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. 2025 Dec 2;267:59–80. doi: 10.3897/phytokeys.267.171699

Taxonomic and nomenclatural history of Neckera (Bryophyta, Neckeraceae), including reinstatement of Rhystophyllum, the correct name for a segregate of this genus

Ryszard Ochyra 1, Vítězslav Plášek 2,3,✉, John C Brinda 4
PMCID: PMC12690376  PMID: 41383388

Abstract

The complex taxonomic history of the moss genus Neckera, whose name was conserved at the Paris Congress in 1954, is reviewed. The issue of typifications of moss generic names by W. Ph. Schimper in 1860 is examined in detail and it is concluded that these typifications comply with the provisions of the "International Code of Nomenclature for algae, fungi and plants". Schimper was the first to typify Neckera with N. pennata, so it is unnecessary to treat this as a conserved type. The unitary designations of moss species proposed by Jakob Friedrich Ehrhart in his exsiccata "Phytophylacium Ehrhartianum", published between 1780 and 1785, are reviewed. It is concluded that two of these designations, Diphyscium and Paludella, are currently in use, having been validly published in the early nineteenth century. The same applies to Rhystophyllum, another of Ehrhart’s unitary designations, which was validated as a subsection of Neckera by C. Müller in 1850 and, subsequently, first elevated to the rank of section by Mitten in 1869, rather than Braithwaite in 1905 as listed in "Index muscorum". Finally, E. Britton raised this infrageneric taxon to the rank of genus in 1904. Rhystophyllum is here reinstated from obscurity, since it is homotypic with the generic name Exsertotheca that was recently introduced for a segregate of Neckera. Accordingly, three new combinations are proposed: Rhystophyllum crispum, Rh. intermedium and Rh. baeticum. Neckera subsect. Leiophyllum is regarded as a heterotypic synonym of Alleniella, another recent segregate of the formerly broadly circumscribed genus Neckera.

Key words: Alleniella , Exsertotheca , generic names, infrageneric taxa, Musci , nomenclature, nomina generica conservanda, pleurocarpous mosses, Madrid Code, taxonomy, unitary designations of species

Introduction

The rapid advancement of phylogenetic taxonomy in recent decades has led to the increasing subdivision of large, traditionally recognised genera. These genera were originally circumscribed using methods of classical descriptive taxonomy, based primarily on morphological and anatomical features. As a result, many have since been split into multiple, smaller genera. The newly-delineated genera are typically assigned new names, a practice permitted under the "International Code of Nomenclature" (ICN; Turland et al. (2025)). However, it occasionally emerges that some generic names, validly published in the distant past, but subsequently forgotten, still meet all the necessary criteria for use and may be appropriately applied to these newly-recognised segregates.

This article examines one such overlooked genus which, despite having fallen into obscurity, warrants reinstatement and application to one of the segregates of Neckera Hedw., a large and well-known genus of pleurocarpous mosses in the family Neckeraceae Schimp. Neckera has recently been subdivided into several smaller genera (Enroth 1991; Olsson et al. 2009a, 2009b, 2011, 2012, 2016; Enroth et al. 2022), for which new names have been proposed. One of these segregates, Exsertotheca S.Olsson, Enroth & D.Quandt, includes a common European calcicolous species previously known as N. crispa Hedw. This article reviews the taxonomic and nomenclatural history of the genus Neckera, with particular emphasis on its infrageneric classification. In many cases, the names of infrageneric taxa can be adopted for segregate genera through a simple change in status, thereby eliminating the need to coin entirely new names and to provide separate diagnoses and descriptions.

Brief taxonomic history of Neckera

From Hedwig to Müller Hallensis

Neckera Hedw. is amongst the earliest generic names in muscology, first introduced by Hedwig (1782) under the orthographic variant Neckeria Hedw. (Fig. 1). The name commemorates Noël Martin Joseph de Necker (1730–1793), a French-born botanist and bryologist who served as the personal physician to the court of the Electoral Palatinate in Mannheim, Germany (Frahm and Eggers 2001). Although Necker strongly opposed Hedwig’s view that mosses reproduce sexually, asserting instead that they do not (Schofield 1985), Hedwig nonetheless recognised Necker’s substantial contributions to botany, a fact he noted in justifying his choice of this eponym.

Figure 1.

Figure 1.

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The original publication of the generic name Neckeria Hedw. (from Hedwig 1782).

Originally, four species were placed in Neckeria, including Hypnum crispum L. A decade later, Hedwig (1792) corrected the orthography of this name to Neckera and expanded the genus to include nine additional species, such as N. pennata Hedw. and N. pumila Hedw. However, neither Neckeria Hedw. (1782) nor Neckera Hedw. (1792) constitute validly published names under modern nomenclatural rules, as both appeared prior to 1801, the year Hedwig (1801) published "Species muscorum frondosorum". This work is the designated starting-point for moss nomenclature (excluding Sphagnaceae) under the ICN (Turland 2013; Turland et al. 2025), in contrast to most other groups whose nomenclature begins with Linnaeus (1753).

Hedwig (1801) validated the generic name Neckera by giving a short diagnosis which included very general features of the peristome, namely that it is double and consists of 16 teeth of the outer peristome alternating with 16 distinct ciliate processes of the inner peristome (Fig. 2). The genus was quickly and widely accepted; however, under this broad circumscription, Neckera could encompass most pleurocarpous mosses that were primarily classified in the former order Isobryales (= Leucodontales). Indeed, this was the case, as throughout the past two centuries, over 600 specific and about 100 varietal epithets, as well as over 100 binary designations, were combined with Neckera (Wijk et al. 1964).

