ABSTRACT
Blechnaceae is a medium-sized family of ferns in New Zealand with one genus, 23 indigenous species, two naturalised species and two nothospecies. A new combination, Blechnum × digenum (Parris) Brownsey & Perrie, is made for the hybrid between Blechnum molle (Parris) Christenh. and B. parrisiae Christenh. 11 lectotypes are chosen for basionyms relevant to New Zealand – Lomaria dura T. Moore, L. filiformis A. Cunn., L. membranacea Colenso ex Hook., L. nigra Colenso, L. oligoneuron Colenso, L. parvifolia Colenso, L. paucijuga Colenso, L. pumila Raoul, L. rotundifolia Colenso, Parablechnum procerum var. acuminatum C. Presl, and Stegania procera var. stipulosa A. Rich. No type material has been found for Blechnum capense var. auriculatum Domin, B. capense var. contractum Domin, B. capense var. hookerianum Domin, Lomaria paleacea Potts, or Lomaria pygmaea Colenso, and the types for these names remain undesignated. Lectotypes are being chosen where possible for all names at specific and subspecific rank, for which no holotype was designated by the original author, in order to fix the application of the name concerned. Justification for the choice is provided in each case. This article is a contribution towards clarifying the taxonomic and nomenclatural status of New Zealand plants for the plant names database (Ngā Tipu Aotearoa) and the electronic Flora of New Zealand.
Introduction
The fern family Blechnaceae is represented in New Zealand by one genus and 23 indigenous species – Blechnum blechnoides (Bory) Keyserl., B. chambersii Tindale, B. colensoi (Hook.f.) N.A.Wakef., B. deltoides (Colenso) T.C.Chambers, B. discolor (G.Forst.) Keyserl., B. durum (T.Moore) C.Chr., B. filiforme (A.Cunn.) Ettingsh., B. fluviatile (R.Br.) Lowe ex Salomon, B. fraseri (A.Cunn.) Luerss., B. kermadecense Perrie & Brownsey, B. membranaceum (Colenso ex Hook.) Mett. ex Diels, B. minus (R.Br.) Ettingsh., B. molle (Parris) Christenh., B. montanum T.C.Chambers & P.A.Farrant, B. neohollandicum Christenh., B. nigrum (Colenso) Mett., B. norfolkianum (Heward) C.Chr., B. novae-zelandiae T.C.Chambers & P.A.Farrant, B. parrisiae Christenh., B. penna-marina subsp. alpina (R.Br.) T.C.Chambers & P.A.Farrant, B. procerum (G.Forst.) Sw., B. triangularifolium T.C.Chambers & P.A.Farrant, and B. zeelandicum Christenh. In addition, there are two nothospecies, Blechnum × aggregatum (Colenso) Tindale and Blechnum × digenum (Parris) Brownsey & Perrie comb. nov., and two naturalised species, B. patersonii (R.Br.) Mett. and B. punctulatum Sw. The classification of Blechnaceae used here follows Perrie et al. (Citation2014) rather than PPG Citation1 (Citation2016), but the choice of classification for the electronic Flora of New Zealand treatment will be discussed there.
The synonymy for Blechnaceae includes 64 basionyms that are relevant to New Zealand. In order to fix the application of these names, their types need to be clarified. Lectotypes have previously been chosen for 17 of these (see ), and holotypes identified and documented for another 31 basionyms by Brownsey & Perrie (in prep.). Lectotypes for 11 of the remaining basionyms are selected here as part of a series on New Zealand’s indigenous ferns (Brownsey and Parris Citation2012; Brownsey and Perrie Citation2012, Citation2013, Citation2014a, 2014b, Citation2015a, 2015b, Citation2016a, 2016b, 2016c, Citation2017, Citation2019; Perrie and Brownsey Citation2015; Brownsey et al. Citation2018, Citation2019) clarifying the taxonomic and nomenclatural status of New Zealand plants for the plant names database (Ngā Tipu Aotearoa) (http://nzflora.landcareresearch.co.nz/default.aspx?NavControl=search&selected=NameSearch) and the electronic Flora of New Zealand (http://www.nzflora.info/). No type material has been found for the following five names – Blechnum capense var. auriculatum Domin, B. capense var. contractum Domin, B. capense var. hookerianum Domin, Lomaria paleacea Potts, or Lomaria pygmaea Colenso.
