Pinaceae in the Herbarium of the Institute of Botany at the Jagiellonian University, Kraków, Poland (KRA)

The Herbarium of the Jagiellonian University in Kraków, Poland (KRA) has extensive collections. The Pinaceae family in KRA embraces 1,057 herbarium sheets and contains representatives of eight out of 11 genera usually distinguished in the family. The collection of the family in KRA contains ca. 54–61% of the 220–250 species occurring in the world. The most numerous species (116 sheets) is Pinus sylvestris . There is one isoneotypus of Larix decidua Mill var. carpatica Domin (KRA 224704) and one syntypus of Tsuga caroliniana Engelm. (KRA 224989) in the collection. There are 706 sheets from Europe, 504 of them come from areas covered by the contemporary borders of Poland, 206 from North America, 98 from Asia, two from Africa, and one from Australia. The herbal material of the family deposited in KRA was collected in the past 200 years. The oldest specimen was collected in 1821. There are 65 sheets which date from the nineteenth century, 56 from the years 1900 to 1918, 173 from 1919 to 1939, 532 from 1944 to 2000, and 139 sheets from the twenty-first century. The most interesting collections include: the exsiccata from the nineteenth century, sheets from China (1925–1926), sheets collected by various Russian expeditions to Siberia, the collection of Professors Jan Kornaś and Anna Medwecka-Kornaś from North America, and collections documenting the scientific activity of the “Kraków geobotanical school” in the twentieth century.


Introduction
Herbaria have long histories and, being collections of dried plants with information attached to labels, they are interesting objects of research for plant systematicians, phytogeographers, and historians of botany. Not only entire herbaria, but also individual collections and even taxonomic units included in a herbarium, are the subject of research in Poland (e.g., [1][2][3][4][5][6][7]) and in the world (e.g., [8]). Herbarium specimens are also an important source of DNA for plant research (e.g., [9]). Herbarium labels, as well as the plant material, contain a lot of valuable information that can be analyzed for many years after collecting a specimen. Based on this information, one can comprise biographies of botanists (e.g., [10]), or analyze contacts between botanists (e.g., [11,12]).
The Herbarium of the Institute of Botany at the Jagiellonian University (KRA), founded in 1780, has the longest history among Polish university herbaria, dating back to the second half of the eighteenth century. It also holds extensive collections [13][14][15]. In 2016-2017, an inventory of herbarium materials belonging to the Pinaceae family located in KRA was made. During the inventory, the quantity and quality of the deposited plant material was recorded, and photos of interesting specimens, labels, and entire sheets were taken. Information from labels served to create a database. The purpose of this study is to present the Pinaceae collections in KRA.

General characteristics
The inventory showed that there are 1,057 sheets with specimens belonging to Pinaceae in KRA; they are included in 32 fascicles. Almost all specimens (995) are determined to species. A slightly lower number of specimens contains information such as locality (95%), collection date (90%), or the name of the collector (89%). Half of the labels feature information about the habitat (51%), and even fewer about the height above sea level of the locality (16%). Some labels (ca. 4%) do not provide any information except for species name and possibly the name of the collector. Almost half of the sheets (615) also contain envelopes with loose plant material.

Taxonomic characteristics
During the inventory, it was found that almost all of the herbarium material (99%) was determined to species. Only nine sheets contained specimens determined to genus (six genera), and only six were undetermined. The previous determination of specimens was not verified. Nevertheless, all species names appearing on labels were checked in the Internet database The Plant List [16]. On this basis, it can be concluded that Pinaceae in KRA embrace ca. 123 species (Tab. S1). This number may of course change in the future, after verifying the determination of the entire herbarium material. Pinaceae collections in KRA contain, depending on the adopted taxonomic approach, from 54% to 62% of all the 220-250 species distinguished within this family worldwide. In the KRA collection, there are representatives of eight out of the 11 genera usually distinguished (e.g., [17] [18].

Geographical characteristics and the origin of specimens
Species of the Pinaceae family occur mainly in cold and temperate regions of the Northern Hemisphere.
In the described collection, 707 sheets were from Europe, of which 503 came from areas covered by the contemporary borders of Poland, and 204 from other European countries (Tab. 1).
Most of the sheets from Poland come from its southern part, mainly the Tatra, Beskidy, and Bieszczady mountains, as well as the Kraków-Częstochowa Upland, the Włoszczowska Basin, and Opoczyńskie Hills. These specimens were collected, for the most part, by naturalists affiliated with the Jagiellonian University: employees of the Institute of Botany, assistants of the Department of Botany, or PhD students. 1 Sources of biographical data of collectors -see Tab. S2.
Tab. 1 Numbers of sheets collected from European countries (other than Poland).