Figure 2.

Figure 2.

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The original publication of the generic name Neckera Hedw. (from Hedwig 1801).

One of the most striking examples of Neckera serving as a catch-all genus is found in the broad concept proposed by Müller (1850, 1851), published in September 1850 and supplemented in October 1851 (Stafleu and Cowan 1981). Müller assigned 199 species, known in the first half of the nineteenth century, to this single genus, subdividing them into nine sections and thirteen subsections, which highlighted its remarkable morphological heterogeneity. In modern classification systems, these species are now distributed across dozens of genera and multiple families of pleurocarpous mosses. In one case, the group even includes an acrocarpous species: Dicnemoloma pallidum (Hook.) Wijk & Margad. Of the 199 species included in Müller's (1850, 1851) synopsis of the genus, only 20 correspond to Neckera as currently circumscribed.

Schimper’s concept of Neckera and its development

By a remarkable coincidence, just three months after publication of Müller’s (1850) concept of Neckera − in December 1850 − Schimper (1850a), in another monograph forming part of the monumental opus "Bryologia europaea" (Stafleu and Cowan 1976), presented a concept of this genus that was diametrically opposed to Müller’s. He assigned seven European species to the genus, including N. pennata, N. pumila, N. crispa, N. oligocarpa Bruch, N. complanata (Hedw.) Huebener, N. sendtneriana Schimp. and N. philippeana Schimp., all of which were described and illustrated in detail. In addition, he referenced five extra-European species: N. menziesii Hook. then known only from North America; N. hornschuchiana Müll.Hal. and N. angustifolia Müll.Hal. from Mexico; N. chilensis Schimp. from Chile; and N. jamesonii Taylor from Ecuador, but without comment. He also listed nine further designations in the protologue, including four from Africa, two each from Mexico and South America and one from Tasmania, all ascribed to W. Ph. Schimper. The species cited in the protologue roughly correspond to the two subsections distinguished by Müller (1850) within section ‘Euneckera ’ Müll.Hal., i.e. the type section of Neckera, namely subsect. Leiophyllum Müll.Hal. and subsect. Rhystophyllum Müll.Hal. This section encompassed a number of species that are now classified under other genera, mostly related to Neckera or the Neckeraceae, such as Homaliodendron M.Fleisch., Porothamnium M.Fleisch., Neckeropsis Reichardt, Himantocladium (Mitt.) M.Fleisch., Calyptothecium Mitt. and Trachyloma Brid.

Schimper’s (1850a) newly-proposed taxonomic concept of Neckera was quickly accepted in both Europe and North America, although the genus is not particularly species-rich in these regions. As exploration of tropical and subtropical areas advanced, new species of neckeroid mosses were discovered. These species often bore only superficial morphological similarities to Neckera, but due to the underdeveloped generic classification within the Neckeraceae, they were provisionally assigned to Neckera anyway (e.g. Mitten (1859, 1869); Dozy and Molkenboer (1863); Bescherelle (1872)). Consequently, many of these species were later transferred to other genera. In "Adumbratio florae muscorum totius orbis terrarum", published in the 1870s, Jaeger and Sauerbeck (1877, 1879) included 92 species and nine designations within Neckera. However, only 50 of those species could be retained in Neckera under the current taxonomic circumscription of this genus. Over the next quarter-century, the number of species assigned to Neckera grew to approximately 150. However, the number of species truly belonging to the genus in the strict sense increased to little more than a dozen (Paris 1896, 1900, 1905).

Finally, Brotherus (1906) published the first global taxonomic overview of the genus Neckera in Engler’s "Die natürlichen Pflanzenfamilien" series, providing general keys to the species. He recognised 127 species within the genus, which were organised into six sections. This concept differed significantly from Schimper’s (1850a) circumscription of Neckera, as nearly half of the species were later found to belong to what are now widely accepted as separate genera − Neckeropsis and Himantocladium. These species were formally transferred to the respective genera in the second edition of the moss treatment in "Die natürlichen Pflanzenfamilien" (Brotherus 1925), reducing the number of Neckera species to 89. This taxonomic framework remained influential throughout the twentieth century, although the number of recognised species continued to fluctuate. Names of many previously described species were synonymised with earlier ones, while others continued to be newly described. According to Crosby et al. (2000), Neckera comprised 71 species, with 27 of these remaining uninvestigated since their original description.

Genus Neckera in the molecular era

A new chapter in the taxonomic study of Neckera began over the past two decades, marked by the integration of phylogenetic taxonomic methods, based on molecular analyses (Olsson et al. 2009a, 2009b, 2010, 2011, 2012, 2016; Enroth et al. 2022). This advancement led to a re-definition of the genus, which was subsequently divided into three segregates. Amongst them are Alleniella S.Olsson, Enroth & D.Quandt, comprising 15 species and Exsertotheca S.Olsson, Enroth & D.Quandt, containing three species (Olsson et al. 2011; Enroth et al. 2022). In addition, six monotypic genera have been newly established or reinstated to accommodate certain Neckera species. These include Caduciella Enroth (Enroth 1991), Pengchengwua S.Olsson, Enroth, Huttunen & D.Quandt (Olsson et al. 2016), Enrothia Ignatov & Fedosov (Fedosov and Ignatov 2019), as well as Indoneckera Enroth, Taiwanobryopsis Enroth and Metaneckera Steere (Enroth et al. 2022). Furthermore, eight species have been transferred to the genus Taiwanobryum Nog. (Olsson et al. 2010; Ma et al. 2018; Enroth and Shevock 2021) and ten species have been re-assigned to Forsstroemia Lindb. (Olsson et al. 2011; Enroth et al. 2019, 2022).

