Currently, the species Bromus alopecuros Poir. has at least two synonyms: B. alopecuroides Poir. and B. contortus Desf.

In describing Bromus alopecuros, Poiret (1789: 100) stated in the protologue diagnosis “Panicula conferta erecta, spiculis oblongis subsessilibus, aristis inferne spiraliter contortis.” followed by a description in French and the locotypic indication indicating this species grows around La Calle [now El Kala, El Tarf province, Algeria].

During our search for original material in the herbaria conserving the plants of Poiret (FI, H, P and UPS), we found only one specimen collected by him. There is a sheet in P (P02622864) from Numidia (Algeria “Numidia” included in the full title of his publication), registered as original material, bearing a single plant annotated as B. alopecuros with five labels transcribed below; one of them handwritten by Poiret:

Label 1: [printed]. Herb. Poiret in Herb. Moquin-Tandom.

Label 2: [handwritten by Poiret] “brom. contortus Desf. Bromus alopecuros (n) Panicula conferta erecta spiculis oblongis subsessilibus, aristis inferne spiraliter contortis. (nobis) h. Poiret ex Numidia.

Label 3: [printed]: barcode Herbier museum Paris P 02622864

Label 4: [handwritten] Bromus alopecuros Poiret, Det. P. Smith 3/72

Label 5: [printed]. Herb. Mus. Paris

As this specimen bears a label handwritten by Poiret with his description of B. alopecuros and the word “nobis” [our], indicating the author is describing a new species, it seems sure that it is original material and it is suitable to be described as a lectotype. In addition, there is no other specimen matching with Bromus alopecuros Poir. in FI (Chiara Nepi pers. comm. 2018); nor in H (Raino Lampinen pers. comm. 1994; Henry Väre pers. comm. 2018) nor in UPS (Dr. Mats Hjertson pers. comm. 2018).

When Desfontaines (1798: 95), describes Bromus contortus, his description closely matches that of Poiret (1789). In his diagnosis, Desfontaines adds that the spikelets are “quindecimfloribus, …pubescentibus”. At the end, he has doubts about his plant and transcribes a reference to the description of Poiret “An Bromus alopecuros? Poiret. Itin. 2. P. 100)”. Subsequently, he includes a more detailed description and asserts “Habitat prope La Calle”. The only known original material for B. contortus is the illustration quoted in Desfontaines (1798: plate 25) and a single sheet in P (MNHN-P P00320328) bearing two specimens mounted on it with a printed label: Herbier de la FLORE ATANTIQUE donné au Museum par M. DESFONTAINES and annotated “Bromus contortus” handwritten (unknown by whom). There is another label, handwritten by Desfontaines, containing the exact diagnosis and description as it appears in Flora Atlantica (Desfontaines 1798: 95). Maybe Desfontaines is surprised one specimen is very similar to B. alopecuros, but not the other one, justifying his doubt in the description where he states “perhaps B. alopecuros?” This sheet bears two plants. The one on the left seems to be Bromus lanceolatus Roth and the one on the right is a fragment (contracted and erect panicle with subsessile spikelets) of a specimen matching B. contortus Desf., that also exemplifies the typical resemblance to B. alopecuros Poir. Therefore, the sheet in question does not represent a specimen as defined in the Code (Turland et al. 2018); but each of the two plants on the sheet is a specimen in its own right. Only one specimen is original material for B. contortus Desf. It is possible that the illustration in Desfontaines (1798: plate 25) was drawn from the plant designated here as lectotype.

Figure 1. 

Lectotype of Bromus alopecuros: It is a complete specimen preserved at the MNHN Collection Vascular plants Specimen with barcode P02622864. (Available at http://coldb.mnhn.fr/catalognumber/mnhn/p/p02622864).

Choosing the specimen on the right of the sheet MNHN-P P00320328 (Figure 2) as lectotype, which taxonomically matches B. alopecuros Poir., the name becomes a taxonomic or heterotypic synonym (Turland et al. 2018) of it, as Persoon (1805, 1: 95) asserted. Moreover, it is also possible that the heterogeneous material in this sheet is the origin of some misidentifications of B. lanceolatus Roth as B. contortus Desf.

Figure 2. 

Lectotype of Bromus contortus Desf. is the specimen on the right having subsessile spikelets conserved in the MNHN Collection Vascular plants with barcode P00320328 (image available at: https://science.mnhn.fr/institution/mnhn/collection/p/item/p00320328?listIndex=1&listCount=4).

