Typification of names published by P. S. Pallas in Astragalus and Phaca, now in genus Oxytropis sect. Orobia (Fabaceae)

Sytin A. K.; Byalt V. V.; Sokolova I. V.

Новости систематики высших растений Novitates Systematicae Plantarum Vascularium

2023

54:41-52

ISSN 0568-5443 (print) ISSN 2687-1564 (online)

Typification of names published by P. S. Pallas in Astragalus and Phaca, now in genus Oxytropis sect. Orobia (Fabaceae)

Типификация названий астрагалов (Astragalus, Phaca), описанных П. С. Палласом, в настоящее время относящихся к секции Orobia рода Oxytropis (Fabaceae)

A. K. Sytin, V. V. Byalt, I. V. Sokolova*

Komarov Botanical Institute of the Russian Academy of Sciences Herbarium of Higher Plants

Professora Popova Str., 2, St. Petersburg, 197022, Russia [email protected]; [email protected]; [email protected] *Corresponding author

https://doi.org/10.31111/novitates/2023.54.03

А. К. Сытин, В. В. Бялт, И. В. Соколова*

Ботанический институт им. В. Л. Комарова РАН Гербарий высших растений

ул. Профессора Попова, 2, Санкт-Петербург, 197022, Россия [email protected]; [email protected]; [email protected] *Автор для переписки

Abstract. The original material used by P. S. Pallas in his descriptions of 7 species in the genus Astragalus L. and 1 species in Phaca L., all now classified within the genus Oxytropis DC. section Orobia Bunge, is analysed and discussed. The lectotypes of Astragalus altaicus Pall. (Oxytropis altaica (Pall.) Pers.), A. argentatus Pall. (O. argentata (Pall.) Pers.), A. candicans Pall. (O. candicans (Pall.) DC.), A. grandiflorus Pall. (O. grandiflora (Pall.) DC.), A. songaricus Pall. (O. songarica (Pall.) DC.), A. spi-catus Pall. (O. spicata (Pall.) O. Fedtsch. et B. Fedtsch.), Phaca sylvatica Pall. (O. sylvatica (Pall.) Willd.), and the epitype of Astragalus spicatus (O. spicata) are designated.

Keywords: Astragalus, Phaca, Oxytropis, section Orobia, Fabaceae, Pallas, typification, lectotype, epitype.

Аннотация. В статье рассматривается первоначальный материал названий 7 видов, описанных П. С. Палласом в роде Astragalus L., и 1 вида, описанного в роде Phaca L., в настоящее время относящихся к секции Orobia Bunge рода Oxytropis DC. Обозначены лектотипы Astragalus altaicus Pall. (Oxytropis altaica (Pall.) Pers.), A. argentatus Pall. (O. argentata (Pall.) Pers.), A. candicans Pall. (O. candicans (Pall.) DC.), A. grandiflorus Pall. (O. grandiflora (Pall.) DC.), A. songaricus Pall. (O. songarica (Pall.) DC.), A. spicatus Pall. (O. spicata (Pall.) O. Fedtsch. et B. Fedtsch.), Phaca sylvatica Pall. (O. sylvatica (Pall.) Willd.) и эпитип Astragalus spicatus (O. spicata).

Ключевые слова: Astragalus, Phaca, Oxytropis, секция Orobia, Fabaceae, Паллас, типификация, лектотип, эпитип.

Introduction

This research completes the work by the first two authors (Byalt, Sytin, 2022) on typification of the names of the taxa originally published by Peter Simon Pallas (1741-1811) within the genera Astragalus L. and Phaca L. (Pallas, 1773b, 1776, 1800-1803), and later transferred to the genus Oxytropis DC. (Can-dolle, 1802; Persoon, 1807; Fedtschenko, Fedtschenko, 1909).

The names of Astragalus later transferred to the sections Arctobia Bunge, Baicalia Steller ex Bunge, Gloe-ocephala Bunge, Janthina Bunge, Ortholoma Bunge, Polyadena Bunge, and Xerobia Bunge of the genus Oxytropis were typified in the previous publication (Byalt, Sytin, 2022). The aim of this research is typification of the names of the species assigned to the section Orobia

Bunge, one of the largest and taxonomically complicated groups within Oxytropis.

Historical background

Pallas described a large number of plant taxa from European part of Russian Empire (Russia, Ukraine, West Kazakhstan), and Northern Asia (Siberia, Kazakhstan, Russian Far East). He was particularly interested in the genus Astragalus s. l., that is especially diverse in South Siberia, and described up to a hundred species of this group. Some of them were published in "Reise durch verschiedene Provinzen des rußischen Reichs" (Pallas, 1771-1776); these and many more newly described ones were included in "Species Astragalorum..." (Pallas, 1800-1803).

The genus Oxytropis was established by Augustin Pyramus de Candolle (1778-1841) based on its

Поступила в редакцию | Submitted: 26.06.2023 Принята к публикации | Accepted: 05.12.2023

Опубликована онлайн | Published online: 17.12.2023 (Страницы | Pages: e03:1-12)

characteristic subulate outgrowth at the keel top: "Carina in mucronem superne desinens" (Candolle, 1802: 661). Currently the genus Oxytropis comprises about 600 species (POWO, 2023), including 25 species described by Pallas as Astragalus, and 7 ones as Phaca. It is noteworthy that Pallas mentioned acuminate keels as a characteristic of certain species ("carina cum acumine", "carina acuminata" or "carina acumine sursum mucronata"), but not of the genus subdivisions. Nevertheless, most of Pallas' species which are now treated as Oxytropis fell into two "phalanges" within his classification of Astragalus: "Astragali anthylloidei" and "Astragali polypterophylli seu verticillares" (Pallas, 1800-1803).

An outline of taxonomic history of Oxytropis including references to some important publications was discussed previously by Byalt and Sytin (2022).

