European Journal of Taxonomy 375: 1–52
https://doi.org/10.5852/ejt.2017.375
ISSN 2118-9773
www.europeanjournaloftaxonomy.eu
2017 · Holstein N. & Weigend M.
This work is licensed under a Creative Commons Attribution 3.0 License.
Monograph
No taxon left behind? – a critical taxonomic checklist of
Carpinus and Ostrya (Coryloideae, Betulaceae)
Norbert HOLSTEIN 1,* & Maximilian WEIGEND 2
1,2
Rheinische Friedrich-Wilhelms-Universität Bonn, Bonn, Nordrhein-Westfalen, Germany.
*
Corresponding author: holstein@uni-bonn.de
2
Email: mweigend@uni-bonn.de
Abstract. Hornbeams (Carpinus) and hop-hornbeams (Ostrya) are trees or large shrubs from the northern
hemisphere. Currently, 43 species of Carpinus (58 taxa including subdivisions) and 8 species of Ostrya
(9 taxa including sudivisions) are recognized. These are based on 175 (plus 16 Latin basionyms of
cultivars) and 21 legitimate basionyms, respectively. We present an updated checklist with publication
details and type information for all accepted names and the vast majority of synonyms of Carpinus
and Ostrya, including the designation of 54 lectotypes and two neotypes. Cultivars are listed if validly
described under the rules of the ICN. Furthermore, we consider Carpinus hwai Hu & W.C.Cheng to be a
synonym of Carpinus fargesiana var. ovalifolia (H.J.P.Winkl.) Holstein & Weigend comb. nov. During
the course of our work, we found 30 legitimate basionyms of non-cultivars that have been consistently
overlooked since their original descriptions, when compared with the latest checklists and loristic
treatments. As regional loras are highly important for taxonomic practice, we investigated the number
of overlooked names and found that 78 basionyms were omitted at least once in the eight regional
treatments surveyed. More seriously, we found 4 basionyms of accepted species being overlooked in a
major loristic treatment.
Keywords. Herbarium digitization, taxon inventory, taxonomy, typiication.
Holstein N. & Weigend M. 2017. No taxon left behind? – a critical taxonomic checklist of Carpinus and Ostrya
(Coryloideae, Betulaceae). European Journal of Taxonomy 375: 1‒52. https://doi.org/10.5852/ejt.2017.375
Introduction
In the ongoing discussions on the loss of biodiversity and taxonomy crisis, inventories of taxon names
are a crucial step. Floristic treatments and annotated checklists for countries, provinces or national parks
are widely used as tools for biodiversity assessments, monitoring, and conservation (Palmer et al. 1995;
Funk 2006). Regional projects are very important, but regional taxonomic treatments may be of limited
use when wide-spread taxa are concerned. The narrow focus of regional treatments carries the risk of
overemphasizing local variability and repeatedly describing more wide-spread taxa (Scotland & Wortley
2003; Michelangeli 2005). Therefore, taxonomically comprehensive lists as a baseline for taxonomic
work are required. This need is addressed by, e.g., the series the World Checklist of Selected Plant
Families (WCSP 2016) or The PlantList (http://www.theplantlist.org), with names and types available
from nomenclatural databases such as IPNI (http://www.ipni.org) or Tropicos (http://www.tropicos.org).
1
European Journal of Taxonomy 375: 1–52 (2017)
The correct and valid application of taxon names (at the rank of family and below), however, is determined
by designated taxonomic types (Art. 7.1. ICN; McNeill et al. 2012). Therefore, lists of names are only
the initial step, since it is the type specimen that anchors the identity of the taxon. Obtaining type
information, often deeply hidden in the historical literature, is a tedious process and the physical type
specimens may be diicult to locate, but a correct and unequivocal typiication is the crucial basis
for disentangling taxonomic problems (Gaston & Mound 1993). Only then, as a feedback mechanism,
treatments will be improved not only by data density, but also by data quality.
Almost 20 years ago, the World Checklist and Bibliography of Fagales (Govaerts & Frodin 1998)
provided a baseline for the taxa of this order containing many species that constitute the temperate
forests. However, critical taxonomic revisions are still scarce, with a revision of the genus Betula L. as
a noteworthy exception (Ashburner et al. 2013). Carpinus L. and Ostrya Scop. belong to Betulaceae
subfam. Coryloideae Hook.f. The species of the two genera are large shrubs or trees in forests nearly
across the northern hemisphere. A few species are used for specialty timber or for landscaping and
hedges, sometimes also as an ornamental, with Carpinus betulus L. and a range of cultivars of that
species being by far the most important taxon. Carpinus and Ostrya are sister to the Chinese Ostryopsis
Decne., a genus of three species (Holstein & Weigend 2016). While some studies ind Ostrya to be
paraphyletic to Carpinus (Grimm & Renner 2013), it is more often found nested in Carpinus (Yoo &
Wen 2002, 2007; Li 2008). Ostrya is easily diferentiated morphologically by having its nuts enclosed
in pale, papery, inlated tubular bracts with a small blunt distal opening. Nuts of Carpinus are more or
less exposed, at most half-covered by a side wing of the one large abaxial bract. An additional, minute,
caducous, adaxial scale bract can be found in some East Asian species (Carpinus sect. Distegocarpus
(Siebold & Zucc.) Sarg.) Currently, the World Checklist of Selected Plant Families (WCSP 2016) lists
42 accepted species of Carpinus and 8 of Ostrya, with both genera having a center of diversity in China.