As a result of these taxonomic revisions, the genus Neckera currently comprises 39 extant and two fossil species (Brinda and Atwood 2025). Of the extant species, only 23 are widely recognised and accepted in taxonomic monographs, revisions and descriptive floras. The remaining 16 species are known solely from their original descriptions and have yet to undergo comprehensive taxonomic evaluation.

Nomenclatural history of the generic name Neckera

Conservation and former typifications

The generic name Neckera has a rather complex nomenclatural history. When Hedwig (1782) first published the name Neckeria for a newly-established genus of mosses, it was predated by an identical name used by Scopoli (1777) for a flowering plant in the family Papaveraceae. Even when later the spelling of this name was corrected to Neckera (Hedwig 1792, 1801), it was, from a nomenclatural perspective, merely an orthographic variant and, thus, considered a later homonym of Neckeria Scop. As such, it could not be validly used as a name for an accepted moss genus. To resolve this issue, Neckera Hedw. was formally proposed for conservation prior to the Sixth International Botanical Congress held in Amsterdam in 1935 (Dixon 1934). While this proposal was recommended by the nomenclature committee at that Congress (Dixon 1935; Magill 1993) and Neckera was placed in the list of "Nomina generica conservanda" in Appendix V of the "Stockholm Code" (Lanjouw et al. 1952), it was not formally acted upon until much later because no bryophyte names were formally conserved prior to ratification at the Paris Congress in 1954 (see below).

The typification of a generic name is fundamental to the taxonomic interpretation of a genus. This is particularly critical for older names, which often encompassed a wide array of diverse and, sometimes, unrelated species. One such name is Neckera, whose typification also has had a complex and convoluted history. According to Crosby (1968), Grout (1934) was the first to designate N. crispa as the type. However, Grout (1939) later “amended” this designation in favour of N. pennata. As noted by Brinda and Davidse (2020), this change appears to have been influenced by discussions amongst members of the nomenclature committee, who selected N. pennata as the type instead (Dixon 1939). However, this decision conflicts with Art. 10.5 of the "Code" (Turland et al. 2025), which would give precedence to the earlier designation of N. crispa.

Nevertheless, the revised typification was accepted by the General Committee and Advisory Board at the Eighth International Botanical Congress in Paris in 1954. Since the publication of the "Paris Code" (Anonymous 1954; Lanjouw et al. 1956), N. pennata has been listed as the type species of Neckera. Considering Grout’s earlier designation of N. crispa in 1934, N. pennata has been included in Appendix III of the "Code" as a conserved type since the "Montreal Code" (Lanjouw et al. 1961). Crosby (1968) opposed this outcome and proposed reinstating N. crispa as the generitype of Neckera, but his proposal was ultimately rejected by the Committee for Bryophytes (Florschütz 1973). Consequently, the recognition of N. pennata as a conserved type has achieved broad consensus (Wijk et al. 1964; Farr et al. 1979a; Greuter et al. 1993).

Schimper’s typifications of moss generic names

In accordance with the delimitation of Neckera in "Bryologia europaea" (Schimper 1850a) and its subsequent usage, N. pennata Hedw. was formally designated as the type of this generic name by Schimper (1860) in the first edition of his "Synopsis muscorum europaeorum". This typification has largely been overlooked by bryologists, despite the relevant information appearing verbatim on the title page of the work: “Accedunt tabulae VIIItypos genericosexhibentes, ...” [= “There are also eight plates showing generitypes” (emphasis ours)]. This critical phrase was first identified and interpreted in the mid-1990s by W. D. Margadant, who prepared a manuscript on Schimper’s typifications, which was submitted to Lindbergia. Pekka Isoviita of Helsinki was invited to serve as a referee for the manuscript and undertook a meticulous review, introducing numerous corrections and amendments. As a result, Margadant proposed that Isoviita be included as a co-author of the paper (Isoviita, personal communication to R. Ochyra in litteris, 23 February 1995; Margadant and Geissler (1995)). Unfortunately, the planned publication was never completed and published. However, typifications for four generic names, Dicranella (Müll.Hal.) Schimp., Hylocomium Schimp., Pylaisia Schimp. and Homalia Brid., were later published by Margadant and Geissler (1995) as part of formal proposals to conserve these names.

For three of these, conservation of the types was necessary to maintain current usage, which had been threatened by Schimper’s (1860) overlooked type designations involving different species. In the case of Homalia, however, the typification of the name was merely confirmed by Schimper (1860), since in "Bryologia europaea", Schimper (1850b) had used the term Grundform, in the German text only, as an equivalent of “type”. All these proposals were accepted by the Committee for Bryophyta (Zijlstra 1998, 1999) and the conserved generic names were subsequently included in Appendix III of the "Saint Louis Code" (Greuter et al. 2000), thereby formally endorsing Schimper’s (1860) typifications.