Later, Poiret (1810: 703) describes Bromus alopecuroides “Bromus panicula conferta, erecta; spiculis oblongis, pubescentibus, quindecimfloris, subsessilibus; aristis infernè spiraliter contortis”. This description is almost identical to the former of B. alopecuros. Its only difference is to include “pubescentibus, quindecimfloris”, the same features Desfontaines (1798: 95) uses to describe B. contortus. Poiret continues adding the references to B. alopecuros Poir. and the synonym B. contortus Desf. Therefore, this name does not need a Lectoype as it is a superfluous and illegitimate renaming (Art. 52. 1) of Bromus contortus Poir. and has the same type we select here for that name.

Based on Schenodorus benekenii Lange.

The current widely used name Bromus benenkenii is an implicit combination by Trimen (1872: 333) of the name described as Schedonorus benekenii Lange in Flora Danica. In the original publication, Lange (1871: 5) describes a perennial Bromus living in forests, with nodding panicle.

An exhaustive search to find the material Lange (1871) cites as models for the illustrations in table 2826 (figs 1, 2), finally had a result. The two syntypes are conserved in C in the Flora Danica subherbarium, that contains specimens drawn in the magnificent work “Flora Danica” (Olof Ryding pers. comm. 2018). Their labels state “insula Lolland legit cl. E. Rostrup” (C10021728)” and “Jonstrup Vang, legit cl. H. Mortensen” (C10021729). Each folder indicates in handwriting that the specimens were drawn for “Flora Danica”.

We choose as lectotype the latter sheet since its spikelets conserve most of its florets. The other sheet is in a more advanced phenological state and conserves, in most of its spikelets, only the glumes. The Rostrup specimen still conserves its basal part, with leaves, that is missing in the lectotype.

Figure 3. 

The original specimen of Schedonorus benekenii Lange illustrated in Flora Danica (in table 2826, fig. 2) is the lectotype selected (C10021729). Reproduced with permission of the Natural History Museum of Denmark.

The lectotype has a handwritten label by Schrader “B. commutatus / Fl. Germ. / Göttingen” and a handwritten indication as “typus-material”. Currently, there is another label on this sheet by H.Scholz 1998 marking it as Neotypus, which was published in Scholz (1999: 436). Although both publications fulfil the conditions for a formal typification of a lectotype or a neotype, the Acedo and Llamas (1999: 73) publication appeared in February and is probably the first typification (ICN, Art. 10. 5). There are two important facts to consider. First, according to Kerguélen (1975: 100), there is original material in GOET. We consulted that herbarium and found two sheets that Schrader sent to GFW Meyer, which are indeed original material. Both bear handwritten labels by Schrader. The first sheet label says “B. commutatus Fl. Germ. Göttingen” and the second sheet label “B. commutatus Fl. Germ. var. spic. paulo brevior. Göttingen”.

As a fire in his house destroyed Hudson’s personal herbarium, most specimens were lost and only those borrowed by other botanists are extant. Some specimens which Hudson sent to Linnaeus are preserved in the herbarium of the Linnean Society of London (LINN) but none of them is Bromus erectus Huds. There is also one sheet conserved in BM that does not seem to be original material. As Hudson did not give any other reference and, as no original material of B. erectus Huds. is available, a neotype must be designated (ICN, Art. 9.13). We searched for material coming from Kent (“Cantium”, England) in K and selected one specimen consistent with the protologue as neotype. It was collected in Kent, near Wye. There is a duplicate of it in BRI, which is an isoneotype.

Figure 4. 

Neotype selected for Bromus erectus Huds. Sheet preserved at Herbarium K, barcode K00618780. [Available at http://specimens.kew.org/herbarium/K000618780]. Image used with permission Board of Trustees of the Royal Botanic Gardens, Kew.

Based on Bromus intermedius Guss. subsp. divaricatus Bonnier & Layens.

