Study region

The Ukrainian part of the Carpathians belongs to the Eastern Carpathians. These are medium-high mountains, where the high-mountain zone that includes both subalpine and alpine altitudinal belts accounts for only about 1.7% of the area (Kruhlov 2008). The Ukrainian part of the Carpathians forms a 280 km long and 110 km wide stripe that stretches from the north-west to the south-east. These mountains are built mostly of sandstone flysch while more diverse geological structure occurs in the Maramuresh (aka Marmarosh) Mts at the border with Romania (Kondracki 1989).

Study species

Alopecurus ‌laguriformis is a narrow-ranged alpine grass species, endemic to the Carpathians. According to POWO (2024), it occurs in Romania and Ukraine, while Euro+Med Plantbase (2024) attributes it to Romania and Poland. However, Şerbănescu & Nyárády (1972) consider it endemic to Romania. Since Tzvelev (1971, 1974, 1976) has downgraded the taxon to subspecies rank, i.e. A. ‌pratensissubsp. laguriformis, many sources (e.g. Clarke 1980; Novikov 2023; Euro+Med Plantbase 2024; POWO 2024) follow this approach.

Alopecurus ‌laguriformis is included in all the Soviet (Tzvelev 1974) and Ukrainian (Chopyk 1976) floras, monographs on grasses (Tzvelev 1976; Prokudin et al. 1977), checklists (Tasenkevich 1998; Mosyakin & Fedoronchuk 1999) and identification keys (Chopyk 1977; Dobrochayeva et al. 1987) as well as surveys on the endemics (Stoyko & Tasenkevich 1991, 1993; Tasenkevich 2003; Kliment et al. 2016; Novikoff & Hurdu 2015; Novikov 2023) concerning the territory of the Ukrainian Carpathians. All mentions of its supposed occurrence in Ukraine refer to the locality at the Black Cheremosh River (Tzvelev 1976; Chopyk 1977; Prokudin et al. 1977; Dobrochayeva et al. 1987). However, there are no corresponding vouchers in any Ukrainian herbarium that could confirm that information. Therefore, some authors (Chopyk 1977; Prokudin et al. 1977; Novikov 2023) admit that these data require verification.

According to literature (Schur 1859, 1866; Şerbănescu & Nyárády 1972; Beldie 1979; Ciocârlan 2009; Sârbu et al. 2013) and information from herbarium labels, A. ‌laguriformis is a high-mountain species occurring at the elevation of c. 2000 m a.s.l. in the grasslands near the lakes, streams and springs, as well as in the alpine pastures, rocky sites, and schists.

Lectotypification

Alopecurus ‌laguriformis was first mentioned by Schur (1850) in his list of Transylvanian plants. However, in that publication, the species name is not followed by any description or a diagnosis, nor a reference to a former one effectively published, and thus is considered a nomen nudum (Novikov 2023). The protologue with the first description of the species (Fig. 1) was provided by Grisebach & Schenk (1852: 362) who on their trip to Transylvania were accompanied and consulted by local botanists Schur and Fuss (Speta 1994). Among other information, the protologue points out the following diagnostic characters of the species: ovoid-oblong panicle, basally connate spikelet glumes, and inflated sheath of the upper culm leaf. Later, Schur (1859, 1866) also published his diagnoses of the species.

The protologue (Grisebach & Schenk 1852) mentions the name of Fuss, which indicates that the species was described using the specimens provided by that collector. Herbarium material used by Grisebach and Schenk in their publications is held in the Herbarium Grisebachianum collection at GOET Herbarium, which holds numerous type specimens of these authors (Ahrens 2020).

Noteworthily, there is only one voucher of Alopecurus ‌laguriformis in this herbarium, and it was collected by Fuss in the Arpash Mts, which corresponds to the protologue, however the toponym there is misspelled as “Argash”. Therefore, this specimen fits as the original material for typification.

Family ‌ Poaceae ‌BarnhartGenus ‌ Alopecurus L.

Alopecurus ‌laguriformis ‌Schur ex Griseb. & Schenkee436792-412b-4654-8df7-199c3f031502

Archiv für Naturgeschichte 18 (1): 362 (Grisebach & Schenk 1852). — Alopecurus laguriformis Schur, Verhandlungen und Mitteilungen des Siebenbürgischen Vereins für Naturwissenschaften zu Hermannstadt 1: 182 (Schur 1850), nomen nudum. — Alopecurus ‌laguriformis f. abbreviatus ‌Schur, Österreichische Botanische Zeitschrift 9 (1): 13 (Schur 1859). — Alopecurus ‌pratensissubsp. laguriformis ‌(Schur) Tzvelev, Novosti Sistematiki Vysshykh Rasteniy 8: 19 (Tzvelev 1971).