Country
Number of sheets Ukraine 37 Together 204

Historical characteristics
The herbal material of the Pinaceae in KRA was collected during the last 200 years by at least 284 collectors (listed in Tab. S2). However, dates of collecting do not fall evenly throughout this period. The oldest sheets come from the first half of the nineteenth century, but most Pinaceae come from the second half of the twentieth century, when the so-called "Kraków geobotanical school" was conducting its full scientific activity [21] (Fig. 1). Information on the most interesting specimens from an historical point of view is given below.
There are several sheets from the first half of the nineteenth century. The oldest specimen (Pinus pinaster KRA 224553) was collected, as its handwritten label informs ( Fig. 2A), in 1821 in Fréjus, a city in southern France, by Jacques Étienne Gay (1786-1864), a Swiss-French botanist, civil servant, collector, and taxonomist. There is no information about how long the specimen has been in KRA. The next two specimens, from a chronological point of view, were part of a collection purchased for KRA in 1851-1852 in Germany [13]. The collection was prepared by Unio Itineraria, a nineteenth-century German joint stock company [22]. A specimen of Pinus halepensis KRA 224546 was gathered in St. Pietro (Sardinia, Italy) in 1827 by Franz August Müller (1798-1871), pharmacist, plant collector, and cryptogam researcher. A specimen of Pinus pinaster KRA 224552 was collected in November 1830 near Bayonne (a town in the Western Pyrenees in France) by Philip Anton Christoph Endress (1806-1831), a German pharmacist and plant collector from Strasburg, during his second (out of three) journey to the Pyrenees. Both specimens have original printed labels (Fig. 2B,C). Pinus teocote KRA 224781 (collected in Chinautla, State of Puebla, Mexico) was part of the exsiccatae "Plantae mexicanae" (Fig. 2D), prepared in Mexico from 1841 to 1843 by the Danish plant collector Frederik Michael Liebmann (1813-1856) and distributed by the botanical museum in Copenhagen from 1845. Pinus leiophylla KRA 224549 was part of the exsiccatae "Plante centroamericanae" prepared in 1845-1848 by the Danish biologist Anders Sandøe Ørsted (1816-1872), and also distributed by the botanical museum in Copenhagen. Both specimens mentioned above were probably purchased for KRA in 1851-1852 [13]. The next two sheets (Pinus sylvestris KRA 224788 and Pinus sylvestris rubra KRA 224787) originate in 1843, probably from the Nikitsky Botanical Garden near Yalta in the Crimea (Russia), in existence since 1812. There is no information about how long the specimens have been in KRA.
Tab. 2 Numbers of sheets collected in North America.  In 1876, Władysław Kulczyński (1854-1919), then a student of the Jagiellonian University, and later zoologist-arachnologist, handed over a set of Galician herbarium consisting of 435 sheets collected in the previous year [13]. Three sheets come from this collection: two of them were collected in the Tatras (Pinus cembra KRA 224666 and Pinus montana KRA 224556), and a specimen of Pinus sylvestris KRA 224799 was collected in Bielany near Kraków.