"Species Astragalorum" (Pallas, 1800-1803) was published during 3 years in separate fascicles, totalling 13. At the same time, Candolle and C. L. Willdenow also studied Astragalus and published respective fundamental treatments almost simultaneously (Candolle, 1802; Willdenow, 1802). Clearly, some names published in these three works may prove to be competing synonyms. To resolve the issues of priority, it is necessary to know the precise dates of publication. Those of Willde-now's and Candolle's works are listed in "Taxonomic literature": November 1-10, 1802 (Stafleu, Cowan, 1988: 303) and November 15, 1802 (Stafleu, Cowan, 1976: 442), respectively, in contrast to "Species Astragalorum". In the 19th century, the books issued in separate parts were traditionally binded by their owners after obtaining the last part, usually with title pages removed. This is the most probable cause of the absence of precise dates of publication for the fascicles of "Species Astragalorum" given in "Taxonomic literature" (Stafleu, Cowan, 1983: 26). P. Perret (1989) determined these dates from indirect evidence, namely, whether certain parts of "Species Astragalorum" were cited by Candolle (1802) and Willdenow (1802). However, copies of "Species Astragalo-rum" with preserved titles of the fascicles exist, e. g. the one available on the website of the Digital library (repository) of the Tomsk State University (Digital library..., 2020: https://vital.lib.tsu.ru/vital/access/manager/Re-pository/vtls:000438986). The comparison of the dates specified by Perret (1989) and printed on the fascicle title pages is presented in the Table. It is disputable, which dates are accurate. It is worth noting that J. G. Georgi (1802: 295) mentioned that he had 9 fascicles of Pallas'

Table. Dates of publication of separate fascicles of "Species Astragalorum" (Pallas, 1800-1803).

Text pages Date

Fascicle Plates Fascicle Perret

title (1989)

I, II viii; 1-16 I-XIV 1800 1800[-1801, before Apr 4]

III, IV 17-32 XV-XXVI 1800 1800[-1801, before Apr 4]

V, VI 33-52 XXVII-XLI 1800 -

VII, VIII, IX 53-76 XLII-LXII 1801 1802 [before Sept-Nov]

X, XI 77-92 LXIII-LXXVI 1801 1802 [before Nov] 1803

XII, XIII 93-124 LXXVII-XCI 1802 [before the end of Feb]

1 The Candolle's "Astragalogia" was simultaneously published in two formats: in folio and in quarto (Stafleu, Cowan, 1976: 441-442). Here and further, we refer to the pages of "Astragalogia" according to the quarto issue, as more accessible (Candolle, 1802).

work with 96 copperplate engravings in January 1802. For ease of search, in this paper we cite the years of publication as specified in appropriate fascicle titles and refer to the fascicle number before the page number.

In relation to the typification of the Pallas' names, the question arises as to the reliability of the images published in "Species Astragalorum". The artist Christian Gottfried Heinrich Geißler (1770-1844) accompanied Pallas during his travel about southern European Russia in 1793-1794, and then served as the manager of his Crimean estate Shulyu in 1795-1798 (Sytin, 2014: 202, 234). Geißler illustrated "Species Astragalorum" and other Pallas' works (Pallas, 1803-1806, 1811-1814), as well as prepared illustrations for the unpublished "Plan-tae selectae rossicae" (see Sytin, 1994) but he did not participate in the travels of 1768-1774, so could not see the plants collected at that time alive. In the preface to his work, Pallas wrote: "Icones omnes et descriptiones ad plantas spontaneas vivas, quamplurimis plerumque speciminibus collatis, paravi, paucas ad specimina sicca, optime asservata et plerumque multiplicata [I have prepared all the icons and descriptions from living spontaneous plants, generally from numerous collated specimens, a few ones from dry specimens, best preserved and generally multiplied]" (Pallas, 1800-1803: viii). On the other hand, he admitted that not every species could be successfully cultivated: "...in hortis autem bene multae species omnem culturae diligentiam frustrant, et quae succedunt, in ollis, ut experientia edoctus sum, humo valde arenosa plenis curandae sunt [in gardens, however, many species negate all care of cultivation, and those that succeed, as I have learned from experience, are to be cared for in pots full of very sandy soil]" (Pallas, 1800-1803: v). From among the species of the section Orobia, Astragalus altaicus Pall. and A. ambiguus

Pall. were mentioned as successfully grown in St. Petersburg. A striking resemblance of many images in "Species Astragalorum" to preserved herbarium specimens suggests that the former were drawn from the latter, e. g. Plates XLVI (A. grandiflorus Pall.), XLVIII (A. argenta-tus Pall.), XLIX (A. candicans Pall.), and LI (A. songari-cus Pall.). Thus, this is a direct evidence of the relevant specimens being a part of original material, but leaves it open whether the plants were collected by Pallas in the wild or subsequently grown in a garden.

As concerns the illustrations in "Reise..." (Pallas, 1771-1776), most copies of this work are provided with black-and-white copper engravings. We only know of one copy with the illustrations hand-painted with watercolours, namely the one stored at the National Library of Russia, in the "Rossica" collection (Figs. 1; 2: B). It is noteworthy that on the plate with Astragalus spicatus Pall. (Pallas, 1773b: tab. W) Pallas left a critical comment on the image colour (Fig. 1: B). The margin of the plate was probably cut off when the volume was rebound; the text, however, is quite restorable: "Die Untertheile der Blumen grünlich grau, nur die Spitzen der Knospen röthlich und viel blauer als hier [The lower parts of the flowers are greenish grey, only the bud tips are reddish and much bluer than here]". This Pallas' note is the evidence of both his high demands on the quality of illustrations, and particular attention to the corolla colour as a key taxonomic character.

Material and methods

Typification of Pallas' species names is quite a complicated task, since his original material is dispersed among different Herbaria, generally imprecisely labelled, and some of the specimens are damaged with time. The most extensive and valuable part of Pallas' collections is kept in the Natural History Museum, London (BM — herbarium acronyms follow Thiers, 2023). Some materials are stored in the Komarov Botanical Institute of RAS, St. Petersburg (LE), with small numbers in the Herbaria of the Botanische Staatssammlung München (M), the Botanic Garden and Botanical Museum Berlin-Dahlem (B), and others. The history of Pallas' herbarium collections is described in detail (Lambert, 1811; Miller, 1970; Belyaeva, Sennikov, 2008; Sytin, 2014).