Bromus hordeaceus L. is a very variable species with a complex nomenclatural history at subspecies level. The origin of many problems in the group is the name B. molliformis J.Lloyd, Fl. Loire-Inf. (Lloyd 1844: 314–315), which was invalidly published as a provisional name “je proposerais, si c’était une espèce nouvelle, de l’appeler Br. molliformis”. The name was later validated as Bromus molliformis J.Lloyd ex Billot, Fl. Gall. & Germ. Exsicc. (Haguenau) 2: Cent. 14. 1854. Billot supplies a brief description for “cette espèce donnée par M.Lloyd dans sa Flore de la Loire-Inférieure sous le nom de B. divaricatitus Rohde?” Billot (Feb. 1854: 297–298) neglects to mention several details of Lloyd’s description of B. molliformis and describes the species as follows:

“Racine fibreuse. Chaume de 2–4 décimètres. Feuilles et graines inférieures mollement poilues. Panicule oblongue, droite, étalée, resserrée après la floraison; pédoncules courts, simples. Epillets oblongs, étalés, velus; arête égalant la glumelle, d’abord droite, à la fin tortillée divariquée, insérée à 1 1/2 millimètre du sommet obtus, échancré”.

Billot (1854) publishes the first validating description for Bromus molliformis. He includes the reference of the features to differentiate the new species from the closely related B. hordeaceus L. and from B. divaricatus Rhode. He sold his exsiccata to several herbaria. There is not a register of those herbaria. Currently, some of them are included in P where some original specimens with its species number (1386) are preserved. Additionally, there are several names and combinations at several ranks, both validly and invalidly published and various nomenclatural changes. Nevertheless, B. molliformis J.Lloyd ex Billot remained untypified and still lacked a stable nomenclature.

Serrafalcus lloydianus Godr. & Gren. (1855: 591) is a superfluous name for B. molliformis J.Lloyd ex Billot. Grenier and Godron (1855) listed several synonyms, including “B. divaricatus Lloyd non Rhode” and B. molliformis J.Lloyd ex Billot [sub. B. molliformis Lloyd]. They also mention Billot’s exsiccate n. 1586; and include his new species in the complex of species having “arête … tordue sur elle-même et divariquée”. Therefore, this name does not need a Lectoype as it is a superfluous and illegitimate renaming (Art. 52.1) of Bromus molliformis J.Lloyd ex Billot and has the same type we select here for that name.

Bonnier and Layens (1894: 369) describe Bromus intermedius Guss. subsp. divaricatus Bonnier & Layens, based on B. divaricatus [sensu] Lloyd, non Rhode, with the characteristics of Lloyd’s plant. Kerguélen (1981: 27) combines it to Bromus hordeaceus subsp divaricatus (Bonnier & Layens) Kerguélen. Bonnier and Layens type material is unknown (Stafleu and Cowan 1976). Kerguélen (1975: 104) mentions Lloyd’s type material is in herbarium NTM, pointing out two localities “Pornic, Saint Brevin” indicated by Lloyd. Nevertheless, no original material is preserved in NTM (Mary Laury Guerin, Com. pers. 2018).

Bromus hordeaceus L. var. molliformis (J.Lloyd ex Billot) Halácsy (1904: 396). This is another combination of B. molliformis J.Lloyd and Halácsy pointed out its main traits “Panicula magis conferta, ramis brevissime, spiculis densius et longius velutinu-pilosis, aristis demum extrorso subcurvatis”.

Bromus hordeaceus L. subsp. molliformis (J.Lloyd) Maire & Weiller (1955: 255). This post-1953 combination, without direct citation to the basionym, nor to a potentially validating Latin description, but only to “Bromus molliformis Lloyd, Fl. Loire-Inf. 315 (1844)”, is another combination not validly published.

As Bromus intermedius subsp. divaricatus Bonnier & Layens is an untypified name, we choose, as lectotype, the figure in Bonnier and Layens (1894: 369) that is definitely original material. However, as is common with figures, it is difficult to observe some diagnostic characteristics. Thus, we select, as epitype, a specimen collected by Lloyd, conserved in BM, consistent with the protologue. It is a sheet bearing two collections. We choose the specimen on the left from Pornichet (barcode BM001067302).

Figure 5. 

The epitype of Bromus hordeaceus L. subsp divaricatus (Bonnier & Layens) Kerguélen is the specimen on the left with barcode BM001067302 (collected and identified by Lloyd as B. molliformis; preserved at BM (Permanent URL: http://data.nhm.ac.uk/object/9a6e0684-9e9b-4951-ac7e-ba13ad34ed31). Reproduced with permission.

Figure 6. 

Lectotype for Bromus molliformis J.Lloyd ex Billot at P. It is a specimen preserved at the MNHN Collection Vascular plants. Specimen with barcode P02381530. (Available at http://coldb.mnhn.fr/catalognumber/mnhn/p/p02381530) Reproduced with permission.