Country and state Number of sheets
Aleksander Jan Śleńdziński (1848-1881) was a botanist-florist. Already in the last year of his studies, he was an assistant at the chair of botany and in the Botanical Garden of the Jagiellonian University under Professor Ignacy Rafał Czerwiakowski (1808-1882). While still at university, he began working with the Physiographical Commission of the Academy of Sciences and Letters, which financed his floristic studies of Eastern Galicia from 1873 [23]. In KRA, there are 18 of his sheets (including one without a given locality) collected from 1872 to 1879: six from Kraków and its surroundings (Abies excelsa KRA 224867, Abies pectinata KRA 223674, Larix decidua KRA 224471, 224470, and 224699, Pinus sylvestris KRA 224798), and 12 from the Eastern Galicia (Abies pectinata KRA 223673, Pinus cembra KRA 224541, Pinus silvestris KRA 224794, etc.) (Fig. 2F). In 1877, he handed his herbarium consisting of about 1,000 sheets over to KRA [13]. Perhaps in later years he also made some donations or maybe after his death some of his herbaria was transferred to KRA, since among these 18 sheets one was collected in 1879, after his donation.
In 1886, Ignacy Szyszyłowicz (1857-1910), while working as an assistant-volunteer at the Imperial Nature Museum in Vienna (K. k. Naturhistorisches Hofmuseum), traveled to Montenegro and Albania. Two sheets (Abies alba KRA 223672 and Pinus leucodermis KRA 224551) collected by this naturalist in 1886 in Montenegro are preserved in KRA (Fig. 2G), but there is no information about when they were handed over to KRA. From 1875, a sheet of Larix decidua KRA 224469 comes from Rakowice near Kraków (currently near the city center). Unfortunately, only the initials of the collector appear on the original label. Perhaps the specimen was collected by the then student of the Jagiellonian University and later assistant to Czerwiakowski, Józef Krupa (1850-1889), who donated his collection of west Galician plants to KRA in 1876 [13].
There are six sheets of plants from the Kamchatka Peninsula (Russia). They were collected in 1879-1882 by Benedykt Dybowski (1833-1930), a Polish zoologist and limnologist, and outstanding researcher of Lake Baikal. Later, Carl Maximovich (1827-1891), a famous Russian botanist who specialized in Asian plants, among others, determined such sheets. There is no evidence regarding their arrival to KRA. Perhaps Marian Raciborski (1863-1917) obtained them from Dybowski when he and Dybowski were working at Lwów University (as the university was known until World War II, currently Lviv University in Ukraine) at the same time, and he brought them with him when moving to the Jagiellonian University in 1912.
Beginning in 1852, the Jagiellonian University was able to allocate income from its plant sales from the University Botanical Garden to enlarge its herbarium collections. Thanks to this possibility, Czerwiakowski purchased various European exsiccatae [13]. Given below is information about the oldest and most interesting exsiccatae preserved in collections in KRA.
"   prepared the first edition of "Flora polonica exsiccata", and the specimen was collected for the exsiccata [24].
A large group of sheets result from the activities of Marian Raciborski and his students who prepared another Polish exsiccatae, "Rośliny polskie" [Polish plants] [25]. In KRA, there are also specimens collected by Polish botanists during their foreign trips before World War I. Władysław Szafer (1886-1970), later a professor of botany at the Jagiellonian University and one of the leading Polish botanists of his time, studied at the Faculty of Philosophy of the University of Vienna, under Richard Wettstein, among others. In 1908, Szafer participated in a several-week-long biological course at the sea station in Trieste, and also went on a floristic trip to nearby Dalmatia. During the trip he collected Pinus halepensis (KRA 224673) in Porto Medolino near Pula in the Istria Peninsula (Fig. 2K). In 1913, Józef Antoni Żmuda (1889-1916), assistant to M. Raciborski at the Botanical Garden and Botanical Institute of the Jagiellonian University, spent some time at the same sea station. In KRA, there are three sheets collected by him at that time, e.g., Pinus halepensis (KRA 224674), and transferred to KRA in 1914 [14]. Based on their label data, one can ascertain that they were part of a documentation of the Istrian flora.
Among the other sheets from the period, many doublets were obtained from other herbaria. Most of these (22) come from the Herbarium of the Institute of Botany, Academy of Sciences of the USSR. They were collected mainly from 1909-1914 during various Russian expeditions carried out in different areas of Siberia, e.g., Larix sibirica (KRA 27589) (Fig. 2L), Larix czekanowskii (KRA 28369), or Picea schrenkiana (KRA 224967). They were purchased in 1933 [14].
A total of 173 sheets were collected in the interwar period. Undoubtedly, the most interesting are specimens (26 sheets) from China. They were gathered by Joseph Francis Charles Rock (1884-1962) during an expedition organized by the

Final remarks
The collections of the Pinaceae family in KRA, their species composition, and geographical origin reflect the changing purposes that the Herbarium of the Jagiellonian University fulfilled over the past 200 years. From the time when it served mainly as a didactic aid, i.e., around the World War I, specimens originating from outside present-day Poland prevail. They were to document mainly the taxonomic diversity within Pinaceae. Sheets of the oldest exsiccatae come from that period as well. Later, the task of documenting Polish flora and research came to the forefront. These studies were conducted (and still are, as evidenced by sheets from the twenty-first century) mainly by members of the so-called "Kraków geobotanical school". Therefore, most of the sheets from this period contain specimens from Poland, and the collectors mostly work at the Institute of Botany of the Jagiellonian University. The enlargement of KRA collections with collections from outside Poland, such as China, Russia, or North America, has not been neglected either, which serve as comparative material.
The taxonomic rank of a herbarium is confirmed by the number of nomenclature types in its possession. Unfortunately, there are only two types of Pinaceae in KRA (for comparison, Kew has at least 178). This difference results from both the history of the KRA and the entire specific character of Polish science. During the period when the most active botanical exploration on Earth was conducted and huge funds were allocated to it, Poland was either not on the map or had no money for expensive research expeditions, seen in its failure in organizing an expedition to Brazil and Peru in 1928 [26].
Thanks to the historical analysis of the Pinaceae collection in KRA, it is possible to supplement unknown biographies of some botanists with details and dates. An example may be the sheets collected by Śleńdziński in Kraków and its surroundings in 1872-1875, which is proof of the previously unknown activity of this botanist in his place of residence (it is a pity that he did not publish anything about it). Raciborski's sheets testify to his research in 1911 in the Rodnei Mountains (in the Inner Eastern Carpathians, now in Romania), and in the Pieniny Mts, while the sheets of Szafer indicate that he was in Switzerland in 1923, etc.
Analysis of data obtained from the labels of the Pinaceae collection in KRA provided previously unknown knowledge about the set itself, as well as its geographical, taxonomical, and historical structure. Furthermore, it contributed to clarifying the biographies of some botanists and disclosed mutual contacts between KRA and other botanical institutions.