Many of Pallas' herbarium collections are available on the Internet resources "Global Plants on JSTOR" (2023), and JACQ (2004) which greatly facilitated our

Fig. 1. Lectotype of Astragalus spicatus (Pallas, 1773b: tab. W; scanned image provided by the National Library of Russia).

A — image of the plant; B — fragment of the plate with a pencil note by P. S. Pallas.

J'alàts Ttebs'ert Tom. JI. 7hi. f*1

À

Fig. 2. Illustration from the protologue of Phaca sylvatica (Pallas, 1776: tab. W, fig. 2; scanned image provided by the National Library of Russia), and herbarium specimens BM000958893 and M-0185384 (lectotype). A — herbarium specimen BM000958893 (https://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen. bm000958893); B — illustration from the protologue; C — herbarium specimen M-0185384 (https://plants.jstor. org/stable/viewer/10.5555/al.ap.specimen.m0185384). Equal numbers indicate corresponding details of the image and plants.

work. The images of the scanned specimens stored at LE are available on the website Herbarium LE (Vascular Plants..., 2023).

The handwriting on each label was identified using the images available in S. Lipschitz and I. T. Vassilczenko (1968) and scanned images of handwritten materials available on the Alvin platform (Alvin, 2023). The latter source, in particular, allowed us to attribute some annotations on Pallas' herbarium sheets at BM to A. B. Lambert (Fig. 3: C, D). The handwriting of C. H. Merck (1761-1799) on the specimen BM000958880 was identified by comparing it with Fig. 489 in "Die Große Nordische Expedition" (1996: 323).

For each specimen cited, we indicate its category (lec-totype, isolectotype, syntype) in accordance with "International Code of Nomenclature" (Turland et al., 2018: Art. 9), or nominate it as an "original specimen" for the specimens belonging to the original material but not cited in the protologue. The provenance of the specimens is cited in abbreviated form according to the "Catalogue of the type specimens of the vascular plants from Siberia and the Russian Far East kept in the Herbarium of the

Komarov Botanical Institute (LE)" (2012). Then textual quotations of the labels follow: the annotations are preceded by authors' names (if identified) in square brackets; the ends of lines in original labels are marked by vertical bars; the text crossed out in the original labels is crossed out in the quotations. Also placed in square brackets are notes and indications of phenological stages of the plants. The barcode numbers of the specimens are included.

Results and discussion

Astragalus altaicus Pall. 1801, Sp. Astragal. VII: 56, tab. XLV.

= Oxytropis altaica (Pall.) Pers. 1807, Syn. Pl. 2, 2: 333.

The specimen LE 01056519 collected by Pallas' correspondent, Petr Ivanovich Shangin, (1745/1748-1816) in the Altai is designated here as the lectotype of Astragalus altaicus. It has an original label written by Shangin with indication of the locality corresponding to the pro-tologue, and the species name in Pallas' handwriting.

Lectotype (Sytin, Byalt, designated here): W Sib., Altai Territory, "[Pallas:] Astragalus montanus

Fig. 3. Illustration from the protologue of Astragalusgrandiflorus (Pallas, 1800-1803: tab. XLVI; courtesy of the Research Library of Tomsk State University) and the herbarium specimen BM000958878 (lectotype). A — illustration from the protologue; B — herbarium specimen (https://plants.jstor.org/stable/viewer/10.5555/ al.ap.specimen.bm000958878), equal numbers indicate corresponding details of the image and plant; C, D — enlarged fragments of the herbarium sheet with inscriptions by A. B. Lambert.

There is some possibility that other specimens which were added to the collection of LE

from F. E. L. Fischer's herbarium are duplicates of LE 01056519 or LE 01056518, i. e. parts of the original material. With some doubt, we also list here the herbarium specimen of the plant grown from the seeds gathered by L. Patrin during his travels around the Altai in the 1780s (BM000958869), since in the protologue the plant(s) growing in a garden in St. Petersburg is(are) mentioned.

?Original specimen: W Sib., Altai Territory / E Kazakhstan, "[Shangin:] Astragalus altaicus |

Растет по всем алтайским | горам во множестве [Occurs abundantly throughout the Altai mountains] | [Fischer:] Schangin | [unknown handwriting:] Herb. Fischer [fl., fr.]" (LE 01056517!).

Note. The specimens LE 01056517, LE 01056518 and LE 01056519 are mounted on the same sheet.

?Original specimen: W Sib., Altai Territory / E Kazakhstan, "[unknown handwriting:] Herb. Fischer | [Fischer:] altaica | Schangin [fl., fr.]" (LE 01056528!).

?Original specimens: W Sib., Altai Territory / E Kazakhstan, "[unknown handwriting:] Herb. Fischer | [Fischer:] Schangin [fl., fr.]" (LE 01056520!, LE 01056521!).

?Original specimen: cultivated, "[unknown handwriting:] E semine Patrinii in hort. botan. floruit. [fl., fr.]"; "[Lambert:] Herb. Pallas"; on the sheet, in pencil: "[unknown handwriting:] Oxytropis brevi-rostris Dec. [Lambert:] Astragalus Altaicus Pall. | Lin. Willd." (BM000958869 — image!).

Protologue quotation: "Astragalus altaicus. Tab. XLV... In alpibus altaicis, praesertim supra Tigeraek, itemque ad Ieniseam, circa lacum Utschum et Abacan fl. frequens planta, alibi vix occurrit et specie distinctis-sima est... Petropoli in hortis optime viget et perennat".

Astragalus ambiguus Pall. 1801, Sp. Astragal. VII: 54, tab. XLIII, XLIII.B.

= Oxytropis ambigua (Pall.) DC. 1802, Astragalo-gia: 70.