Alopecurus ‌brachystachyus auct., Linnaea 30: 615 (Janka 1860), non M. Bieb.

Colobachne gerardi sensu Schur, Enumeratio plantarum Transsilvaniae: 728 (1866), non Link.

Type material

Romania • Arpaser Carpaten; s.d.; Fuss s.n.; lectotype (designated here): GOET Herbarium Grisebachianum [GOET064596] (Fig. 2).

Other Historical material

Romania • In alpibus, Frecker Alpe Csortu; 10.VII. sine anno; Fusss.n.; W[W1916-0035870]) • In alpibus Principatis Transilvaniae. Frecker... ? Csortu; 30.VI. sine anno; Fuss s.n.;WU • In alpibus calcareis Transilvaniae. Fundu Buliu Forumei; 7.VIII.sine anno; Fusss.n.;WU • In pascuis alpinis Transsilvan. in monte Arpás; VII.sine anno; Schur s.n.; W[W0025395] • Arpaser Alpen; VII-VIII.sine anno; Schur s.n.; WU • In pascuis alpinis sum. Transsilvan. in monte Podruschel; VII.sine anno; Schur s.n.; WU.

Remarks

Among these vouchers, only two Schur’s specimens were collected in the locus classicus, i.e. in the Arpash Mts, but according to Art. 9.4 of the ICN (Turland et al. 2018), they cannot be considered original material because there is no evidence of their relation to the preparation of Grisebach & Schenk’s (1852) protologue. The same applies to Fuss’s specimens (other than the one from GOET) because they do not come from the locus classicus. Nevertheless, two vouchers in W were labeled as “typus probabiliter” by Pignotti in the 2010s, vizW1916-0035870 collected by Fuss and W0025395 collected by Schur, but for the reasons above they are not eligible for typification.

Revision of the information about Alopecurus ‌laguriformis in the Ukrainian Carpathians

A thorough examination of the information on the distribution of A. ‌laguriformis in the Ukrainian Carpathians showed that it is based on the old record made by Zapałowicz (1889: 342). He claimed that the species sporadically occurs in the Upper Black Cheremosh River Valley on the lush riparious meadows below the inflow of the Balasyniv Stream from 1180 to 1255 m a.s.l. He also mentioned that some plants are over 1 m tall. These data seem doubtful per se because the species is known as a rather short-statured alpine grass (Şerbănescu & Nyárády 1972; Beldie 1979; Ciocârlan 2009; Sârbu et al. 2013), while the above-mentioned altitude refers to the montane forest zone. Thorough surveys performed later in that area (Pawłowski & Walas 1949) did not reveal the species’ occurrence. I also failed to find A. ‌laguriformis during field surveys carried out in that region in 2003-2006 in search of rare, endangered, and endemic species (Kobiv et al. 2007). This area is situated in the Eastern Carpathians, namely in the Chyvchyny Mts that belong to the Maramuresh orographic unit (Breman et al. 2020), which Ukraine shares with Romania. In the 1920-1930s, the region belonged to Poland, while still earlier – to the Austro-Hungarian Empire.

In his later publication, Zapałowicz (1906) referred to that finding as “A. ‌brachystachyus ‌M. Bieb.” synonymizing it with A. ‌laguriformis. Such a nomenclatural approach was also used by Szafer (1919) in the ‘Flora of Poland’ and Szafer et al. (1924) in the first edition of ‘Polish plants…’ concerning the plants from the same locality in the Chyvchyny Mts.

Synonymization of A. ‌laguriformis with A. ‌brachystachyus is based on Janka’s (1860) publication, where he also considers it conspecific with A. ‌castellanus ‌Boiss. & Reut. described from the Pyrenean Peninsula, which is now regarded as a synonym for A. ‌arundinaceus ‌Poir. (Aedo 2020). In the late 19th century, that treatment was accepted by a number of European authors (e.g. Nyman 1882; Ascherson & Graebner 1898). However, Hackel (1902) proved the difference between these species, and it is currently accepted that the range of A. ‌brachystachyus does not extend beyond Asia, while A. ‌laguriformis is restricted to the Carpathians (Tzvelev 1971; Clarke 1980).

