FloraOfNewZealand-SeedPlants-6-Heenan-2020-Cardamine
Cardamine in New Zealand has a long and complex taxonomic history, which is outlined below, based on a more detailed account provided by Heenan (2017). The first Cardamine species to be recognised in New Zealand was Sisymbrium heterophyllum G.Forst. (Forster 1786), and this was transferred to Cardamine by Schulz (1903). However, the combination Cardamine heterophylla (G.Forst.) O.E.Schulz is preoccupied in Cardamine, and so the new name C. forsteri Govaerts was provided (Govaerts 1999). Candolle (1821) provided the name C. debilis DC. and although this was not widely used in the years after it was published (e.g., by Hooker 1844, 1864), it was adopted extensively in New Zealand for nearly 60 years (e.g., Allan 1961; Webb et al. 1988). Cardamine debilis DC. is an illegitimate name because it includes Sisymbrium heterophyllum in synonymy.
The name Cardamine hirsuta L. was applied to New Zealand plants by Hooker (1844, 1853, 1864) and this was followed by Kirk (1899) and Cheeseman (1906). Hooker (1844) named the new variety C. hirsuta var. subcarnosa Hook.f. and provided the new species names C. corymbosa Hook.f., C. depressa Hook.f. and C. stellata Hook.f. Only a few years later Hooker (1853) provided a treatment accepting only two species, and he did not accept any of the species he had named in 1844. He accepted C. hirsuta and included Sisymbrium heterophyllum and C. debilis DC. in synonymy. The second species Hooker (1853) accepted was the newly named C. divaricata Hook.f., but this was later treated as a synonym of C. stylosa DC. (Candolle 1821). This species is now placed in Rorippa as R. divaricata (Hook.f.) Garn.-Jones & Jonsell (Garnock-Jones & Jonsell 1988). For further discussion on the Australian and New Zealand Cardamine names now placed in Rorippa, see Garnock-Jones (1978) and Garnock-Jones & Jonsell (1988).
Hooker (1864) provided a treatment of Cardamine that was quite different from his 1844 and 1853 classifications. He continued to recognise C. hirsuta var. subcarnosa, but added three more varieties: C. hirsuta var. debilis (DC.) Hook.f. and C. hirsuta var. corymbosa (Hook.f.) Hook.f., and the new variety C. hirsuta var. uniflora Hook.f. He also reduced C. stellata to the rank of variety, providing the new combination C. depressa var. stellata (Hook.f.) Hook.f., and he accepted C. stylosa DC. and the newly named and described C. fastigiata Hook.f. Cardamine stylosa is treated by Garnock-Jones (1978) as a synonym of Rorippa gigantea (Hook.f.) Garn.-Jones (Garnock-Jones 1978), and C. fastigiata is treated as Pachycladon fastigiatum (Hook.f.) Heenan & A.D.Mitch. (Heenan et al. 2002).
In his treatment of Cardamine, Kirk (1899) followed Hooker (1864) in the recognition of C. hirsuta (five varieties), C. depressa (two varieties), C. stylosa, and C. fastigiata. Kirk (1899) also included C. latesiliqua Cheeseman (Cheeseman 1883) and C. enysii Cheeseman ex Kirk (Kirk 1899); both species are now treated in Pachycladon (Heenan et al. 2002). Kirk (1899) described the new species C. bilobata Kirk. Kirk's treatment of C. hirsuta is notable in that he recognised the autonym C. hirsuta var. hirsuta. This is the first recognition of this taxon being present in New Zealand and being distinguished from the indigenous varieties of C. hirsuta. In his treatment of C. hirsuta, Kirk (1899) emphasised variation in stamen number, stating for C. hirsuta var. hirsuta "stamens usually 4", thereby distinguishing this naturalised taxon from the indigenous varieties with six stamens. Cheeseman's (1906) treatment of Cardamine was based on that of Kirk (1899), with the only difference being that under C. hirsuta he did not include C. hirsuta var. hirsuta. The reason for this is quite straightforward, with Kirk (1899) including naturalised species, whereas Cheeseman (1906) comprised only indigenous species.