None of the specimens known to us bears the name Astragalus ambiguus written in Pallas' handwriting. The specimen BM000958874 is quite well preserved and has an original Pallas' label "ad Altaicos montes". Its provenance and identity, however, are doubtful. The plant's robust habit shows more resemblance to the Plate XLIII.B (Astragalus ambiguus horten-sis) than to XLIII (Astragalus ambiguus spontaneus) in Pallas' publication (1800-1803), which suggests that it was probably grown in a garden rather than collected in the wild. Moreover, the name Astragalus ambiguus is written on the sheet by Lambert, not by Pallas, and Altai is not mentioned in the protologue (Arzamas and the Bolshoi Cheremshan River are located in the European Russia). So, we cannot be certain that Pallas treated this specimen as Astragalus ambiguus. One of the four labels mounted on the sheet LE 01051468 has an annotation by Pallas "Astragalus uralensis", as do the labels by Shangin and Fischer. The name Oxytropis ambigua on the specimens LE 01051470 and LE 01051471 is written later. So, despite Pallas' remark about similar habit of Astragalus ambiguus and A. uralensis, we cannot confidently

ascribe these specimens to the original material, and refrain from designating the lectotype.

?Original specimen: W Sib., Altai Territory / E Kazakhstan, "[Pallas:] ad Altaicos montes [fl.]"; "[Lambert:] Herb. Pallas"; on the sheet, in pencil: "[unknown handwriting:] Oxytropis ambigua Dec. [Lambert:] Astragalus ambiguus Pall. | Lin. Willd." (BM000958874 — image!).

Note. There are 4 plants and 4 labels mounted on the sheet but it is not possible to find out which plant corresponds to each label.

?Original specimen: W Sib., Altai Territory / E Kazakhstan, "[unknown handwriting:] Herb. Fischer | [Fischer:] ambigua | Schangin | [Bunge:] (Bge!) [fl.]" (LE 01051471!).

Protologue quotation: "Habitu proxime similis A. Uralensi... Crescit haec species frequens per omnem Sibiriam, imo passim in Rossia ad orientem Moscuae occurrit, mihi v. gr. lecta in sylvis circa Arsamas et ad fluv. Tscheremschan... Habeo varietatem valde luxuri-antem... Petropoli in hortis optime viget, perennat et floret Majo".

Astragalus argentatus Pall. 1801, Sp. Astragal. VII: 60, tab. XLVIII.

= Oxytropis argentata (Pall.) Pers. 1807, Syn. Pl. 2, 2: 331.

The Pallas' specimen stored at BM (BM000958881) matches the provenance specified in the protologue, and most probably was the plant from which Plate XLVIII in "Species Astragalorum" (Pallas, 1800-1803) was drawn. Here, we designate it as the lectotype of Astragalus ar-gentatus.

Lectotype (Sytin, Byalt, designated here): E Kazakhstan, "[Pallas:] Astragalus incanus | Incipis Kirgisen [fl.]"; "[Pallas:] E des[ertis]. kirgis[icis]. Sievers"; "[Lambert:] Herb. Pallas" (BM: BM000958881 — image!).

Two specimens stored in the type collection of LE with labels: (1) "[unknown handwriting:] Herb. Fischer [Fischer:] argentata | Schangin" and (2) "[label 1,

unknown handwriting:] Herb. Fischer | [Fischer:] Schangin | [unknown handwriting:] O. argyracea", "[label 2, unknown handwriting:] Herb. Fischer | [Fischer:] Schangin" are listed as types by T. Krestovskaya and V. Byalt (1998) mistakenly, as Shangin was not mentioned in the protologue.

Protologue quotation: "Astragalus argentatus... Astragalus incanus, Sievers adnot. MS. ...Hanc plantam in desertis elatis Kirgisiorum, circa Irtin fl. et Altaica pro-montoria legit Diligentiss. Joh. Sievers".

Astragalus candicans Pall. 1801, Sp. Astragal. VIII: 61, tab. XLIX.

= Oxytropis candicans (Pall.) DC. 1802, Astragalo-gia: 72.

The specimen stored at BM (BM000958880) is designated here as the lectotype of Astragalus candicans. Its provenance specified on both labels corresponds to the protologue. According to I. P. Borodin (1908), Merck lived in Irkutsk and visited Yakutsk, among other places. The village "Chotingtumul" (Khatyng-Tumul) is located on the Lena River, about 450 km upriver from Yakutsk. When Merck travelled from Irkutsk down the Lena, he inevitably passed Khatyng-Tumul. Merck's annotation "in trockener schwarzer Erde zwischen Steinen" corresponds to the phrase in the protologue: "locis saxosis, humo tectis, siccioribus". In addition, this plant was obviously represented in Table XLIX of "Species Astragalorum" (Pallas, 18001803), as evidenced by the resemblance of the specimen to the illustration, as well as the annotation "De-lin. [depicted]" on its label.

Lectotype (Sytin, Byalt, designated here): E Sib., Yakutia, "[Pallas:] Astragalus candicans | Ex orientali Sibiria | Delin. [fl.]"; "[Lambert:] Herb. Pallas"; "[Merck:] № 41. | Bey dem dorpfe Chotingtumul | des Flusses Lena, am Ufer in trockener | schwarzer Erde zwischen Steinen" (BM: BM000958880 — image!).

Protologue quotation: "Astragalus candicans. Tab. XLIX... Provenit haec species ad Lenam fl. et in maxime orientalibus Sibiriae, locis saxosis, humo tectis, siccioribus. Lecta a D. D. Merk [sic!]".

Astragalus grandiflorus Pall. 1801, Sp. Astragal. VII: 57, tab. XLVI.

= Oxytropis grandiflora (Pall.) DC. 1802, Astragalo-gia: 71.

The Pallas' specimen from East Siberia stored at BM (BM000958878) is designated here as the lec-totype of Astragalus grandiflorus. Apparently, it was this plant from which the illustration on Plate XLVI of "Species Astragalorum" (Pallas, 1800-1803) was drawn. With time, the root and caudex were lost,

the shoots fragmented, but the specimen still demonstrates its resemblance with the image (Fig. 3: A, B).

Lectotype (Sytin, Byalt, designated here): E Sib., "[Lambert:] Herb. Pallas. [fl.]"; on the sheet, in pencil: "[Lambert:] Astragalus grandiflorus Pall. | Lin. Willd. | Oxytropis grandiflora Decand. 103" (BM: BM000958878 — image!).