The revision of Carpinus for China by Hu (1933) is the latest critical taxonomic treatment on a larger
geographical scale, while Furlow (1987) worked on the four North American taxa. The latest global key
for Carpinus and Ostrya (Rushforth 1987) is an extension of the key from Li’s Chinese loral treatment
(Li 1979a). Here, we amend the current lists of taxa of Carpinus and Ostrya with further names based on
extensive literature and in silico research, compare the taxonomic status of the names in various loras,
and present typiication data as far as available. Furthermore, we check the number of names not treated
in the various treatments despite occurrences in the covered area to analyze the rate of overlooked taxa.
Material and methods
We assembled an initial list from the World Checklist of Selected Plant Families (WCSP 2016) and
complemented it with names from IPNI, The Plant List, Tropicos and other sources. We took the
acceptance status of World Checklist of Selected Plant Families as a baseline and compared the status
and the basionyms of the corresponding synonyms with the recent literature (Furlow 1987; Chang et al.
2014a), and the appropriate loras, such as the Flora of China (Li & Skvortsov 1999), Flora of Taiwan
(Liao 1996), Flora Sichuanica (Editorial Board of the Flora Sichuanica 2012), Flora of Japan (Ohwi
1965), Flora of North America (Furlow 1997), and Flora of the USSR (Bobrov 1936).
In order to calculate the number of overlooked basionyms, names relating to cultivars (hanging branches,
leaf color and maculation, and leaf serration mutants) were excluded. Illegitimate and invalid names
were listed in the Supplementary File as they are sometimes mentioned and may represent evolutionary
signiicant entities, but they were not included in the calculations either. A name was tagged as “missing”
for a speciic publication if it was not cited, although it should have been covered as it occurs in the focus
area of the publication.
Type information for the names was identiied from the protologues. The following herbaria were
searched for type specimens in situ: B, BM, BR, E, GOET, K, LIV, M, MANCH, MSB, W, and WU.
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HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
Additional specimens were obtained via personal communication (DWC, LE, TBI) or in silico: JSTOR
Plant Science (http://plants.jstor.org/), Chinese Virtual Herbarium (CDBI, IBK, IBSC, KUN, LBG, N,
NAS, PE, WUK; http://www.cvh.org.cn/), Hungaricana (BP; http://gallery.hungaricana.hu/en/), NHN
Naturalis (L, U, WAG), Virtuella Herbariet (LD, S; http://www.herbarium-ume.se/virtuella_herbariet),
and the homepages of the following herbaria: A, F, GH, HK, LE, LINN, MICH, MO, NY, P, PH, TAI,
TAIF, and US. Types of Korean taxa in Korean and Japanese herbaria are cited from Chang et al. (2014b).
Results
We found 175 legitimate basionyms for Carpinus (plus 16 for cultivars described under the rules of
the ICN, two of them not formally mentioned as cultivars) and 21 for Ostrya. The list of non-cultivar
basionyms with acceptance and taxonomic status, and the digital availability of at least one type specimen
per basionym is found in the Supplementary File. In total, 78 basionyms were overlooked in at least one
of the ten treatments where they should have been covered, plus the recently described C. langaoensis
(Lu et al. 2017). We found 32 legitimate basionyms not included in the World Checklist of Selected
Plant Families yet (WCSP 2016). The World Checklist of Fagales (Govaerts & Frodin 1998) missed
46 basionyms, plus three basionyms, C. chingiana, C. fargesiana var. tchouana and C. henryana var.
chuana (Yang 1997). The latter might, however, have been published between the copy deadline and the
publication of the checklist. Six relevant basionyms have been overlooked in the Flora of USSR (Bobrov
1936), ten in the Flora of Japan (Ohwi 1965), two in the Flora of Taiwan (Liao 1996), 34 in the Flora
Europaea (Tutin & Walters 1993), three in the Flora of North America (Furlow 1997), 14 in the Flora
of China (Li & Skvortsov 1999, including C. chingiana, C. fargesiana var. tchouana and C. henryana
var. chuana), one in the Flora Sichuanica (Editorial Board of the Flora Sichuanica 2012), and three in
the Checklist of Korea (Chang et al. 2014a). The four Chinese species C. lipoensis, C. luochengensis,
C. mengshanensis, and C. shimenensis are accepted in the World Checklist (Govaerts & Frodin 1998),
but not mentioned in the Flora of China (Li & Skvortsov 1999). The acceptance status of 21 names in
Carpinus and one name in Ostrya difers among the diferent treatments. Five basionyms in Carpinus
and one in Ostrya difer in the taxon to which the name is supposed to be synonymous only.