Table 1. Basionyms of New Zealand Blechnaceae for which holotypes have previously been identified or lectotypes designated.
Materials and methods
Herbarium abbreviations follow Thiers (Citation2019). Type material was sought in herbaria known to house the main collections of relevant authors – AK, K and WELT for William Colenso, BM for Allan Cunningham, K for Thomas Moore, P for E.F.L. Raoul and Achille Richard, PR for Karel Domin, PRC for C.B. Presl, and W for Ignaz von Szylszlowicz. Images of some of the types can be viewed through the relevant institutional websites or JSTOR Global Plants (http://plants.jstor.org). Syntypes for each species comprise only single morphological entities unless otherwise stated.
New combination
Blechnum × digenum (Parris) Brownsey & Perrie, comb. nov. ≡ Doodia × digena Parris, New Zealand J. Bot. 10: 596 (1972). Holotype: Kitekite Falls, Piha, Auckland, B.S. Parris, 26.5.1966, CHR 202054 (isotype: BM).
Notes: Doodia × digena was described by Parris (Citation1972) as the hybrid between Doodia caudata (Cav.) R.Br. and D. media subsp. australis Parris. Under the classification of Blechnaceae proposed by Perrie et al. (Citation2014) and adopted here, Doodia was subsumed into an expanded Blechnum. For most New Zealand species of Doodia new combinations have already been made in Blechnum, but one is still required for the nothospecies Doodia × digena, now treated as the hybrid between B. molle (Parris) Christenh. and B. parrisiae Christenh.
Lectotypifications
Lomaria dura T.Moore, Gard. Chron. 290 (1866) ≡ Blechnum durum (T.Moore) C.Chr. Index Filic. 153 (1905)
Lectotype (designated here): Chatham Island, Hort. Standish, 1864, K! (photo WELT E 467/23)
Notes: Lomaria dura was described by Moore (Citation1866) who stated ‘This very distinct fern was introduced from the Chatham Islands some few years since by Mr Watson, from whom it passed into the hands of Mr Veitch and Mr Standish, to both of whom we are indebted for specimens’. Four syntype collections are mounted on one sheet in Thomas Moore’s fern herbarium at K, and all are candidates for lectotype. There are three sterile fronds labelled ‘Hort. Veitch 1859, ex Chatham Island’, ‘Hort. Veitch 1860, ex Watson from Chatham Island 1858’, and ‘Hort. Veitch 1861, ex Chatham Island’. There is also a collection labelled ‘Hort. Standish 1864, ex Chatham Island’ comprising one fertile and one sterile frond. Moore’s protologue specifically refers to the sterile fronds being longer than the fertile and, therefore the Standish collection, which is the only one to include a fertile frond, is designated here as the lectotype.
Lomaria filiformis A. Cunn, Companion Bot. Mag. 2: 363 (1837) ≡ Blechnum filiforme (A.Cunn.) Ettingsh., Denkschr. Kaiserl. Akad. Wiss., Math. Naturwiss. Kl. 23: 57, t. 6, f. 5 (1864)
Lectotype (designated here): New Zealand, Cunningham, 1826, BM 001048229!
Notes: Lomaria filiformis was described by Cunningham (Citation1837) from material collected by him at Whangaroa in 1826. Allan (Citation1961) stated that the type was at K, but that appears to have been an error because no relevant sheets have been located there. However, there are three syntype collections at BM. Two of them (BM 001048227–8) are labelled ‘New Zealand, A. Cunningham’ without date or further locality. The third is similarly labelled, but with the date ‘1826’ (BM 001048229). The latter sheet is designated here as the lectotype because of the additional collection information.