In addition to the article by Margadant and Geissler (1995), published in the November issue of Taxon, Volume 44, Margadant’s discovery was also acknowledged in the monograph on the genus Racomitrium Brid. in Poland (Bednarek-Ochyra 1995). Notably, this monograph was published even earlier, on 1 October 1995, than the aforementioned article. As mentioned above, the information regarding the typification of generic names in Schimper’s (1860) work had been communicated to R. Ochyra by P. Isoviita, who also granted permission for its inclusion in the monograph of Racomitrium. This case is of particular interest because Schimper (1860) designated R. lanuginosum (Hedw.) Brid. as the type, thereby rendering superfluous a later typification by Pfeiffer (1874), who had proposed R. canescens (Hedw.) Brid. instead. This change in type somewhat affected the nomenclature of infrageneric taxa within Racomitrium and subsequently influenced the naming of segregate genera derived from this heterogeneous and broadly circumscribed genus (Bednarek-Ochyra et al. 2014; Sawicki et al. 2015).

For the next quarter-century, bryologists largely overlooked the typification of generic names in Schimper’s (1860) opus. Zijlstra (2015) reignited the debate by claiming that the phrase on the book’s title page implies the figures represent typical species. This, however, is a clear overstatement: Schimper (1860) explicitly used the term type (typus), not typical (typicus). While it is possible that Schimper meant “exemplar”, not our modern nomenclatural type, that is irrelevant under the current wording of the "Code" (Turland et al. 2025). Notably, the same terminology appears in "Nomenclator botanicus" by Pfeiffer (1873a), who included a general statement on the unnumbered second page of the Prefatio: “Species plantarum in libro meo omnino negliguntur, excepta indicatione illarum, quaetypumgeneris novi aut novo modo circumscripti vel sectionis offerunt” [= “Plant species are entirely neglected in my book, except for indication of those which are presented as the type of a new or re-circumscribed genus or of a section” (emphasis ours)]. Like Schimper (1860), Pfeiffer (1873a) clearly indicated type-species for generic or sectional names and these typifications are acceptable under the "Code" (Art. 7.11, Ex. 18; Turland et al. (2025)). Of course, it occasionally emerges that Schimper’s or Pfeiffer’s choices were erroneous or based on a misinterpretation of the protologue. However, such cases do not undermine the fundamental validity of their typifications. Similar issues occur even in contemporary practice and erroneous choices can be corrected in accordance with Art. 10 of the "Code" (Turland et al. 2025).

Typification of moss generic names by Schimper (1860) once again drew the attention of the Committee for Bryophytes in the early 2020s, during discussions surrounding the proposal to conserve the name Oreas Brid. (Brinda and Fedosov 2020). Although the name was formally proposed for conservation with the conserved type Oreas martiana (Hoppe & Hornsch.) Brid., based on the then-prevailing assumption that Oreas had first been typified by Pfeiffer (1873b) with “Weissia Mielichhoferi Schwägr.” (≡ Weissia mielichhoferiana Funck), the Committee ultimately accepted Schimper’s (1860) earlier typification of this generic name with O. martiana as effectively published (Klazenga 2023).

This decision was subsequently accepted by the General Committee with the suggestion that “bryologists should check whether this acceptance of Schimper’s typification in his 1860 paper has consequences for typifications of other names in that paper” (Wilson 2023). However, most of the issues arising from Schimper’s (1860) typifications, which had significantly affected the status of several of the aforementioned generic names, have in fact already been resolved via conservation proposals (Margadant and Geissler 1995).

Schimper’s (1860) typifications have recently gained broader recognition, as reflected in the TROPICOS database (https://www.tropicos.org/), where an attempt has been made to correctly list the types of all bryophyte genera. This includes the generic name Neckera, which was first typified with N. pennata (https://www.tropicos.org/name/35000848). In Table VI, Schimper (1860) presented an illustration labelled only with the name Neckera. However, the accompanying legend specifies N. pennata and includes captions identifying the illustrated details. Consequently, Dixon’s (1939) seemingly arbitrary selection of N. pennata aligns with Schimper’s earlier, but long-overlooked typification. This finding clarifies that Dixon’s (1939) choice was indeed correct and that Grout’s (1934) typification was not the earliest. Therefore, N. pennata should not be listed as a conserved type in Appendix III of the "Code", as suggested by Brinda and Davidse (2020), but rather its current status should be maintained. As noted above, N. pennata was treated as the conserved type of Neckera from the "Montreal Code" (Lanjouw et al. 1961) through to the "Tokyo Code" (Greuter et al. 1994). However, in the mid-1990s, W. D. Margadant re-discovered Schimper’s (1860) earlier typifications of many generic names, including Neckera. As a result, treating N. pennata as a conserved type became unnecessary. Accordingly, beginning with the "Saint Louis Code" (Greuter et al. 2000), the phrase "typus conservandus" associated with N. pennata was editorially removed.