Much confusion about the correct circumscription of Alopecurus ‌laguriformis emerged from Schur’s (1859, 1866) publications. It should be admitted that Schur was keen on describing new species and intraspecific taxa or just naming them (i.e. assigning nomina nuda) often without strong reasons, which also concerns the genus Alopecurus. For instance, A. ‌altissimus ‌Schur (1859, 1866) was later synonymized with A. ‌pratensis (Pawłowski 1939; Şerbănescu & Nyárády 1972). An example of a name misapplied by Schur is “Colobachne ‌gerardii ‌Link.” Currently, the accepted name for it is Alopecurus ‌gerardi ‌Vill., but that species is distributed in the mountains of Central and Southern Europe and does not occur in the Carpathians (Clarke 1980). That Schur’s record proved to be Alopecurus ‌laguriformis ‌(Şerbănescu & Nyárády 1972). Interestingly, both these species are mentioned by Schur (1866: 727-728) next to each other as separate taxa, but gathered in the same location. That shows that some of Schur’s taxonomic referrals need critical revision.

Schur’s circumscription of A. ‌laguriformis is rather ambiguous and assumes its significant intraspecific diversity, which is not implied in Grisebach & Schenk’s (1852) protologue. On the one hand, Schur (1859, 1866) described a typical A.laguriformis, which in the first publication (Schur 1859) he also considered an intraspecific form, viz. f. abbreviatus. It is characterized by 6-12 line (1.2-2.5 cm) long, ovoid-oblong, compressed at base panicle; culms 8-15 inch (20-40 cm) long, which fits the protologue and most of modern descriptions of A. ‌laguriformis (Şerbănescu & Nyárády 1972; Prokudin et al. 1977; Clarke 1980).

In addition, Schur (1859, 1866) described f. elongatuswith much longer, 1.5-3 inch (c. 4-7.5 cm) long, cylindrically elongated, attenuate at base panicle and culms 2-3 ft (c. 60-90 cm) long.

These taxa significantly differ even in their habit, while the description of the latter one contradicts the protologue, namely concerning the shape of the panicle, which is supposed to be ovoid-oblong (Grisebach & Schenk 1852). The main reason why such different entities were treated as the same species is based on the assumption that in both of them the glumes are connate only at base while in A. ‌pratensis they are connate for 1/3 to 1/2 of their length (Szafer 1919; Şerbănescu & Nyárády 1972; Ciocârlan 2009).

Locus classicus of the typical A. ‌laguriformis is Mt. Arpash in the Fagaras Massif, Southern Carpathians, while f. elongatus was first described by Schur (1859) from Mt. Piscu in the same massif but later that author referred it to Mt. Corongisu in the Rodna Mts, Eastern Carpathians (Schur 1866).

Kulczyński (1932) and Pawłowski (1939), who examined the original Schur’s herbarium specimen of A.laguriformis f. elongatus in LW herbarium (unfortunately, missing now), found out that it should be identified as A. ‌pratensis ‌sensu stricto. According to Kulczyński (1932: 5), it has all the characters of the typical A. ‌pratensis concerning dissection, indumentum, and apexes of glumes as well as the lack of inflation of leaf sheaths.

Confusion about the circumscription of A. ‌laguriformis is revealed in the herbarium exsiccata distributed among the main European herbaria at the beginning of the 20th century. It concerns “Gramina Hungarica”, where issue #306, collected by Zsák in 1910 in the Fagaras Mts fits the species description, whereas issue #108 labeled as A. ‌laguriformisf. elongatus and collected by Degen in 1902 in the Rodna Mts proved to be A. ‌pratensis. These plants are over 60 cm high, with oblong-cylindrical c. 5 cm long panicles and strongly connate glumes that does not match the protologue of A. ‌laguriformis ‌(Grisebach & Schenk 1852). They were collected on Mt. Craciunel, Mt. Cisia, and Mt. Corongisu in the Rodna Massif, while the latter locality is the site where Schur (1866) reported A. ‌laguriformis f. elongatus, which could be the reason for that misidentification.

A similar situation occurred with Porcius’s (1913: 129) contribution to “Flora Exsiccata Austro-Hungarica”. His issue of the herbarium specimens #3997, which allegedly presented A. ‌brachystachyus and were synonymized with A. ‌laguriformis, have been later re-identified as A. ‌pratensis ‌sensu stricto. They were collected on Mt. Craciunel in the Rodna Massif.