Lomaria membranacea Colenso ex Hook., Sp. Fil. 3: 34, t. 145 (1859) ≡ Blechnum membranaceum (Colenso ex Hook.) Mett. ex Diels in Engler & Prantl, Nat. Pflanzenfam. 1(4): 249 (1899)
Lectotype (designated here): New Zealand, Bay of Islands, J.D. Hooker, Herb. Hooker., K! (photo WELT E467/11)
Notes: Lomaria membrancea was described by W.J. Hooker (Citation1859) but attributed to Colenso, based on material collected by Colenso and J.D. Hooker in the Bay of Islands, and by Joliffe from Waiheke Island. Three potential syntype collections are in the Hookers’ Herbarium at K. One was collected by J.D. Hooker in the Bay of Islands; a second is labelled ‘Waiheki [Waiheke] Island, near Auckland, Joliffe, 1852’; and the third (K 001092718) is labelled ‘2017, Lomaria membranacea, n. sp., W.C.’ in Colenso’s hand-writing. Although the latter specimen has been annotated ‘holotype’ by Chambers, it is clear that Colenso No. 2017 was sent to W.J. Hooker in September 1848 (St. George Citation2009, p. 240) and was collected at Te Hawera (now Hamua, between Masterton and Woodville), not in the Bay of Islands. It cannot therefore be considered part of the original material seen by W.J. Hooker. Of the remaining two syntypes, the J.D. Hooker collection is more extensive, comprising two whole fertile plants. It is designated here as the lectotype.
Lomaria nigra Colenso, Tasmanian J. Nat. Sci. 1: 375 (1843) ≡ Blechnum nigrum (Colenso) Mett., Ann. Sci. Nat. Bot. Sér. IV, 15: 69 (1861)
Lectotype (designated here): near Tauranga, Jan. 1842, Herb. Colenso, WELT P003243!
Notes: Lomaria nigra was described by Colenso (Citation1843) from plants he collected in ‘humid places in the dense forest between Tauranga and Rotorua’ in January 1842. There are syntype collections in K and WELT. The specimen at K is in the Hookers’ Herbarium and is labelled ‘291, Lomaria nigra, n. sp. W.C.’ in Colenso’s hand-writing. Colenso No. 291 was sent to W.J. Hooker in September 1842 (St. George Citation2009, p. 166) and was collected on 13 January 1842 ‘from the stony banks of Mangarewa, a small river running in a deep ravine’. The locality is near to Rotorua and is consistent with the protologue. The second specimen in WELT is labelled ‘“Lomaria nigra, n. sp. W. Colenso, humid spots, dense forest, near Tauranga, Jan. 1842’, and is also consistent with the protologue. Both sheets bear two fertile plants and are good candidates for lectotype. However, the collection details on the WELT specimen are somewhat closer to those in the protologue, and it is designated here as the lectotype.
Lomaria oligoneuron Colenso, Trans. & Proc. New Zealand Inst. 16: 346 (1884) = Blechnum membranaceum (Colenso ex Hook.) Mett. ex Diels in Engler & Prantl, Nat. Pflanzenfam. 1(4): 249 (1899)
Lectotype (designated here): Great Barrier Island, C.P. Winkelmann, Herb. Colenso, WELT P003335!
Notes: Lomaria oligoneuron was described by Colenso (Citation1884) from specimens collected on Great Barrier Island by C.P. Winkelmann in 1883. There are syntype collections in K and WELT. The sheet at K (001092719) is labelled ‘Lomaria oligoneuron (sp. nov.)’ in Colenso’s hand-writing and ‘New Zealand, W. Colenso, rec’d 12/1885’ in a different hand. It bears several fertile and sterile fronds, and parts of rhizomes. The sheet in WELT is labelled ‘Gt Barrier, Winkelmann’ in Colenso’s hand-writing, and ‘Great Barrier Island, type of L. oligoneuron Col.’ in Cheeseman’s writing. It has four complete plants with fertile and sterile fronds. The sheet at WELT has more complete collection data linking it to the protologue, and is designated here as the lectotype.