Synonyms of the generic name Neckera

In summary, the generic name Neckera currently has two heterotypic synonyms and ten homotypic synonyms (Brinda and Atwood 2025). The first heterotypic synonym is Alsia Sull., formerly treated as a monotypic genus endemic to the Pacific coast of North America (Schofield 1980). It included only A. californica (Hook. & Arn.) Sull., a species originally described as Neckera californica Hook. & Arn., which has recently been transferred back to Neckera (Olsson et al. 2011). The second is Neckera sect. Douglasiella (Kindb.) Paris, typified by the North American species N. douglasii Hook. Three of the homotypic synonyms are homonyms and seven represent names of homotypic infrageneric taxa. Of these, Distichia (Brid.) Brid. and Braunia Hornsch. are considered nomina rejicienda, while Cryptopodium Fürnr. is both a homonym and a superfluous name by typification. Additionally, Cryptopodia (Röhl.) Fürnr. is a superfluous name by typification and Eleutera P.Beauv. ex Stuntz is illegitimate, as it included the previously designated type of Neckera.

Taxonomic and nomenclatural history of Rhystophyllum

Earliest plant exsiccatae

The name Rhystophyllum first appeared in bryological literature through the work of Jakob Friedrich Ehrhart (1780–1785, 1789), who used it on the label of a specimen distributed in 1780 as No. 79 in the eighth decade (a set of ten numbered specimens) of his exsiccata "Phytophylacium Ehrhartianum" (Fig. 3). Ehrhart (1742–1795), a German botanist of Swiss origin and a gifted student of Linnaeus, held the position of Royal Botanist in the Electorate of Hanover (Braunschweig-Lüneburg) during the final decade of his life. Ehrhart is best known in the history of botany as one of the pioneers in publishing exsiccatae, numbered collections of dried and pressed plant specimens, distributed under a common title, precisely identified and accompanied by printed labels, intended for circulation amongst botanists and institutions.

Figure 3.

Figure 3.

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The specimen No. 79 from "Phytophylacium Ehrhartianum" from UPS with the unitary designation Rhystophyllum of Neckera crispa Hedw.

Although Johann Balthasar Ehrhart (1700–1756) is regarded as the first publisher of an exsiccata, his "Herbarium vivum recens collectum", issued in 1732, contained only medicinal plants and no cryptogams. Today, only three copies of this earliest exsiccata are known to exist (Stafleu 1972). On the other hand, Jakob Friedrich Ehrhart – the similarity of names is coincidental – published for commercial purposes seven series of exsiccatae in the years 1780–1793, covering about 1620 species of flowering and cryptogamic plants (Britten 1922, 1923; Stafleu and Cowan 1976; Sokoloff et al. 2002). For bryologists, the first two series are particularly important, the above-mentioned "Phytophylacium Ehrhartianum" and "Plantae cryptogamae Linneae" (Sayre 1969), comprising 320 specimens of mosses, liverworts, lichens, algae, club-mosses, horsetails and ferns, distributed in decades (Ehrhart 1785–1793, 1792). In the first of these series, only six species of mosses and ten lichens were distributed from the cryptogams. However, they are particularly interesting from a nomenclature perspective. On all labels, the standard Latin name of a species followed by a locality precede, as if in the form of a title, the specimen number and Latin uninomial on the top line (Fig. 3). Ehrhart (1789) calls them "nomina usualia", i.e. common names and treats them as an attempt to give each plant a distinct name that would be used independently of the binomial name. Although the author himself does not consider them equivalent to generic names, in later years, various authors began to consider them as such, although from a nomenclature perspective, they were typical "nomina nuda", as Ehrhart never published any diagnoses of them.

Ehrhart’s unitary designations of species

The confusion surrounding generic names was addressed at the International Botanical Congress held in Montreal in 1959. The resulting "Montreal Code" (Lanjouw et al. 1961) expanded Article 20 by adding a note clarifying what should not be considered generic names. The second point of this note referred to “unitary designations of species”, exemplified by Ehrhart’s (1780–1785, 1789) names in "Phytophylacium Ehrhartianum", which imitated generic names. According to the "Code", such designations were to be rejected unless they have been validly published as generic names by subsequent authors. This provision remained in force until the Tokyo Congress in 1993, when a new Appendix V, "Opera utique oppressa", was introduced (Greuter et al. 1994). This Appendix listed publications considered suppressed works, meaning that names at specified ranks within them were not validly published. Amongst the works included was Ehrhart’s exsiccata "Phytophylacium Ehrhartianum" and its associated species index, published in 1789. As a result, Article 20.4, which had provided examples of names that should not be treated as generic names, was removed from the "Code". At the Melbourne Congress in 2012, Appendix V was renamed " Suppressed works" and re-designated as Appendix VI (Wiersema et al. 2015).

Of the six unitary designations for moss species listed in "Phytophylacium Ehrhartianum" (Ehrhart 1780–1785, 1789), only two, Hippopodium and Tristichitis, have fallen into complete obscurity. The former, applied to Buxbaumia aphylla Hedw. in the first decade under No. 10, was originally borrowed from Fabricius (1743). However, the generic name Buxbaumia Hedw. had already been introduced a year earlier by Haller (1742) and was subsequently accepted as the older name by Schmidel (1758) and Linnaeus (1760, 1763). Nevertheless, Röhling (1808) validated Hippopodium Röhl. as a generic name with H. aphyllum (Hedw.) Röhl. as its generitype. This makes Hippopodium homotypic with Buxbaumia which has nomenclatural priority. Despite evident priority of Buxbaumia, Röhling (1813) continued using Hippopodium, but since his death, this name has fallen into oblivion.