Protologue quotation: "Astragalus grandiflorus. Tab. XLVI... a Jenisea fluv. per omnem ulteriorem Si-biriam, praesertim in Trans-Baicalensi regione aprica...".

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Astragalus songaricus Pall. 1801, Sp. Astragal. VIII: 63, tab. LI.

= Oxytropis songarica (Pall.) DC. 1802, Astragalo-gia: 73.

The Pallas' specimen from East Siberia stored at BM (BM000958875) is designated here as the lectotype of Astragalus songaricus. We believe that it was the plant from which the illustration in "Species Astragalorum" (Pallas, 1800-1803: tab. LI) was drawn, as evidenced by its undeniable external resemblance to the image, as well as by Pallas' annotation "Delin. [depicted]" on the original label. To note, the inflorescence taking the central position in the image is now separated from the plant and mounted to the sheet on its right side, and its former place is taken by another, shorter inflorescence (Fig. 4). Another Pallas' specimen stored at BM (BM000958876) is treated here as a syntype.

A specimen of this taxon from the collection of Johann Friedrich Pott (1738-1805) sent to him in 1789 by Georg Lindenthal (1749-?) is stored at LE. Lindenthal worked in Altai, and in particular led one of the parties of the "gem travel" (an expedition of 1786 in Altai, mainly aimed at search for deposits of ores and gemstones — see Harms, 2013), which explored Altai mountains along with P. I. Shangin's party, and was undoubtedly acquainted with Shangin, so there is a small possibility that this specimen is a duplicate of a collection sent by Shangin to Pallas, i. e. part of the original material.

The other specimens stored in the type collection at LE and listed by Krestovskaya and Byalt (1998) were collected later and do not belong to the original material.

M. S. Knjasev (2001: 141) had not seen the herbarium specimens of Oxytropis songarica that belong to the original material and proposed that the type of this name should be the illustration in Pallas' monograph (Pallas, 1800-1803: tab. LI). The typification, however, was not effected by him.

Lectotype (Sytin, Byalt, designated here): W Sib., Altai Territory / Khakassia, "[Pallas:] Astragalus Sieversianus | Delin. [alab., fl.]"; "[Pallas:] Sievers"; "[Lambert:] Herb. Pallas"; on the sheet, in pencil: "[unknown handwriting:] Astragalus songaricus.

Fig. 4. Illustration from the protologue of Astragalus songaricus (Pallas, 1800-1803: tab. LI) and the herbarium specimen BM000958875 (lectotype).

A — illustration from the protologue (courtesy of the Research Library of Tomsk State University); B — herbarium specimen (https://plants.jstor.org/stable/viewer/10.5555/al.ap.specimen.bm000958875).

Pall. | Lin. Willd. | Oxytropis songarica. Decand." (BM: BM000958875 — image!).

Original specimens: W Sib., Altai Territory, "Locis montosis et apricis | ad Tigeräk, Schangin [fl., fr.]"; "[Lambert:] Herb. Pallas" (BM000958876 — image!).

Note. There are two plants mounted on the sheet BM000958876. Judging from their phenological stages (left plant with flowers, right plant with ripened fruits) they represent different gatherings. There is also a paper capsule with seeds labelled: "Astragalus Sibiriae uralensi potius". Most probably, it is not related to these plants.

?Original specimen: W Sib., ?Altai Territory, "[Pott:] Astragalus songaricus Pall. Astrag. 4 | Hab. in Sibiriae campis elatis circa promontoria | Altaica

| Ibidem lectum Dn. de Lindenthal misit 1789 [alab., fl.]" (LE 01083347!).

Protologue quotation: "Astragalus songaricus. Tab. LI... Provenit in campis elatis circa Promontoria Altaica, missa a diligentissimis Florae Sibiricae scrutatoribus P. Schangin et Jo. Sievers".

Astragalus spicatus Pall. 1773, Reise Russ. Reich. 2, 2: 742, tab. W.

= Astragalus caudatus Pall. 1801, Sp. Astragal. VIII: 62, tab. L, pro nom. nov., illeg., superfl.

= Oxytropis caudata DC. 1802, Astragalogia: 73, nom. illeg.

= Oxytropis spicata (Pall.) O. Fedtsch. et B. Fedtsch. 1909, Consp. Fl. Turkest. 2: 188.

No undoubted herbarium specimens belonging to the original material have been found so far. The only currently known original material is Plate W in "Reise..." (Pallas, 1773b), which we are designating here as the lectotype of the name. Knjasev (2001: 141) wrote concerning the type of this name: "Возможно, типом О. spi-cata следует считать рисунок Pallas (1776: tabl. 4) [Probably, the type of О. spicata should be considered a drawing in Pallas ([corr.: 1773, tabl. W])]". We, however, do not agree with Knjasev (2001) concerning the dates when Pallas found the plant and its exact locus classicus. Pallas travelled in the left-bank area of the Ui (Uy) River from June 12 to 13, 1770, and mentioned some rivers and streams of its basin in his itinerary, such as Uvelka, Bach Kamennaja (Kamenka River), Sanarka, Tieplaja Rietschka (Teplaya River), etc.1 (Pallas, 1773a: 104-113). On June 13, he reached Kukuschevskoi Rud-nik (literally "Kukusha's mine") located on the left bank of the Ui River (Pallas, 1773a: 114-116), east of the modern Tungatarovo village, Republic of Bashkortostan (Ankushev et al., 2016). The only legume plant mentioned on these pages of "Reise... " was referred to as "Hedysarum saxatile" with a touch of uncertainity: "Ich glaube in der Benennung dieser Pflanze nicht zu irren [I believe I am not wrong in naming of this plant]" (Pallas, 1773a: 107). It is very likely that this plant, initially confused by Pallas with the West European-Mediterranean Onobrychis saxatilis (L.) Lam. (= Hedysarum saxatile L.), was later described as Astragalus spicatus (Pallas, 1773b). In favour of this, is a comparison of a brief outline of the plant (Pallas, 1773a: 107) with the protologue of A. spicatus (Pallas, 1773b: 742). In particular, both texts mention woolly radical leaves: "auf einer holzigten Wurzel viele Blatter... etwas wolligt" (Pallas, 1773a) and "folia tomentosa, radicalia" (Pallas, 1773b); leafless scapes: "den Blumenstengel ohne Blatter" (Pallas, 1773a) and "scapi... aphylli" (Pallas, 1773b); spike-like raceme resembling that of Onobrychis: "die Blumenohre von Onobrychis" (Pallas, 1773a) and "spica Onobrychidis simili forma et colore" (Pallas, 1773b). Also, its habitat, namely exposed rocky slopes, is described with partly similar words: "in dem freyen Gebürge" (Pallas, 1773a) and "in acclivibus saxosis australioribus et apricis" (Pallas, 1773b), respectively. Thus, we believe that the most probable locus classicus of A. spicatus is the area between the Kamenka River and the upper reaches of the Ui River near Tungatarovo village, where Pallas was 1213th June 1770.