This taxon has a long history full of nomenclatural problems. Duval-Jouve (1865: 208), who only lists, without naming them, the variations in spikelet size and hairiness of Bromus species, e.g. Bromus mollis “microstachys glabre” and Bromus mollis “microstachys pubescent”. This is the first mention of this taxon. Later Rouy (1913: 236) proposed a named variety under the genus Serrafalcus Parl.: S. mollis ß microstachys Rouy, giving “Bromus microstachys Duval-Jouve” (1865: 207) as a synonym and adding a diagnosis. Therefore, there is no doubt this is the same plant cited by Duval-Jouve (l. c.). Afterwards, Krösche (1924: 329) described B. gracilis Krösche, which unfortunately is a posterior homonym as Weigel (1772: 15) previously proposed this name for a different plant.

Finally, Holmberg (1924: 326) solves this unfortunate situation and names simultaneously the new taxa Bromus lepidus Holmb. and B. lepidus f. lasiolepis Holmb. The herbaria having Holmberg material are K, LD and S. There are 13 sheets in LD collected before 1924, five in S and three in K. It is reasonably certain that Holmberg studied all those plants before the description of the species and form and that all of them are original material. Therefore, we decided to limit our choices to sheets with the annotations “B. lepidus mihi” and “B. lepidus f. lasiolepis Holmb.”, as this annotation indicates that Holmberg is interpreting them as the new taxa he is going to describe.

Figure 7. 

Lectotype of Bromus lepidus Holmberg, preserved at the Herbarium of the biological museum of the Lund University (LD1136295), reproduced with permission.

Figure 8. 

Lectotype of Bromus lepidus f. lasiolepis Holmberg, preserved at the Herbarium of the Biological Museum of Lund University (LD 1136235). Reproduced with permission.

Hudson (1762: 40) proposed the name Bromus ramosus for the wood Brome-grass from England. The protologue is a short diagnosis “BROMUS panicula ramosa nutante scabra, spiculis linearibus decemfloris arista longioribus, foliis scabris” followed by three polynomials as “synonyms”, but without indication of the geographical area in which the new species lives, except an indication to “Anglia”. As pointed out before, the fire destroyed Hudson’s house as well as his personal herbarium. We were not able to find any sheet that could be “original material”. Accordingly, we select a neotype of Bromus ramosus Huds. An annotation label indicates that Carmen Fraile verified and chose it as a neotype in 1994, but this designation was never published (ICN, Art. 7. 10). As this sheet agrees with the protologue, we accept her choice and make it effective here.

Figure 9. 

Neotype selected for Bromus ramosus Huds. preserved at the general collection of the Swedish Museum of Natural History (S) (S-G-1020!). Reproduced with permission.

(designated here): Bromus sterilis L., Sp. Pl.: 77 (1753).

Grisebach (1844: 448) described Bromus sect. Stenobromus including B. maximus Desf., B. madritensis L., B. tectorum L. and B. sterilis L. Hackel (1887: 75) described B. subg. Stenobromus including only two species: B. tectorum L. and B. sterilis L and gave Anisantha K. Koch as a synonym. Afterwards, some other species were included. Until now, B. subg. Stenobromus has been used in some works (Stebbins 1981, Acedo and Llamas 1999) because there is no correct name available for this taxon at subgenus level. As a consequence, B. subg. Stenobromus is treated as a new combination at new rank because a potential basionym exists, B. sect. Stenobromus, applying to the same taxon ICN Art. 41.4.

This subgenus is variable enough to have two sections:

(designated by Tournay 1961: 294): Bromus sterilis L. in Sp. Pl. 77 (1753).

Dumortier (1824) describes the section including Bromus sterilis L., B. tectorum L., B. diandrus Roth, B. rigidus Roth. and B. rigens L., a synonym for B. scoparius L., now belonging to B. sect. Triniusa (Steud.) Nevski.

Europe, temperate Asia and North Africa.

Panicle dense, compact, with short branches. The erect spikelets seem to be sessile. Awns more or less divaricate at maturity.

Bromus rubens L. in Cent. Pl. I: 1 (1755).

B. rubens L., B. fasciculatus C.Presl and B. matritensis L.

Southern Europe, temperate Asia and North Africa. Etymology: Named from the Latin “penicillus”, meaning brush, referring to the morphology of the inflorescence, especially in B. rubens, the name of which provides the type.