Lomaria parvifolia Colenso, Trans & Proc. New Zealand Inst. 20: 224 (1888) = Blechnum penna-marina subsp. alpina (R.Br.) T.C.Chambers & P.A.Farrant, Fern. Gaz. 15: 96 (1996)
Lectotype (designated here): Tongariro, H. Hill, com. W. Colenso 5/1890, K 001092741!
Notes: Lomaria parvifolia was described by Colenso (Citation1888) from specimens collected by Henry Hill in 1887 from high slopes of Tongariro Mountain Range, County of East Taupo. There are two syntype collections in K and AK. The sheet in K (001092741) bears two sterile fronds and is labelled ‘Lomaria parvifolia Col. from Tongariro, per Mr Hill, Trans. N.Z. Inst., vol. XX’ in Colenso’s hand-writing, and ‘com. W. Colenso 5/1890’ in a different hand. The sheet in AK (143480) also bears two sterile fronds and is labelled ‘type of L. parvifolia Col., Tongariro, H. Hill’ in Cheeseman’s hand-writing. Both sheets are very similar, but the sheet in K has the label data in Colenso’s hand-writing and is designated here as the lectotype.
Lomaria paucijuga Colenso, Trans. & Proc. New Zealand Inst. 20: 222 (1888) ≡ Blechnum deltoides (Colenso) T.C.Chambers, Telopea 22: 48 (2019)
Lectotype (designated here): Tongariro, H. Hill, Herb. Colenso, WELT P003332!
Notes: Lomaria paucijuga was described by Colenso (Citation1888) from specimens collected by Messrs Owen and Hill in 1887 from the sides of Mt Tongariro, County of East Taupo. There are two syntype collections in K and WELT. The sheet in K (001092696) bears a plant consisting of four juvenile fronds and a piece of rhizome. It is labelled ‘Lomaria paucijuga Col., Trans. N.Z. Inst., vol. XX’ in Colenso’s hand-writing and ‘com. W. Colenso, 5/1890’ in a different hand. The sheet at WELT (P003332) bears two small, partly eaten fronds, and is labelled ‘Lomaria paucijuga Col.’ in Colenso’s hand-writing, and ‘type of L. paucijuga Col., Tongariro, H. Hill’ in Cheeseman’s writing. The specimen at K is more complete, but that at WELT has more complete collection data. Chambers and Wilson (Citation2019) cited the WELT specimen as holotype, and the K specimen as isotype, but Colenso (Citation1888) did not indicate a holotype or the herbarium where material was deposited. A lectotype therefore needs to be designated, and the WELT specimen is chosen here, in line with its citation as holotype by Chambers and Wilson (Citation2019).
Lomaria pumila Raoul, Ann. Sci. Nat. Bot. Sér. III, 2: 115 (1844), nom. illeg., non Lomaria pumila Kaulf. (1824) = Blechnum penna-marina subsp. alpina (R.Br.) T.C.Chambers & P.A.Farrant, Fern. Gaz. 15: 96 (1996)
Lectotype (designated here): Nouvelle-Zélande, Presqu’ȋle de Banks [New Zealand, Banks Peninsula], Raoul, 1843, P 01618104 (image!)