Pallas had not seen plants in fruit: "Legumina perfecta non vidi" (Pallas, 1773b: 742). The characters of

1 All the watercourses are located in the modern Chelyabinsk Region; the original spelling follows Pallas (1773a).

fruits, however, are essential for the systematics of the genus Oxytropis, namely, the legume size, degree of its pubescence, shape of trichomes, and especially features of the septum dividing the fruit into two locules. For this reason, we are designating a specimen with ripened legumes as the epitype of the name (Turland et al., 2018: Art. 9.9), namely, the plant collected on the Iren-dyk Range (southeastern Bashkortostan), in the zone of feathergrass-sheep's fescue steppes, about 200 km from both Tungatarovo and the Kamenka River. This plant has the characters typical of the species, namely, semi-rosulate stems; leaves up to 20-22 cm long with petioles 12-13 cm long; 12-14(21) pairs of sessile, ovate-lanceolate, acuminate leaflets; robust sulcate pilose peduncles (the description of Astragalus caudatus in "Species Astragalorum" (Pallas, 1800-1803: 62): "Scapus... nu-dus", is obviously erroneous, disagreeing with the image on Plate L in the same work, where the pubescence of short patent hairs is depicted) almost three times longer than leaves, 30-40 cm long; inflorescence a dense many-flowered raceme (30-60 flowers!); legumes 1214 mm long, 3-4 mm broad, pubescent with white patent hairs.

L e c t o t y p e (Sytin, Byalt, Sokolova, designated here): Icon: Pallas, 1773, Reise Russ. Reich. 2, 2: tab. W. — Fig. 1: A.

Epitype (Sytin, designated here): Europ. Russia, Bashkiria, "Plantae Baschkiricae. Зилаирский кантон. Ковыльно-разнотр[авная] степь пологих склонов увала к СЗ от д. Аныковой [Zilairsky canton (district), feathergrass-forb steppe of gentle slopes of an elongated hill NW of Anykova village], 1 VIII 1929 [fr.], Крашенинников, Афанасьев [Krasheninnikov, Afa-nasyev], № 1156" (LE: LE 01205148!). — Fig. 5.

Protologue quotation: "Astragalus spicatus. Tab. W... Crescit in acclivibus saxosis australioribus et apricis ad orientales limites montium Uralensium, praesertim circa Ujum fluvium, florens Junio".

Phaca sylvatica Pall. 1776, Reise Russ. Reich. 3, 2: 744, tab. W, fig. 2.

= Astragalus sylvicolus Pall. 1802, Sp. Astragal. XII: 95, tab. LXXVIII, pro nom. nov., suprefl., illeg.

= Oxytropis sylvatica (Pall.) Willd. 1-10 IX 1802, Sp. Pl. 3, 2: 1300; DC. 15 IX 1802, Astragalogia: 82, isonymum.

The specimen BM000958893, though partly destroyed during storage, in some features resembles the image on Plate W in the Pallas' "Reise..." volume 3 (Pallas, 1776), in particular, in a split caudex, and the presence of a young inflorescence with undeveloped axis and unopened buds. There is a possibility that this herbarium specimen was the plant from which Plate W

Fig. 5. Epitype of Astragalus spicatus (LE 01205148). A — habit; B — fruits; C — leaf.

LE 01205148

(Pallas, 1776) was drawn despite the absence of any annotation by Pallas. On the other hand, the specimen stored at M (M-0185384) has an original Pallas' label "Astragaloides sylvatica Pall. it. III", and the earlier epithet "sylvatica" (rather than later "sylvicola") suggests that this specimen can also belong to the original material. In addition, the long bracts depicted on Plate W (Pallas, 1776) are more clearly visible on the Munich specimen, as well as some verticillate leaflets. Thus, it is not unlikely that the image was "collective", sharing the features of both plants (Fig. 2). Here we designate the specimen M-0185384 as the lectotype.

The specimens determined as Oxytropis sylvatica stored in the type collection at LE and listed by Kres-tovskaya and Byalt (1998) are not original Pallas' specimens as they were gathered by later collectors.

Original specimen: E Sib., "[Lambert:] Herb. Pallas [alab., fl.]" (BM000958893 — image!).

Protologue quotation: "Phaca? sylvatica Tab. W fig. 2... In pinetis Davuriae cum Trifolio hedysaroide fre-quens occurrit".

Acknowledgements

The work was carried out within the framework of the State assignment of the Komarov Botanical Institute "Vascular plants of Eurasia: systematics, flora, plant resources" (project № ААААА19-119031290052-1). The help of the internet community "Deutsche Palaog-raphie" with deciphering Merck's and Pallas' handwritten notes is highly appreciated. The authors are most grateful to anonymous reviewers for valuable comments, and to Dr. Keith Chamberlain (UK) for linguistic correction of the text.

Литература | References

Alvin: Platform for digital collections and digitized cultural heritage. 2023 / Uppsala University Library; University Library, Lund University; Gothenburg University Library etc. https://www.info.alvin-portal.org/ (Accessed 16.01.2023).

