Bruch et Schimp., a lecto­type had to be chosen for the genus Grimmia, and accor­ding to Mår­tensson (195­6), this is Grimmia plagiopo­dia Hedw. Alt­hough predominantly occurring in the moderate zones, repre­sentatives of this cosmopo­lithic genus may be found in all parts of the world, from Alaska to the most sout­hern point of Chili, and from Siberia to South Afri­ca, though in tropic regions e.g. Hawaii and Indo­nesia, Grim­mia species only occur high up in the moun­tains.

Grimmia is a notorious difficult genus, and in all herbaria investigated, I found a considerable number of species misidentified. The American bryologist Geneva Sayre (1911-1992), who worked for many years on a monograph of the North American Grimmias, indicated in an original way these difficulties, as she said: “it contains an ambigua, a varia, a decipiens, a controversa, a revisa and at least two anomalas” (Richards, 1993). According to Frahm & Frey (1987), the genus Grimmia s.s. counts ± 150 spe­cies. However, the Index Muscorum (Wijk et al., 1962) com­prehends more than 800 published Grimmia taxa. Cause for this large number is the concep­tion of some 19th century bryologists that all acrocar­pous saxico­lous species should belong to the genus Grimmia, and they included Racomitrium, Ptychomitrium, Eucladium, even Seligeria and many Potiaceae.

Johan Zettersted (1785-1874)
Johan Zettersted was probably the first Grimmiologist. In 1861, he published ”Revisio Grimmiearum Scandinaviae”. In this work he recognized 20 species: Grimmia anodon,    G. pulvinata, G. fuscolutea, G. decipiens, G. funalis, G. trichophylla, G. elatior, G. donniana, G. longirostris, G. laevigata, G. ovalis, G. montana, G. alpestris, G. torquata, G. hartmanii, G. elongata, G. mollis, G. unicolor, G. atrata and G. incurva (nomenclature according to Greven (2003).

Leopold Loeske (1865-1935)
Leopold Loeske, a professional watchmaker, started his bryological career in 1896 with a paper on the moss flora of the Harz. He died in 1935 by a heart attack during a bryological excursion in, of all places, the Harz. He published many bryological papers but his most important work was ”Monographie der Europäischen Grimmiaceen”, (Loeske 1930). Since my first paper on Grimmia (Greven 1990), Loeske’s book was continuously on my desk, I read and reread all his detailed remarks about on the European species, in my opinion the best book on Grimmia ever published.

George Neville Jones
Jones (1933) treated the Grimmiaceae in the Moss Flora of North America, North of Mexico. He acknowledged with appreciation his indebtedness to Leopold Loeske. In this work, he recognized 72 species, varieties and forms for North America north of Mexico. The work is much more outdated than that of Loeske because Jones included in the Grimmiaceae genera such as: Schistidium (with a large number of varieties), Coscinodon, Braunia, Campylostelium, Glyphomitrium, Hedwigia, Ptychomitrium, Rhacomitrium, and Scouleria.

Hironori Deguchi
Deguchi (1979) revised the genera Grimmia, Schistidium and Coscinodon of Japan. He remarked that the Japanese flora included a large number of doubtful species that needed to be revised taxonomically. All taxa are described in detail, and drawed carefully. I have frequently consulted this excellent work, Deguchi recognized 15 species: Grimmia affinis, G. anomala, G. apiculata, G. atrata, G. brachydictyon, G. curvata, G. donniana, G. elongata, G. funalis, G. incurva, G. muehlenbeckii, G. olympica, G. percarinata, G. pilifera and G. subsulcata. Grimmia olympica has to be treated as Brachydontium olympicum. Grimmia brachydictyon is a endemic to Japan.

Akira Noguchi
A second, but less detailed treatment of the Japanese Grimmias was, as part 2 of the “Ilustrated moss flora of Japan”, elaborated by Noguchi (1988). He recognized the same species as Deguchi, but added three Schistidium species as Grimmia: G. apocarpa, G. maritima and G. subconferta. He deleted one doubtful record of Grimmia funalis and treated G. muehlenbeckii as a variety of G. trichophylla.