Cheeseman (1925) provided some significant changes to his 1906 treatment of Cardamine, introducing some of the taxonomy proposed in a worldwide revision of the genus by Schulz (1903). Two changes were that Schulz (1903) and Cheeseman (1925) both excluded C. hirsuta from the indigenous flora of New Zealand, and they also applied C. heterophylla (G.Forst.) O.E.Schulz. The treatment of C. heterophylla provided by Schulz (1903) included five infraspecific taxa at various ranks, but these were not mentioned by Cheeseman (1925), because he did not consider Schulz’s treatment able to be applied to the diversity of Cardamine in New Zealand. The application of the taxonomy proposed by Schulz is still problematic today, because the type specimens of Schulz’s five C. heterophylla varieties are missing, presumed to have been destroyed when the Berlin Herbarium was damaged by fire in a bombing raid during March 1943 (Heenan 2017). The Schulz (1903) names were published at various infraspecific ranks, so they would not have any priority over the new species names published by Heenan (2017).
Schulz (1903) also reinstated Cardamine corymbosa at species rank, and accepted C. stellata, C. depressa var. depressa and C. depressa var. acaulis. Cheeseman (1925) accepted C. corymbosa, but differed from Schulz (1903) in accepting C. depressa var. depressa and C. depressa var. stellata, as well as C. bilobata which wasn’t mentioned by Schulz (1903). Cheeseman (1925) accepted C. heterophylla var. uniflora (Hook.f.) Cockayne (Cockayne 1909), and Schulz treated C. hirsuta var. uniflora as a synonym of C. heterophylla. Cheeseman (1925) followed Schulz (1903) in treating C. subcarnosa as C. glacialis var. subcarnosa (Hook.f.) O.E.Schulz. Three species treated by earlier authors in Cardamine (C. enysii, C. fastigiata, and C. latesiliqua) were considered by Cheeseman (1925) to be species of Nasturtium Brown in Aiton (1812), having been formally transferred to that genus by Cheeseman (1911). These three species are now placed in Pachycladon (Heenan et al. 2002).
Pritchard (1957) studied morphological variation of Cardamine in New Zealand and recognised several unnamed entities that provided the basis for subsequent studies. Pritchard (1957) recognised three unnamed forest races in C. debilis ('glossy leaf', 'long style' and 'narrow petal'), C. corymbosa and an affiliated 'mainland coastal race', and two alpine races ('tussock race' and 'scree race'). The two most recent Flora of New Zealand treatments for Cardamine accepted six (Allan 1961) and five (Webb et al. 1988) species, with both treatments accepting C. bilobata, C. corymbosa, C. debilis, C. depressa, and C. subcarnosa. Allan (1961) also accepted C. uniflora (Hook.f.) Allan, but this was treated as a synonym of C. corymbosa by Webb et al. (1988); Allan (1961) also accepted C. depressa var. stellata but this was reduced to a synonym of C. depressa by Webb et al. (1988). Both of these treatments reinstated at species rank C. subcarnosa and recognised two distinct populations of this species. The population of C. subcarnosa from Auckland Islands was named as C. latior Heenan (Heenan 2008), with C. subcarnosa restricted to Campbell Island.
Phylogenetic analyses by Mitchell & Heenan (2000) and artificial hybrids (Heenan 2002) were used as the basis to transfer Iti lacustris Garn.-Jones & Johnson (Garnock-Jones & Johnson 1987) to Cardamine as C. lacustris (Garn.-Jones & P.N.Johnson) Heenan (Heenan 2002). The diminutive and much reduced C. cubita Molloy, Heenan & Smissen (Heenan et al. 2013) has been named and described as a new species. Most recently, a monograph of Cardamine in New Zealand was published, and this accepts 41 indigenous taxa, an additional four species are accepted as naturalised, and several putative unnamed taxa are discussed (Heenan 2017). One of these putative unnamed taxa was named as C. panatohea by Heenan & de Lange (2018).
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