Ankushev M. N., Yuminov A. M., Zaikov V. V., Kotlyarov V. A., Blinov I. A. 2016. Starinnyye mednyye rudniki Nikolskogo rudnogo polya (Yuzhnyi Ural) [Ancient copper mines of Nikolskoye ore field (South Ural)] // Metallogeny of ancient and modern oceans-2016. From mineral genesis to deposits. Miass: IMin UB RAS. P. 96-101. [In Russian] (Анкушев М. Н., Юминов А. М., Зайков В. В., Котля-ров В. А., Блинов И. А. 2016. Старинные медные рудники Никольского рудного поля (Южный Урал) //

Металлогения древних и современных океанов-2016. От минералогенеза к месторождениям. Миасс: И Мин УрО РАН. С. 96-101).

Belyaeva I. V., Sennikov A. N. 2008. Typification of Pallas' names in Salix // Kew Bull. Vol. 63, № 2. P. 277-287. https://doi.org/10.1007/s12225-008-9015-0

Borodin I. 1908. Kollektory i kollektsii po flore Sibiri [Collectors and collections on the flora of Siberia]. St. Petersburg: Tipogr. Imp. Akad. nauk. iv + 247 p. [In Russian] (Бородин И. П. 1908. Коллекторы и коллекции по флоре Сибири. СПб.: Тип. Имп. Акад. наук. iv + 247 с.).

Byalt V. V., Sytin A. K. 2022. Lectotypification of some names, published by P. S. Pallas, in Astragalus L. and Phaca L. (Fabaceae) // Skvortsovia. Vol. 8, № 2. P. 25-57. https://doi.org/10.51776/2309-6500_2022_8_2_25

Candolle A.-P. de. 1802. Astragalogia nempe Astragali, Biser-rulatae et Oxytropis, nec non Phacae, Colutae et Lessertiae, historia iconibus illustrata. [Ed. quarto.] Parisiis: J. B. Car-nery. [viii]+270 p., 50 ills. https: //doi.org/10.5962/bhl.title.517

Catalogue of the type specimens of the vascular plants from Siberia and the Russian Far East kept in the Herbarium of the Komarov Botanical Institute (LE). Part 1. 2012 / ed.-in-chief I. V. Sokolova. Moscow; St. Petersburg: KMK Sci. Press. 443 p. [In Russian] (Каталог типовых образцов сосудистых растений Сибири и российского Дальнего Востока, хранящихся в Гербарии Ботанического института им. В. Л. Комарова РАН (LE). Ч. 1. 2012 / отв. ред. И. В. Соколова. М.; СПб.: Т-во науч. изд. КМК. 443 с.).

Die Große Nordische Expedition: Georg Wilhelm Steller (1709-1746) — ein Lutheraner erforscht Sibirien und Alaska. 1996 / W. Hintzsche, T. Nickhol et al. Halle: Franckesche Stiftungen; Gotha: Perthes. 347 S.

Digital library (repository). 2020 / Tomsk State University. [In Russian] (Электронная библиотека (репозито-рий) Томского государственного университета. 2020). https://vital.lib.tsu.ru/vital/access/manager/Index (Accessed 29.03.2023).

Fedtschenko O., Fedtschenko B. 1909. Conspectus Florae Turkestaniae. Pt. 2 // Izv. Turkestansk. Otd. Imp. Russk. Geogr. Obshch. Suppl. to vol. 6. Yuryev: Typogr. K. Mattisen. P. 97-300. [In Russian] (Федченко О. А., Фед-ченко Б. А. 1909. Перечень растений дико растущих в Русском Туркестане. Ч. 2 // Изв. Туркестанск. отд. Имп. Русск. Геогр. общ. Прилож. к т. 6. Юрьев: Типогр. К. Маттисена. С. 97-300).

Georgi J. G. 1802. Nachträge für dessen geographisch-physikalische und naturhistorische Beschreibung des Russischen Reichs. Königsberg: F. Nicolovius. x+[11]-444 S.

Global Plants on JSTOR. 2023. https://plants.jstor.org/ (Accessed 18.05.2023).

Harms O. Ya. 2013. Gemmy travel of ober-huttenverwalter P. I. Shangin (to the 227 anniversary of his famous expedition of 1786). Barnaul. 309 p. (Trudy Tigireksk. Zapov. Vol. 6). [In Russian] (Гармс О. Я. 2013. Самоцветное путешествие обер-гиттенфервальтера П. И. Шангина (к 227-летию его знаменитой экспедиции 1786 года). Барнаул. 309 с. (Тр. Тигирекского заповедника. Вып. 6).

JACQ consortium. 2004-. Virtual Herbaria Website.

https://www.jacq.org/ (Accessed 23.06.2023). Knjasev M. S. 2001. Systematic and chorologic notes on the species of the genus Oxytropis (Fabaceae) in the Urals. IV. Species related to Oxyropis songorica // Bot. Zhurn. Vol. 86, № 4. P. 140-148. [In Russian] (Князев М. С. 2001. Заметки по систематике и хорологии видов рода Oxytropis (Fabaceae) на Урале. IV. Виды родства Oxytropis songorica // Бот. журн. Т. 86, № 4. С. 140-148). Krestovskaya T., Byalt V. 1998. The Siberian and Far East section of the Komarov Botanical Institute Herbarium: Guide to the microform collection. Leiden: IDC Publishers. 49 p. Lambert A. B. 1811. Some account of the herbarium of Professor Pallas // Trans. Linn. Soc. London. Vol. 10, pt. 2. P. 256-265. https://doi.org/10.1111/j.1096-3642.1810.tb00016.x Lipschitz S., Vassilczenko I. 1968. Herbarium centrale URSS (Herbarii Instituti Botanici nomine V. L. Komarovii Academiae Scientiarum URSS historia ab initiis ad tempora nostra. Leningrad: Nauka. 141 p. [In Russian] (Липшиц С. Ю., Васильченко И. Т. 1968. Центральный гербарий СССР. Исторический очерк. Л.: Наука. 141 с.). Miller H. S. 1970. The herbarium of Aylmer Bourke Lambert. Notes on its acquisition, dispersal, and present whereabouts // Taxon. Vol. 19, № 4. P. 489-553. https://doi.org/10.2307/1218947 Pallas P. S. 1771-1776. Reise durch verschiedene Provinzen des Rußischen Reichs. Theilen 1; 2 (Büchen 1, 2); 3 (Bü-chen 1, 2). St. Petersburg: Kaiserl. Acad. Wiss. 1773a. Th. 2, Buch 1: vom Jahr 1770. [6]+368 S.; 1 Karte. 1773b. Th. 2, Buch 2: vom Jahr 1771; Anhang. S. [i-ii]+[369]-744; 32 Taf.; 1 Karte.