Tong Cao and Dale Vitt
A taxonomic revision and phylogenetic analysis of Grimmia and Schistidium in China was published by Cao & Vitt (1986). In this high standard work, the authors recognized 22 species: Grimmia affinis, G. anodon, G. apiculata, G. atrata, G. caespiticia, G. decipiens, G. donniana, G. elatior, G. elongata, G. handelii, G. incurva, G. laevigata, G. macrotheca, G. mammosa, G. montana, G. obtusifolia, G. ovalis, G. pilifera, G. pulvinata, G. sessitana, G. tergestina, and G. unicolor.

Henk Greven
In 1988, working on a thesis ”Changes in the Dutch Bryophyte Flora and Air Pollution”, (Greven 1992), I found on roofs of concrete bunkers three Grimmia species, new to The Netherlands: Grimmia orbicularis, G. ovalis and G. tergestina. During identification, I detected that the taxonomy and nomenclature of the genus Grimmia was a mess, with over 800 names in the Index Muscorum (Wijk et al. 1962).

Since Loeske (1930), no revision of the European species had been carried out.

I started with a revision of the Grimmia collections from B, BM, BP, DUIS, E, EGR, H, JE, GRO, ICEL, K, KRAM, L, LGHF, MADS, PC, RO, S, U, and Z. The result of this revision, combined with fieldwork in the Alps, Pyrenees, Tatra and other European mountainous areas, was published in ”Grimmia Hedw. (Grimmiaceae, Musci) in Europe” (Greven 1995). In this work, I recognized 41 Grimmia species. Subsequently, I started a world revision, examining the collections from AD, AK, AKU, ALTA, AMNH, BISH, BR, BRI, FH, C, CANB, CBG, CHR, INA, MEL, MELU, MEXU, MICH, NY, OTA, HO, MO, NSW, PERTH, PRE, UC, UWA and WELT.

Up to 2002, about 20.000 herbarium specimens, including hundreds of types, had been revised, and about 50 Grimmia-collecting trips were made in all continents. The results of the herbarium revisions, combined with about 2000 Grimmia samples, collected in the field, was published in ”Grimmias of the World” (Greven 2003). Apart from the two above mentioned books, about 50 papers on Grimmia, including synopsi of Grimmia in Australia, New Zealand and Mexico, were published.

An important contribution was the revision of Grimmia in North America, carried out by Roxanne Hastings and me, resulting in a chapter on Grimmia in the Flora of North America, Volume 27, Bryophyta, part 1 (Hastings & Greven 2007).
 
In the past 15 years, the following Grimmia species have been discovered and described:

Grimmia maido (Greven 1996)
Grimmia macroperichaetialis (Greven 1998)
Grimmia wilsonii (Greven 1998)
Grimmia mexicana (Greven 1999)
Grimmia indica (Goffinet & Greven 2000)
Grimmia nevadensis (Greven 2002)
Grimmia lesherae (Greven 2003)
Grimmia mauiensis (Greven 2003)
Grimmia maunakeaensis (Greven 2003)
Grimmia shastae (Greven 2003)
Grimmia milleri (Hastings & Greven 2007)
Grimmia texicana (Greven 2010)

Redescription of Grimmia fuscolutea Hook.
Greven (1996) published Grimmia sanii spec. nov. from South Africa, Drakensberg. In the protologue, the new species was contrasted with Grimmia flexicaulis Müll. Hal. (a synonym of G. fuscolutea Hook.), described from South America. Because Deguchi (1987) described G. flexicaulis with an erect, flexuose seta, smooth capsule, and undivided or occasionally cleft peristome teeth, characters that contrasted with G. sanii, G. fuscolutea was incorrectly redescribed.

Eva Maier
Maier & Geissler (1995) publiced ”Grimmia in Mitteleuropa: Ein Bestimmungsschlüssel”. The study was based upon revision of 1243 herbarium specimens. In this work, Grimmia alpestris was included as Grimmia sudetica, and Grimmia arenaria, G. austrofunalis,     G. crinitoleucophaea (= G. poecilostoma), G. lisae and G. triformis were not recognized.