Nevertheless, issue #1155 of A. ‌laguriformis from “Flora Romaniae Exsiccata“ collected in the Retezat Massif, Southern Carpathians, and distributed by Borza and Nyárády in the 1930s (a duplicate occurs at P [P02222096]) matches the description of that species.

The above review shows that there was much ambiguity in the understanding of A. ‌laguriformis by botanists in the late 19th-early 20th century, which might have influenced Zapałowicz’s (1889, 1906) reports on the species’ occurrence in the Chyvchyny Mts. The issue of the real identity of these findings could be best resolved by examining his original herbarium material.

Fortunately, I discovered two vouchers gathered by Zapałowicz in the KRAM herbarium and labeled by him as “Alopecurus ‌laguriformis ‌Schur” and “Alopecurus ‌brachyctachyus ‌M. Bieb.” as synonyms, which clearly correspond to the location mentioned by that author (Zapałowicz 1889, 1906). The first specimen (KRAM0003667-V) comes from “the Upper Black Cheremosh” and dates to August 1881 (Fig. 3). Another one (KRAM0003666-V) was also collected at the Black Cheremosh River “below the inflow of the Popadia Stream” in August 1906. Evidently, based on these very findings Zapałowicz (1889, 1906) claimed that Alopecurus ‌laguriformis occurs in the Chyvchyny Mts. Moreover, there is a mark on the label of the first voucher, which states that it was cited in Zapałowicz’s (1906) “Conspectus…” In fact, both specimens proved to be A. ‌pratensis. Moreover, in the ‘Atlas of the Polish Flora’ Kulczyński (1932) used Zapałowicz’s herbarium specimen KRAM0003667-V as one of the exemplary illustrations of A. ‌pratensis. The icons, prepared by Baecker (Fig. 4) depict the habit and morphological characters, including oblong-cylindrical panicles and strongly connate short-ciliate glumes peculiar to A. ‌pratensis.

Therefore, Kulczyński (1932: 3) denied the occurrence of A. ‌laguriformis in the Chyvchyny Mts and pointed out that the species had not been found in the territory of Poland, which included that area in the 1920-1930s. The Atlas also contains the illustration of L. laguriformis with several icons, which depict its distinguishing characters accompanied by textual explanations (Kulczyński 1932: 4).

Pawłowski, who performed comprehensive studies of the flora and vegetation of the Chyvchyny in the 1930s and paid special attention to the distribution of endemics in that region (Pawłowski 1947; Pawłowski & Walas 1949), did not mention A. ‌laguriformis. However, in his publication on floristic records from Romania, Pawłowski (1939) gives a rather detailed description of A. ‌laguriformis in comparison to A. ‌pratensis and A. ‌arundinaceus accompanied by illustrations of their spikelets and lemmas. Defining the distribution of the former species he refers only to the Romanian Carpathians. Examining of the available herbarium vouchers of A. ‌laguriformis carried out within the present study confirms that opinion stated also by Şerbănescu & Nyárády (1972).

As follows from the above, the information about occurrence of A. ‌laguriformis in the Ukrainian Carpathians proved to be incorrect.

Alopecurus ‌laguriformis is a distinctive species clearly restricted to the alpine zone and well-distinguishable from A. ‌pratensis by the shape of panicle (ovoid-oblong, < 3 cm long in A. ‌laguriformis vs cylindrical, > 4 cm long), height (< 0.5 m vs > 0.5 m), glume pubescence (hairs 2-3 mm vs < 2 mm) and connation of glumes (connate only at the base vs connate to c. ⅓-½ of their length) and culm leaf sheath (inflated vs non-inflated).

The degree of connation of glumes is a taxonomically important character in the genus Alopecurus, which is used in the infrageneric identification keys, e.g. by Tzvelev (1971, 1974, 1976) and Clarke (1980), however these authors did not apply it to distinguish between A. ‌pratensis and A. ‌laguriformis and have downgraded the latter taxon to subspecies rank, i.e. A. ‌pratensissubsp. laguriformis, which does not seem reasonable. On the other hand, in Romania, where the species occurs, floras and identification keys (Şerbănescu & Nyárády 1972; Ciocârlan 2009; Sârbu et al. 2013) emphasize that character as distinguishing for A. ‌laguriformis. Most importantly, it is highlighted in the species’ protologue (Grisebach & Schenk 1852) as well as in other publications (Szafer et al. 1924; Kulczyński 1932; Pawłowski 1939; Chopyk 1977).