Notes: Lomaria pumila was described by Raoul (Citation1844) from Akaroa but is illegitimate because of an earlier homonym. There are three syntype collections in P (1618102–1618104), all labelled ‘Nouvelle-Zélande – Presqu’ȋle de Banks’ [New Zealand – Banks Peninsula], two of which are additionally labelled ‘Ann. Sc. Nat., Aout 1844’ – a reference to the place of publication of the new name in Annales des Sciences Naturelles (Raoul Citation1844). There is a fourth collection in the Hooker’s herbarium at K labelled ‘Akaroa, Raoul, Herb. Mus. Paris’, which is clearly a duplicate sent to Kew. Raoul’s main herbarium is in P, so the best choice for lectotype would be one of the two specimens at P which also indicate the place of publication. Both sheets bear a good range of sterile and fertile fronds, but P 01618104 is designated here as the lectotype because it also includes four whole plants with portions of the rhizome
Lomaria rotundifolia Colenso, Tasmanian J. Nat. Sci. 1: 377 (1843) nom. illeg., non Lomaria rotundifolia Blume (1828) = Blechnum fluviatile (R.Br.) Lowe ex Salomon, Nomencl. Gefasskrypt. 115 (1883)
Lectotype (designated here): woods near Poverty Bay, W. Colenso, Dec. 1841, WELT P003245!
Notes: Lomaria rotundifolia was described by Colenso (Citation1843) from specimens he collected in ‘dense woods near Waikare Lake, in the mountainous district in the interior of the North Island; five days journey from Poverty Bay, on the east coast, December, 1841’. There are two potential syntype collections at WELT and a third in K. The sheet in the Hooker’s Herbarium at K (001092725) bears one whole plant and two detached fertile and sterile fronds. It is labelled ‘265, Lomaria rotundifolia, n. sp., W.C., large specimen from forest on mountains’ in Colenso’s hand-writing. Colenso No. 265 was sent to W.J. Hooker in September 1842 (St. George Citation2009, p. 161) and was collected in December 1841 from the vicinity of Lake Waikaremoana. One sheet in WELT bears one fertile and one sterile frond, and is labelled ‘L. rotundifolia var., woods, mountains, near Waikare Lake’ in Colenso’s hand-writing (WELT P003244). The other, bearing two complete plants and two detached sterile fronds, is also labelled in Colenso’s writing, ‘Lomaria rotundifolia, n. sp., W. Colenso, woods near Poverty Bay, December 1841’ (WELT P003245). WELT P003244 is labelled by Colenso as a variety of L. rotundifolia and is therefore not a good choice for lectotype. Neither of the other two specimens are particularly well localised, although both are consistent with the protologue and are correctly dated. WELT P003245 is designated here as the lectotype because it comprises more complete material.
Parablechnum procerum var. acuminatum C.Presl, Epimel. Bot. 109 (1851) = Blechnum novae-zelandiae T.C.Chambers & P.A.Farrant, New Zealand J. Bot. 36: 14 (1998)
Lectotype (designated here): Tab. 177–178 in Icones Plantarum, Vol. 5 (Hooker Citation1841)
Notes: Parablechnum procerum var. acuminatum was described by Presl (Citation1851). He included Lomaria procera var. ß Hook., Icon. Pl. 5: t. 177–178 (1841) in synonymy, but cited no other specimen or locality that specifically represented his new variety, although he included a long list of material for P. procerum itself. No material has been identified in Presl’s herbarium at PRC. Similarly, Hooker’s description of Lomaria procera var. ß included no cited specimens or localities. However, it did include a very clear illustration, which is undoubtedly referable to Blechnum novae-zelandiae. In the absence of other original material, the Plate 177–178 in Icones Plantarum (Hooker Citation1841) is designated here as the lectotype.
Stegania procera var. stipulosa A.Rich., Voy. Astrolabe, Essai 86, t. 13 (1832) = Blechnum novae-zelandiae T.C.Chambers & P.A.Farrant, New Zealand J. Bot. 36: 14 (1998)
Lectotype (designated here): Fertile frond in Tab. 13 in Voyage de découvertes de l’Astrolabe, Botanique (Richard Citation1832).