The designation Tristichitis was applied to Mnium triquetrum L. (≡ Meesia triquetra (L. ex Jolycl.) Ångstr.; see Ochyra (2001)), a species distributed in the sixth decade under No. 59. It was later only once considered by Röhling (1800) in a synopsis of the German moss genera, as a taxon close to Meesia Hedw. He provided a short diagnosis of Tristichitis, but no species were included. As a pre-starting point name, Tristichitis is not validly published and has been supplanted by the generic name Meesia Hedw. (Hedwig 1785–1787), which was validated in "Species muscorum frondosorum" (Hedwig 1801).

Finally, the unitary designation Ampullaria was assigned to Splachnum ampullaceum Hedw., published in the fifth decade under No. 49 (Ehrhart 1780–1785). Müller (1848) subsequently validly published this name as a section within Splachnum Hedw.; however, it is now superfluous given that Schimper (1860) selected Splachnum ampullaceum Hedw. as the type of this genus name.

Of the three remaining unitary designations, all have persisted to the present day, with two still in common use. The earliest legitimised name was Diphyscium, which Ehrhart (1780–1785) used to designate a specimen of Webera diphyscium, published in the tenth decade as No. 99. Notably, this binomial was first published in that work, not in Ehrhart’s (1779) earlier publication describing the genus Webera, as incorrectly reported by Limpricht (1894) and Wijk et al. (1969). Mohr (1803) later validated the generic name Diphyscium, assigning to it a single species: D. foliosum (Hedw.) D.Mohr. While Mohr acknowledged Ehrhart in his discussion: “Itaque triviali Ehrharti vocem, tanquam substantivum, pro novo conficiendo generi nomine lubenter recepi” [= “Therefore, I gladly accepted Ehrhart’s trivial word, as a substantive, for the newly-coined genus name”], he ascribed the generic name to himself alone as “mihi”.

The second unitary designation, Paludella, was established by Ehrhart (1780−1785) and remains in common use today. It was originally proposed for Bryum squarrosum L., a strikingly beautiful peatland moss, which was published as No. 69 in the seventh decade of his exsiccata. Initially, this species was placed in several broad genera, including Bryum Hedw., Mnium Hedw. and Hypnum Hedw. Eventually, Bridel (1817) transferred it to the monotypic genus Paludella Ehrh. ex Brid., adopting the uninomial Ehrhart had introduced 37 years earlier.

Validation of Rhystophyllum

The third and final unitary designation that has withstood the test of time is Rhystophyllum. Ehrhart (1780–1785) originally proposed this name for the calciphilous moss Hypnum crispum, which is widely distributed throughout Europe (Ochyra et al. 1988). He included it as No. 79 in the eighth decade of his first exsiccata, "Phytophylacium Ehrhartianum" (Fig. 3). The name was later revived by Müller (1850), who adopted it for one of two subsections within Neckera sect. ‘Euneckera ’. In defining subsect. Rhystophyllum Müll. Hal., Müller provided a brief diagnosis − "Folia transversim rugosa" − to distinguish it from subsect. Leiophyllum Müll.Hal., whose species possess "Folia laevia, haud transversim rugulosa". Subsect. Rhystophyllum was subsequently elevated to the rank of section, though there is considerable confusion regarding the identity of the author, as well as the date and place of publication. According to the Index muscorum (Wijk et al. 1964) and Crosby (1968), the name was validated by Braithwaite (1905). However, the elevation of subsect. Rhystophyllum to the sectional rank was actually carried out much earlier by Mitten (1869).

The "Index muscorum" (Wijk et al. 1964) suggests that Lindberg (1879) most likely treated this taxon as a subgenus of Neckera. According to the compilers of this compendium, although this author does not explicitly indicate the rank of the subdivision in that publication, they assume that all subdivisions of genera designated by capital letters are most probably subgenera. Indeed, Lindberg (1879) does not define the rank of these subdivisions in the main text. It is only in the final chapter, entitled "Comparatio denominationum in editione decima Florae Hartmanianae cum iisdem in hoc opere nostro datis" (pp. 45–48), that he clearly refers to them as sections. For example, Racomitrium [in Hartman (1871)] corresponds to Grimmiasect. A [in Lindberg’s work] (emphasis ours). Thus, following Mitten (1869) and Bescherelle (1872), this constitutes the third isonym of Neckera sect. Rhystophyllum (Müll.Hal.) Mitt. Formally, the status of Rhystophyllum as a subgenus of Neckera should be attributed to the compilers of the "Index muscorum", as reflected in the TROPICOS database [https://www.tropicos.org/name/35171285], where it is cited as Neckera subg. Rhystophyllum (Müll.Hal.) Lindb. ex Wijk, Margad. & Florsch. However, this is an invalid name, erroneously used for Neckera sect. Rhystophyllum.

