Lachnagrostis batesii in Europe as Agrostis distans , 160 years before being described for South Australia (Poaceae)

Mid-19th Century collections of the endemic South Australian Lachnagrostis batesii A.J.Br. were found in Vienna (W), Paris (P) and Saint Petersburg (LE), labelled as ‘ Agrostis distans Kze’. The species was described in the 1847 Leipzig index seminum and listed on several European indexes from 1851–1867. Although Kunze’s description was brief and the type destroyed in WWII, his taxon is very likely the same as that later described as L. batesii . Consequently, the new combination Lachnagrostis distans (Kunze) A.J.Br. is made. It is proposed that A. distans raised in Europe was originally derived from seed separated from a collection made by Hans Behr in the 1844–1845 summer and described by Dietrich von Schlechtendal as Deyeuxia aemula Kunth.


Introduction
consultation of relevant historical documents and searches for further specimens at other European herbaria. Herbarium codes per Thiers (2022).

Results and Discussion
Examination and interpretation of the labels of the Agrostis distans sheets reveal that they derived from cultivation of Australian seed in various European botanical gardens from 1847 to 1869 (Table 1).
Annual index seminum (i.e. published index or catalogue to seed collections) have been produced by botanic gardens since the mid-16th Century, and continue to be a common output (Aplin et al. 2007). During the 19th Century, new plants and seed collected from the New World were highly sought after in Europe, and propagation of such was a major activity for many institutions. The index seminum served as an opportunity for horticulturists to exchange plant material and increase the range of species on display for their visitors. Previously unknown taxa were commonly described in these indexes and serve as  (Otto 2018).
Behr returned to South Australia in 1848-1849 and made further plant collections, of which some are thought to have been made with Mueller. The repository for the collections from this trip is uncertain, as Behr didn't return to Germany but travelled onto the Philippines and then settled in California (Kraehenbuehl 1981, Bretag 2016. It is probable that he left his collections with Mueller, destined for Joseph Hooker at Kew Gardens, but which may instead have been sent to Sonder. Of the 19 Behr collections imaged in the Kew Herbarium Catalogue (2020), all but two (ex Mueller) are from Sonder's herbarium.
The evidence therefore indicates that Behr collected Lachnagrostis batesii instead of L. aemula or L. filiformis in his 1844-1845 trip to South Australia and that either field harvested seed or disarticulated seed (i.e. florets and caryopsis) from the plant collection was passed from Behr or Schlechtendal to Richter in a batch of Australian material for the Luisium Gardens. However, it was Kunze who recognised that the plants raised from seed at Leipzig ex Richter did not conform to Deyeuxia aemula Kunth. It may be coincidence, but the only year between 1838 and 1849 that Kunze offered D. aemula seed in the Leipzig index seminum was in the same year in which he described A. distans, suggesting that he obtained seed of D. aemula with the express purpose of comparing its resulting plants with A. distans. If the A. distans seed Kunze received from Richter was harvested in the northern summer of 1846 (i.e. Jun-Aug), then Richter must have received and sown the seed very soon after Behr returned to Germany in May 1846.
The origin of Lachnagrostis filiformis and/or L. aemula seed in Europe is unknown. Bentham (1878) noted that Australian seed (of unknown source) was used to raise plants in Continental Gardens, which Willdenow (1809) described as Agrostis retrofracta, Poir (1810) as Agrostis debilis Poir and Kunth (1829) as Deyeuxia retrofracta. Koch (1831) notes that he had been growing A. aemula in the greenhouse at Erlangen. A review of online surviving index seminum (Lut 2017) shows that seed of A. retrofracta or D. retrofracta was sporadically offered for sale or exchange from as early as 1818 through to 1868, from gardens including, Breslau, Halle, Dorpat, Hamburg, Schwetzingen, Gottingen, Turin, Pisa, Berlin, Madrid, St. Petersburg, Vienna and Uppsala, and A. aemula or D. aemula seed was offered from 1824 to 1871 from Vienna, Budapest, Erlangen, Breslau, Hamburg, Gottingen, St Petersburg, Dresden, Heidelberg, Pisa, Basel, Angers, and as noted above, from Leipzig. While the seed of A. retrofracta and D. retrofracta was likely to be of L. filiformis, seed of A. aemula and D. aemula could have been either L. filiformis or L. aemula or not even consistent among botanical gardens. This is not surprising given the confusion concerning the morphological distinctions between these taxa and   (Lut 2017). Perhaps Richter did receive seed of D. billardieri from Behr's collection, but it failed to establish or survive the cold central European winter. Although all these Lachnagrostis taxa are temperate species, L. billardierei, being predominately a coastal grass, may be less tolerant of cold altitudes.
There are no specimens of A. distans lodged at Naturkundemuseum Leipzig (Karl Heyde 2019, pers. comm., 17 th Jan.) or at MNVD in Dessau-Roßlau, nor any documents relating to correspondence between Richter and Kunze in the archives of the former principality and state of Anhalt (Timm Karisch, pers. comm., 17 Apr. 2019).
The earliest collection of Agrostis distans that has been found, is at P ( Table 1). The collection was made in 1849 by Jean Charles Marie Grenier (1808-1875), a Professor at the Faculty of Sciences at Besançon University and Director of the Jardin botanique de Besançon (Botanical Garden of Besançon) and received from Grenier's herbarium in the year of his death. It consists of two small specimens to about 15 cm tall with just emerging inflorescences and cultivated from seed received from nearby Dijon, probably from the Jardin botanique de l'Arquebuse (Arquebuse Botanical Garden). Although the sheet provides no further information, presumably Dijon received seed from Leipzig to grow plants during the 1848 season. If this is the case, the Grenier collection represents second generation offspring of the parent plants described by Kunze and fourth generation offspring from Behr's original collection.
The index seminum for Hortus Botanicus Imperialis Petropolitanus first offered seed of Agrostis distans in 1853 and then in 1856-1858, 1860-1861 and 1864-65. Specimen sheets at LE ( Although it is not certain, the sheets at LE probably represent collections made of cultivated plants at Saint Petersburg, rather than collections made elsewhere and sent from donating gardens. If so, those from Berlin and Leipzig were collected in March 1852, while those from Karlsruhe were collected in 1865. A further collection (Fig. 2), probably derived from F6 or F7 generation plants of the pre-1852 seed, was made in 1857 by the Bavarian, Ferdinand von Herder (1828-1896, who worked at the Saint Petersburg Botanical Gardens from 1856-1891. All these collections coincide with seed offerings through the index seminum. While LE 01072060 and LE 01072061 have single labels relating to one seed lot, the other sheet labels present a confused picture (Table 1). The label for LE 0107259 refers to both Reichenbach's and Braun's seed, despite the single sheaf of relatively large and robust specimens which were obviously made from a single gathering. The collection is undated, and the seed sources may have been mixed prior sowing. While LE 01072060 depicts relatively small and stunted plants derived from Braun's seed lot, LE 01072062 consists of 16 similar stunted specimens of uniform size, primarily labelled in pencil as Reichenbach's seed lot, but with Braun added in a separate but later and different hand in ink. In addition, LE 01072062 has a separate label referring to Herder's collection of 1865 based on the Karlsruhe seed lot. The implication from this mixed labelling is that the specimens were not mounted until a later era and matching all the labels to their correct respective collections was not possible.

Note:
The neotype was chosen as the earliest of the collections found in Europe and therefore the closest cultivated generation to the specimen described by Kunze.