Notes: Stegania procera var. stipulosa was described by Richard (Citation1832) from material collected at the Bay of Islands during the Voyage of the Astrolabe. Both the sterile and fertile fronds are characterised in the description, but in the observations that follow, it is the stipule-like expansions at the base of the fertile pinnae that are emphasised as distinguishing var. stipulosa from Stegania procera sens. strict. The only potentially relevant specimen found at P comprises a single fertile frond which shows reduced sterile basal pinnae and stipule-like expansions at the base of the fertile pinnae. It belongs to the species currently recognised as Blechnum novae-zelandiae. However, the original label states only ‘Stegania procera Br., Astrolab. bot. t., Nlle Zelande’. The reference to ‘Astrolab. bot. t.’ could be taken to mean that this was the specimen illustrated with the original description in Voyage de découvertes de l’Astrolabe, Botanique (Richard Citation1832). On the other hand, there is nothing on the label that indicates that it is Richard’s var. stipulosa, or that the specimen was collected from the Bay of Islands prior to 1832. Therefore there is too much uncertainty about the specimen to designate it as the lectotype. However, accompanying the original description is an illustration (Richard Citation1832, tab. 13) of both a sterile and fertile frond of what is labelled Stegania procera. The protologue (Richard Citation1832, p. 86) states that ‘nous avons figuré ici cette belle variété’, making it clear that the illustration represents Stegania procera var. stipulosa rather than the type variety. Unfortunately, it depicts a mixed collection – the sterile frond belonging to what is now recognised as Blechnum discolor and the fertile frond to Blechnum novae-zelandiae. This is reflected in the description which accurately characterises sterile and fertile fronds from these two different species. Nevertheless, the emphasis in the protologue on the stipule-like expansions at the base of the fertile pinnae is enough to determine that it was the characters of the fertile frond which induced Richard to nominate his new variety. In the absence of any convincing original specimen, the illustration of the fertile frond in Richard’s tab. 13 in Voyage de découvertes de l’Astrolabe, Botanique is here designated as the lectotype.
Types not designated
No type material has been located for the following basionyms, and their identity remains unresolved.
Blechnum capense var. auriculatum Domin, Biblioth. Bot. 20(85): 115 (1913)
Notes: No type material of Blechnum capense var. auriculatum has been located at PR, the main repository of Domin’s Australasian types, and its identity remains unresolved.
Blechnum capense var. contractum Domin, Biblioth. Bot. 20(85): 115 (1913)
Notes: No type material of Blechnum capense var. contractum has been located at PR, the main repository of Domin’s Australasian types, and its identity remains unresolved.
Blechnum capense var. hookerianum Domin, Biblioth. Bot. 20(85): 115 (1913)
Notes: No type material of Blechnum capense var. hookerianum has been located at PR, the main repository of Domin’s Australasian types, and its identity remains unresolved.
Lomaria paleacea Potts, New Zealand Country J. 6: 84 (1882)
Notes: Lomaria paleacea was described by Potts (Citation1882) from Banks Peninsula. No type material has been located at AK, CHR, K or WELT and its identity remains unresolved. The dimensions of the frond and pinnae given in the protologue suggest that the name relates to a species in the Blechnum procerum complex, but it is impossible to be certain which one.
Lomaria pygmaea Colenso, Trans. & Proc. New Zealand Inst. 25: 322 (1893) = Blechnum membranaceum (Colenso ex Hook.) Mett. ex Diels in Engler & Prantl, Nat. Pflanzenfam. 1(4): 249 (1899)
Notes: Lomaria pygmaea was described by Colenso (Citation1893) from a specimen collected by Henry Hill in 1892 from the area between Dannevirke and the East Coast. No type material has been located in AK, K or WELT, the usual repositories of Colenso collections. Its synonymy with B. membranaceum is based on historical precedent first determined by Cheeseman (Citation1906), which is at least partly consistent with the protologue.
Acknowledgements
We are grateful to staff at AK, CHR, K, W, and WELT for access to collections in their care, to Germinal Rouhan at P and Ota Sida at PR for endeavouring to locate type specimens in their herbaria, and to an anonymous referee who corrected some errors in the original text.
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References
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