Given that various genera proposed by Ehrhart were previously accepted and that Linnaeus’s (1753) "Species plantarum" was still the starting point for moss nomenclature at the time, Britton (1905) concluded that Rhystophyllum also merited recognition as a distinct genus and that it predated Neckera. Although she did not acknowledge that Ehrhart (1780–1785) failed to provide a formal description or diagnosis of this “genus”, she nonetheless included a very brief diagnosis on page 4 of her text: “…as Ehrhart’s genus Rhystophyllum is monotypic, […], there is no question as to its meaning or the application of the name, seeing that its derivation from two Greek words meaning ‘Wrinkled-Leaved’ indicates one of the most noticeable characters of the genus as limited in modern times” (emphasis ours). This single morphological trait may be accepted as sufficient to serve as a validating description of the genus. In fact, this character was also used by Müller (1850) when describing subsect. Rhystophyllum. As the requirements for valid publication of Rhystophyllum are fulfilled in Britton’s (1905) paper, it may be considered a cryptic reference to the subsectional name Neckera subsect. Rhystophyllum as used by Müller (1850: p. 46), in accordance with Art. 41.4 of the "Code" (Turland et al. 2025). Notably, the citation accompanying the genus name Rhystophyllum − “Crypt. Exsic. No. 97. 1780” − constitutes an indirect reference to Müller’s (1850) work, as it even reproduces his error in the cited specimen number: “No. 97” should correctly read “No. 79”.

Britton (1905) considered Hypnum crispum (≡ Neckera crispa) to be the generitype of Rhystophyllum as well as Hedwig’s Neckera. She was, therefore, in agreement with Grout’s erstwhile selection for Neckera, but in conflict with Schimper’s. She was likely unaware that Pfeiffer (1874) had previously typified Müller’s subsect. Rhystophyllum with the same species, but this merely reinforces the interpretation that Britton’s Rhystophyllum should be treated as a status nova rather than the name of a new taxon or a nomen novum for Neckera. Given that Britton’s Rhystophyllum has a legitimate basionym with a type that is different from the type of Neckera, the name must be considered legitimate even though Britton incorrectly included the previously designated type of Neckera in her circumscription (Art. 52.4, Turland et al. (2025)). For most of the last century, Neckera has been treated as a large, heterogeneous genus typified by N. pennata and Rhystophyllum was considered a heterotypic synonym (Farr et al. 1979b). However, following the division of Neckera into three segregate genera, Neckera s.str., Exsertotheca and Alleniella (Olsson et al. 2011), Rhystophyllum (Müll.Hal.) Ehrh. ex E.Britton should be recognised as the correct name for Exsertotheca. Both names are homotypic synonyms typified by Neckera crispa, but Rhystophyllum has nomenclatural priority.

The second segregate of Neckera, Alleniella, also has a related subsectional name. In addition to subsection Rhystophyllum, which includes species with distinctly undulate leaves, Müller (1850) recognised subsection Leiophyllum, comprising species with smooth, non-undulate leaves. According to Art. 10.8 of the "Code" (Turland et al. 2025), this subsection is automatically typified by Neckera leiophylla W.Gümbel ex Müll.Hal. This species is conspecific with N. besseri (Łobarz.) Trevis., which is the type species of Alleniella. However, elevating subsect. Leiophyllum to the rank of genus is pre-empted by Leiophyllum (Pers.) R.Hedw. (1806, Ericaceae), so it may only be used at the infrageneric ranks.

Nomenclatural implications

. Neckera

Hedw., Spec. Musc. Frond.: 200. 1801 nom. cons.

0EE0A593-8384-5741-A379-7309C513D74C

  • ≡ Neckera [unranked] Cryptopodia Röhl., Borkhausen’s Ringen: 148. 1808 ≡ Neckera [unranked] Distichia Brid., Muscol. Recent. Suppl. 4: xvi, 137. 1819 ≡ Distichia Brid., Bryol. Univ. 1: xxxvii. 1826; 2: 238, 757, 787, 811. 1827 ≡ Neckera subg. Distichia (Brid.) Brid., Bryol. Univ. 2: 238, 757. 1827 ≡ Cryptopodium Fürnr., Flora 12(2, Beil. 1): 81. 1827 ≡ Braunia Hornsch., Jahrb. Wiss. Krit. 1828(59/60): 467. 1828, nom. illeg. ≡ Neckera subg. Cryptopodia (Röhl.) Rchb., Consp., Regn. Veg. 1: 33. 1828 ≡ Cryptopodia (Röhl.) Fürnr., Flora 12(2, Ergänzugsbl.): 49. 1829 ≡ Neckera sect. Distichia (Brid.) Müll.Hal., Linnaea 18: 707. 1844 ≡ Eleutera P.Beauv. ex Stuntz, Bull. Torrey Bot. Club 27: 202. 1900, nom. illeg. ≡ Neckera sect. Cryptopodia (Röhl.) Broth. in Engl. & Prantl, Nat. Pflanzenfam. 1(3): 843. 1906. Typus: Neckera pennata (videSchimper 1860: t.p., 732).

  • = Alsia Sull., Not. Sp. Moss.: 16. 1855. Typus: Alsia californica (Hook. & Arn.) Sull. (Neckera californica Hook. & Arn.)

  • = Neckera [unranked] Douglasiella Kindb., Eur. N. Amer. Bryin. 1: 15. 1897 ≡ Neckera sect. Douglasiella (Kindb.) Paris, Index Bryol. Suppl.: 132. 1900. Typus: Neckera douglasii Hook. (videWijk et al. 1964: 432).