1776. Th. 3, Buch 2: ...im 1773sten Jahr. S. [455]-760+[32]; 45 Taf.; 2 Karten. Pallas P. S. 1800-1803. Species Astragalorum descriptae et iconibus coloratis illustratae, cum appendice. Lipsiae: G. Martini. 124 p., 99 tabs. [I-XCI]. https://doi.org/10.5962/bhl.title.46625 1800 [publ. 1800-1801]. Fasc. 1-2. [2] + viii+ 16 p.; tab. I-XIV.

1800 [publ. 1800-1801]. Fasc. 3-4. P. 17-32; tab. XV-XXVI.

1800 [publ. ?]. Fasc. 5-6. P. 33-52; tab. XXVII-XLI.

1801 [publ. 1802]. Fasc. 7-9. P. 53-76; tab. XLII-LXII.

1801 [publ. 1802]. Fasc. 10-11. P. 77-92; tab. LXIII-LXXVI.

1802 [publ. 1803]. Fasc. 12-13. P. 93-124; tab. LXXVII-XCI.

Pallas P. S. 1803[-1806]. Illustrationes plantarum imperfecte vel nondum cognitarum, cum centuria iconum. Lipsiae: G. Martini. iii + 68 p.; 59 plates by C. G. H. Geissler. Pallas P. S. 1811 [-1814]. Zoographia Rosso-Asiatica, sistens omnium animalium in extenso Imperio Rossico et adjacen-tibus maribus observatorum recensionem, domicilia, mores et descriptiones, anatomen atque icones plurimorum. Pet-ropoli: Offic. Caes. Acad. Sci. T. I-III.

Perret P. 1989. Précisions sur la publication du "Species Astragalorum" de P. S. Pallas. In: Burdet H. M. (éd.). Med-Checklist Notulae Bibliographicae, 14 à 17 // Candollea. T. 44, № 2. P. 571-573. http://doi.org/10.5169/seals-879629 Persoon C. H. 1807. Synopsis plantarum, seu enchiridium bo-tanicum... Ps. 2. Parisiis Lutetiorum: Treuttel et Würtz; Tubingae: J. G. Cottam. P. 273-657. https: //doi.org/10.5962/bhl. title. 638 POWO. 2023. Plants of the World Online / Royal Botanic Gardens, Kew. http://www.plantsoftheworldonline.org/ (Accessed 20.06.2023). Stafleu F. A., Cowan R. S. Taxonomic literature: A selective guide to botanical publications and collections with dates, commentaries and types. 2nd ed. https: //doi.org/10.5962/bhl.title.48631 Vol. 1: A-G. 1976. Utrecht: Bohn, Scheltema & Holkema. Junk. xl + 1136 p. (Regnum Veg. Vol. 94). Vol. 4: P-Sak. 1983. Utrecht; Antwerpen: Bohn, Scheltema & Holkema; The Hague; Boston: W. Junk. x + 1214 p. (Regnum Veg. Vol. 110).

Vol. 7: W-Z. 1988. Utrecht; Antwerpen: Bohn, Scheltema & Holkema; The Hague; Boston: W. Junk. lvi + 653 p. (Regnum Veg. Vol. 116). Sytin A. K. 1994. An unpublished work by P. S. Pallas "Icones plantarum selectarum cum descriptionibus" or "Plantae selectae rossicae" // Bot. Zhurn. Vol. 79, № 7. P. 92-98. [In Russian with English abstract] (Сытин А. К. 1994. Неопубликованное сочинение П. С. Палласа "Icones plantarum selectarum cum descriptionibus" или "Plantae selectae rossicae" // Бот. журн. Т. 79, № 7. С. 92-98). Sytin A. K. 2014. Peter Simon Pallas as a botanist. Moscow: KMK Sci. Press. 456 p. [In Russian] (Сытин А. К. 2014. Ботаник Петр Симон Паллас. М.: Т-во науч. изд. КМК. 456 с.).

Thiers B. M. 2023- (updated continuously). Index Herbario-rum: A global directory of public herbaria and associated staff. New York Botanical Garden. https://sweetgum.nybg. org/science/ih/ (Accessed 31.03.2023). Turland N. J., Wiersema J. H., Barrie F. R., Greuter W., Hawk-sworth D. L., Herendeen P. S., Knapp S., Kusber W.-H., Li D.-Z., Marhold K., May T. W., McNeill J., Monro A. M., Prado J., Price M. J., Smith G. F. (eds.). 2018. International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Glashütten: Koeltz Botanical Books. 254 p. (Regnum Veg. Vol. 159). https://doi.org/10.12705/Code.2018 Vascular Plants Herbarium of the Komarov Botanical Institute RAS — Herbarium LE. 2023. (Гербарий высших растений Ботанического института им. В. Л. Комарова РАН (БИН РАН). 2023). https://en.herbariumle.ru/ (Accessed 21.06.2023). Willdenow C. L. 1802. Species Plantarum. T. 3, ps. 2. Berolini. P. 851-1474. https://doi.org/10.5962/bhl.title.37657

Typification of names published by P. S. Pallas in Astragalus and Phaca, now in genus Oxytropis sect. Orobia (Fabaceae) Текст научной статьи по специальности «Биологические науки»

Аннотация научной статьи по биологическим наукам, автор научной работы — Sytin A. K., Byalt V. V., Sokolova I. V.

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Типификация названий астрагалов (Astragalus, Phaca), описанных П. С. Палласом, в настоящее время относящихся к секции Orobia рода Oxytropis (Fabaceae)

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