Maier (2002a) studied 483 Grimmia trichophylla specimens, resulting in a paper with two new species: Grimmia dissimulata and Grimmia meridionalis. It is regrettable that she did not enter into the detailed discussions on the Grimmia trichophylla-complex by Loeske (1930) and Greven (1995). Maier sent me material of Grimmia dissimulata and Grimmia meridionalis. The first one has frequently been collected by me in Mediterranean countries, especially on the islands Cyprus, Corsica, Sardinia and Mallorca. So far, I ihad always identified these plants as forms of the variable G. lisae, but provisionally, I accept this taxon as a good species, because there are indeed differences wiith G. lisae. The second one is conspecific with Grimmia austrofunalis, a taxon not recognized for Europe by Maier. Muñoz (2006) synonymized G. meridionalis with G. trichophylla.

Maier’s taxonomic view is predominantly based upon the structure of the costa. According to Maier, you must start with good transverse sections of the costa before you can distinguish Grimmia lisae (6 guide cells) from Grimmia trichophylla, G. meridionalis and  G. dissimulata (4 guide cells). In G. meridionalis, the guide cells are distinctly elliptic, while these in G. dissimulata are more or less rounded. It is my experience, that this is not the way to identify Grimmia taxa. The quality of the cross sections (not correctly cutted sections can show rounded cells as elliptic), and the location, where these sections have been made, are important.

Further, Maier did not discuss the extreme morphological variability within the Grimmia trichophylla-complex. As in all taxa of this complex, leaf form, margins, cell pattern, and hair-points are extreme variable, why should the costa be excluded from variability. In my opinion, there are some more steady characters that must be used, to identify variable Grimmia taxa. However, because there have been many discussions on the Grimmia trichophylla-complex, first of all, the type specimens of the 8 Grimmia trichophyl-la subspecies and the 21 G. trichophylla varieties (Wijk et al. 1962), must be examined before new species can be described within this complex.

Maier (2002b) presented the revision of 597 Grimmia specimens collected by the Scottish bryologist David Long, on excursions in the Himalaya (Nepal, Sikkim, Bhutan and adjacent China: Yunnan and Qinghai). She also had included 140 herbarium specimens from ALTA, BM, G, E, GZU, H-Broth, H-SOL, NY, S, W, WU, Z and ZT. In this publication, the Himalayan endemic Grimmia ochyriana was identified as Grimmia atrata, Grimmia crinitoleucophaea (G. poecilostoma) was synonymized with Grimmia tergestina. Grimmia capillata was identified as Grimmia crinita, Grimmia arenaria was synonymized with G. doniana, and Grimmia redunca, a Himalayan endemic, was synonymized with Grimmia elongata.

During fieldwork in the Himalaya, I studied richly fruiting cushions of Grimmia ochyriana and Grimmia redunca, in respectively the Khumbu valley in Nepal, and along Tsongo Lake in Sikkim. The field observations showed that these species are not synonymous with Grimmia atrata (Greven 2008), respectively Grimmia elongata.

Maier (2009) republished the central European Grimmia species. In this paper, she recognized 32 European Grimmia taxa. Her concept of taxa deviates far from that of Muñoz & Pando (2000) and Greven (2003), therefore her remark: ”Die Auswahl der Typen stützt sich weitgehend auf die Arbeit von Muñoz & Pando (2000)” is not supported by the 39 European Grimmia taxa, recognized in this work. Although Muñoz and Greven both agree that: Grimmia arenaria, G. capillata, G. crinitoleucophaea (the correct name for Grimmia poecilostoma), G. mollis, G. reflexidens, G. teretinervis, and G. ungeri are good European Grimmia species, Maier rejects all of them, however, without any argumentation.

Jesùs Muňoz
Director of “Instituto Asturiano de Taxonomía y Ecologia Vegetal”
Apartamendo 8, Pravia, Spain.