. Rhystophyllum

(Müll.Hal.) Ehrh. ex E.Britton, Bryologist 8(1): 4−5. 27 Dec 1904

0ABC4A82-5736-5CC6-A22B-D105D9F382FA

  • ≡ Neckera subsect. Rhystophyllum Müll.Hal., Syn. Musc. Frond. 2: 46. Sept 1850 ≡ Neckera sect. Rhystophyllum (Müll.Hal.) Mitt., J. Linn. Soc. Bot. 12: 453. 1869 ≡ Neckera subg. Rhystophyllum (Müll.Hal.) Wijk, Margad. & Florsch., Regnum Veg. 33: 432. 1964, nom. inval. err. pro sect. ≡ Exsertotheca S.Olsson, Enroth & D.Quandt, Taxon 60(1): 45. 2011. Typus: Rhystophyllum crispum (Hedw.) Ochyra, Plášek & Brinda (Neckera crispa Hedw., videPfeiffer 1874: 965).

. Rhystophyllum baeticum

(J.Guerra, J.F.Jiménez & J.A.Jiménez) Ochyra, Plášek & Brinda comb. nov.

6EBB17B2-F6D0-5B93-B932-F273986ADEA3

Basionym.

Neckera baetica J.Guerra, J.F.Jiménez & J.A.Jiménez, Nova Hedwigia 91: 259. 2010.

. Rhystophyllum crispum

(Hedw.) Ochyra, Plášek & Brinda comb. nov.

232BC628-A1CB-55F9-995F-919F83CCD13D

Basionym.

Neckera crispa Hedw., Spec. Musc. Frond.: 206. 1801.

. Rhystophyllum intermedium

(Brid.) Ochyra, Plášek & Brinda comb. nov.

65FF4B75-3A02-5113-9EB0-88E7A0A7DF62

Basionym.

Neckera intermedia Brid., Muscol. Recent. Suppl. 2: 24. 1812.

. Alleniella

S.Olsson, Enroth & D.Quandt, Taxon 60(1): 45. 2011

684DF9ED-73A7-5191-846A-AC4C1B29C1BF

  • = Leskea [unranked] Complanatae Brid., Muscol. Recent. Suppl. 2: 50. 1812 ≡ Hypnum sect. Complanata (Brid.) Arn., Disp. Méth. Mousses: 58. 1825. Typus: Leskea complanata Hedw., syn. nov.

  • = Neckera subsect. Leiophyllum Müll.Hal., Syn. Musc. Frond. 2: 41. Sept 1850 ≡ Neckera sect. Leiophyllum (Müll.Hal.) Lindb., Musci Scand.: 40. 1879 ≡ Neckera subg. Leiophyllum (Müll.Hal.) Lindb. ex Wijk, Margad. & Florsch., Regnum Veg. 33: 432. 1964, nom. inval. err. pro sect. Typus: Neckera leiophylla W.Gümbel. ex Müll.Hal. [= Alleniella besseri (Łobarz.) S.Olsson, Enroth & D.Quandt], syn. nov.

Supplementary Material

XML Treatment for Neckera
XML Treatment for Rhystophyllum
XML Treatment for Rhystophyllum baeticum
XML Treatment for Rhystophyllum crispum
XML Treatment for Rhystophyllum intermedium
XML Treatment for Alleniella

Acknowledgements

We thank Dr Åsa Kruys, Curator in the Museum of Evolution (UPS), Uppsala University, Sweden, for availability of digitised specimen of Neckera crispa from "Phytophylacium Ehrhartianum".

Citation

Ochyra R, Plášek V, Brinda JC (2025) Taxonomic and nomenclatural history of Neckera (Bryophyta, Neckeraceae), including reinstatement of Rhystophyllum, the correct name for a segregate of this genus. PhytoKeys 267: 59–80. https://doi.org/10.3897/phytokeys.267.171699

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Use of AI

No use of AI was reported.

Funding

The study of R. Ochyra has been financially supported by the statutory fund of the W. Szafer Institute of Botany of the Polish Academy of Sciences in Kraków. Work of V. Plášek was carried out at MCBR UO (International Research and Development Center of the University of Opole), which was established as part of a project co-financed by the European Union under the European Regional Development Fund, RPO WO 2014–2020, Action 1.2 Infrastructure for R&D. Agreement No. RPOP.01.02.00-16-0001/17-00 dated 31 January 2018.

Author contributions

RO, VP, JCB prepared the draft version of the manuscript, which was reviewed, edited and approved by all co-authors. The authors have contributed equally.

Author ORCIDs

Ryszard Ochyra https://orcid.org/0000-0002-2541-0722

Vítězslav Plášek https://orcid.org/0000-0002-4664-2135

John C. Brinda https://orcid.org/0000-0001-9083-1235

Data availability

All of the data that support the findings of this study are available in the main text.

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

XML Treatment for Neckera
phytokeys.267.171699-treatment1.xml (9.1KB, xml)
XML Treatment for Rhystophyllum
phytokeys.267.171699-treatment2.xml (5.1KB, xml)
XML Treatment for Rhystophyllum baeticum
phytokeys.267.171699-treatment3.xml (2.8KB, xml)
XML Treatment for Rhystophyllum crispum
phytokeys.267.171699-treatment4.xml (2.7KB, xml)
XML Treatment for Rhystophyllum intermedium
phytokeys.267.171699-treatment5.xml (2.8KB, xml)
XML Treatment for Alleniella
phytokeys.267.171699-treatment6.xml (5.2KB, xml)

Data Availability Statement

All of the data that support the findings of this study are available in the main text.

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