In the Bryological Times No. 75, October 1993, Jesùs Muňoz presented a new research institute. The institute had as start 8500 plants (bryophytes and vascular plants), the bryophyte herbarium contained 5750 specimens world-wide, including 20 types, interesting specimens were duplicates of collections by Luisier, Mönkemeyer and Warnstorf. It is in my opinion rather questionable how a junior bryologist could obtain so many specimens and especially from where came all those types?

In August 1995, I visited him in Pravia. As a present, I brought with me my just published “Grimmia Hedw. in Europe”. During this visit, he told me that his institute was only established to receive Grimmia specimens from herbariums all over the world. I don’t know what happened but since my visit, he refused any contact and did not reply anymore my letters. 

The Missouri Botanical Garden has a website with ”A checklist of the Mosses”.
On this checklist, species treated in a monograph, revision, or desciptive flora are recognized by four stars. Muñoz was responsible for the the genus Grimmia. A look at this checklist, in february 2010, learned that all Grimmia species, described by Greven, are absent or not recognized as good species, because not any species has 4 stars. Grimmia maido (Greven 1996) is absent, Grimmia macroperichaetialis, G. wilsonii, and Grimmia mexicana have only 2 stars. Grimmia ochyriana, described by Muñoz from material, that he found intermingled in a herbarium sample of G. subdonniana from the Himalaya, was ranked with four stars, although Muñoz never revised Grimmias from the Himalaya.

A revision of Grimmia (Musci, Grimmiaceae) in the Americas: 1: Latin America
Muñoz (1999) published a revision of Grimmia in Latin America.
My comment on this revison can be summarized into three points:

1. The list of species is incomplete, the following species are absent:

  • Grimmia arizonae Renauld & Cardot ( considered conspecific with G. pilifera P. Beauv.).
  • Grimmia bernoullii C. Muell. (considered conspecific with G. ovalis (Hedw.) Lindb.
  • Grimmia herzogii Broth. (considered conspecific with G. longirostris Hook.).
  • Grimmia incrassicapsulis B.G. Bell (Chile, Townsend 97/207)
  • Grimmia lawiana J.H. Willis (excluded from Grimmia )
  • Grimmia mexicana H.C. Greven (considered conspecific with G. ochyriana J. Muñoz)
  • Grimmia orbicularis Wils. (Argentina, Townsend 97/246, Chile, Townsend 97/212)
  • Grimmia torquata Hornsch. (Mexico, Greven M163-165, Argentina, Townsend 97/228)

2. The list contains taxa that do not occur in Latin America :

  • Grimmia ochyriana J. Muñoz is a Himalayan endemic.
  • Grimmia ovalis (Hedw.) Lindb. Plants identified as G. ovalis are G. bernoullii C. Muell.
  • Grimmia pilifera P. Beauv. Plants identified as G. pilifera are G. arizonae Ren. & Card.
  • Grimmia crinitoleucophaea Card. (G. poecilostoma Card. & Seb.). This species is has never been recorded from Latin America.
  • Grimmia americana Bartr. This species has never been recorded from Latin America..
  • Grimmia ungeri Jur. Two Mexican samples, identified as G. ungeri, are G. montana Bruch & Schimp. This is shown in the drawings of G. ungeri and G. montana, that do not differ in any aspect.

3. The key is unworkable
The group of species with recurved margins are divided into:
24a: cross section of costa reniform (G. longirostris and G. lisae)
24b: cross section of costa semicircular, never reniform (G. pulla, G. pulvinata, G. reflexidens, G. pilifera,G. navicularis, G. molesta, G. bicolor, G. humilis, G. trichophylla, and G. austrofunalis).

However, the drawings of the costa of G. lisae do not differ from those of G. pilifera,
G. trichophylla, G. navicularis, and G. reflexidens shows even a costa reniform in cross section.

A World Synopsis of the genus Grimmia

Muñoz & Pando (2000) published ”A World Synopsis of the genus Grimmia” (Missouri Botanical Garden Press Vol. 83). In this work, they recognized 71 taxa. Because there are no species descriptions or drawings, the work cannot be titled as a synopsis, it is more or less a nomenclatural enumeration. Although they claimed to have studied all relevant literature on Grimmia, from the, at that moment 35 published papers on the genus Grimmia by Greven, only two are mentioned in the Literature Cited. On page 70, Muñoz wrote: ”Greven (1998: 397) blindly followed Dixon’s publication”, and on page 79 again: ”he blindly followed Dixon”.

Such remarks are not scientific and it is incomprehensible that a high standard institute as the Missouri botanical Garden published a scientific work with such personal remarks. The work is very incomplete, e.g. ”names with type material not seen”, number 204 taxa. In my opinion, the study is so full of taxonomical and morphological misunderstandings that it is rather useless, even misleading in many places.

The work can be conceived as a refreshment of the Grimmia chapter in the Index Muscorum however, with many gaps and traps. Study of this work revealed the following incorrect data:

  • Grimmia abyssinica is good species, not a synonym of G. fuscolutea
  • Grimmia afroincurva is a good species, not a synonym of G. fuscolutea
  • Grimmia argyrotricha is a good species, type not seen by Muñoz
  • Grimmia arizonae is a good species, not a synonym of G. pilifera
  • Grimmia asperitricha is a good species, not a synonym of G. australis
  • Grimmia attenuata is a good species, not a synonym of G. arcuatifolia
  • Grimmia bernoullii is a good species, not a synonym of G. ovalis
  • Grimmia brittoniae is a good species, not a synonym of G. funalis
  • Grimmia brachydictyon is a good species, not a synonym of G. hartmanii
  • Grimmia crassifolia is a good species, not a synonym of G. crinitoleucophaea
  • Grimmia herzogii is a good species, not a synonym of G. longirostris
  • Grimmia khasiana is a good species, not a synonym of G. longirostris
  • Grimmia maido is a good species, not a synonym of G. longirostris
  • Grimmia mexicana is a good species, not a synonym of G. ochyriana
  • Grimmia moxleyi is a good species, not a synonym of G. orbicularis
  • Grimmia nivalis is not a good species, it is synonymous with G. caespiticia
  • Grimmia obtusolinealis is a good species, type not seen by Muñoz
  • Grimmia pilosissima is a good species, not a synonym of G. montana
  • Grimmia sessitana is a good species, not a synonym of G. reflexidens
  • Grimmia stenobasis is a good species, not a synonym of G. kidderi
  • Grimmia stolonifera is a good species, not a synonym of G. orbicularis
  • Grimmia torngakiana is a good species, not a synonym of G. incurva
  • Grimmia ungeri is endemic to Cyprus, not occurring everywhere
  • Grimmia trinervis is not a Grimmia but a Coscinodon

The majority of these incorrect data are addressed and corrected by papers from: Delgadillo (2000), Delgadillo & Villaseñor (2002), Maier (2002b), Price et al. (2003), Greven (2003), and Hastings and Greven (2007).

Redescription of Grimmia capillata De Not.
Muñoz (2002). published Grimmia exquisita spec. nov., based on two herbarium samples, one from Turkmenistan, identified as G. mesopotamica (a synonym of G. capillata), and one from Tajikistan, identified as G. crinita. In the protologue, G. exquisita was contrasted with G. pulvinata and G. orbicularis, taxa that belong to a different group with clearly exserted capsules on arcuate setae. Because gametophytes and sporophytes of G. exquisita correspond in all aspects with the type specimen of G. capillata, it is herewith put in synonymy with G. capillata De Not.

Redescription of Grimmia leibergii Paris
Ignatova, E., H. Bednarek- Ochyra, O. Afonina & J. Muñoz (2003) published Grimmia jacutica spec. nov. from eastern Asia and Alaska. Hastings and Greven examined three samples of the new species. They found only small forms of G. leibergii with slightly more sinuous mid-leaf cells. As a result of this, G. jacutica was put in synonymy with G. leibergii Paris (Hastings & Greven 2007).