One hundred and two taxa of Carpinus and 16 of Ostrya (including all accepted ones) had original or
type material available online, most of them freely. However, almost all herbaria surveyed in situ for
type specimens and all herbaria with non-type digitization programs harbored hitherto unrecognized
type material to a considerable degree. Complete digitization of herbaria, such as in the Paris herbarium,
Naturalis or the China Virtual Herbarium, was most rewarding for this work, due to the manifold of
unrecognized type material and the time-unlimited access.
Checklist of Carpinus
Order Fagales Engl.
Family Betulaceae Gray
Subfamily Coryloideae Hook.f.
Tribe Carpineae A.DC.
Carpinus L.
Species Plantarum: 998 (Linnaeus 1753). ‒ Ostrya J.Hill, The British Herbal: 513 (Hill 1757) [“1756”]
nom. rejic. – C. sect. Eucarpinus Sarg., The Silva of North America 9: 40 (Sargent 1896) nom. inval.
[Art. 22.2, McNeill et al. (2012)]. – C. subg. Eucarpinus (Sarg.) Nakai, The Botanical Magazine
[Tokyo] 29: 37 (Nakai 1915) nom. inval. [Art. 22.2, McNeill et al. (2012)]. – C. subg. Carpinus (L.)
Hu, Acta Phytotaxonomica Sinica 9: 282 (Hu 1964). – C. ser. Betulae Hu, Acta Phytotaxonomica
Sinica 9: 282 (Hu 1964), nom. inval. [Art. 22.2, 22.6, McNeill et al. (2012)]. – C. subsect. Carpinus
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European Journal of Taxonomy 375: 1–52 (2017)
(L.) P.C.Li & S.H.Cheng, Flora Reipublicae Popularis Sinicae 21 (2): 65 (Li 1979a). – C. subsect.
Betulae (Hu) C.J.Wang, Flora Yunnanensis 5: 182 (1991) nom. inval. [Art. 22.2, 22.6, McNeill et al.
(2012)]. ‒ Type species: C. betulus L., lectotypiied by Britton & Brown (1913: 606).
Distegocarpus Siebold & Zucc., Abhandlungen der Mathematisch-Physikalischen Klasse der Königlich
Bayerischen Akademie der Wissenschaften 4 (3): 226 (Siebold & Zuccharini 1846). – Carpinus sect.
Distegocarpus (Siebold & Zucc.) Sarg., The Silva of North America 9: 40 (Sargent 1896). – C. subg.
Distegocarpus (Siebold & Zucc.) Nakai, The Botanical Magazine [Tokyo] 29: 37 (Nakai 1915). ‒
Type species: Distegocarpus carpinus Siebold & Zucc. (C. japonica Blume).
C. sect. Brachyspicae Nakai, The Botanical Magazine [Tokyo] 29: 37 (Nakai 1915). – Type species:
C. paxii H.J.P.Winkl.
C. sect. Elongatae Nakai, The Botanical Magazine [Tokyo] 29: 37 (Nakai 1915). – Type species: Not
designated.
C. ser. Monbeigianae Hu, Acta Phytotaxonomica Sinica 9: 285 (Hu 1964) [“Monberigianae”]. – C.
subsect. Monbeigianae (Hu) P.C.Li, Flora Reipublicae Popularis Sinicae 21 (2): 70 (Li 1979a). –
Type species: C. monbeigiana Hand.-Mazz.
C. ser. Polyneurae Hu, Acta Phytotaxonomica Sinica 9: 294 (Hu 1964). – C. subsect. Polyneurae (Hu)
P.C.Li, Flora Reipublicae Popularis Sinicae 21 (2): 84 (Li 1979a). – Type species: C. polyneura
Franch. [Art. 22.6, McNeill et al. (2012)].
C. ser. Pubescentes Hu, Acta Phytotaxonomica Sinica 9: 289 (Hu 1964). – Type species: C. pubescens
Burkill [Art. 22.6, McNeill et al. (2012)].
C. subg. Distegocarpus ser. Fangianae Hu, Acta Phytotaxonomica Sinica 9: 281 (Hu 1964). – Type
species: C. fangiana Hu.
C. subg. Distegocarpus ser. Cordatae Hu, Acta Phytotaxonomica Sinica 9: 282 (Hu 1964). – Type
species: C. cordata Blume.
1. Carpinus betulus L.
Species Plantarum: 998 (Linnaeus 1753). – Original citation: “Europe, Canada.” – Type: Sweden,
Småland, s. coll., s.n. (lecto-, designated by Browicz (1972: 3): LINN! ex herb. Linn. no. 1131.1). –
Additional original material: Sweden, Skâne, C. Linnaeus s.n. (S09–40045).
C. vulgaris Mill., The Gardeners Dictionary ed. 8, no. 1 (Miller 1768). – Original citation: “In Germania
copiose, rarius in Italia, in Scania & inferiore parte Smolandiae.” – Type: Dodoens (1616: 841)
[igure]. – Additional original material: Sweden, Småland, s. coll., s.n. (LINN! ex herb. Linn. no.
1131.1). S. loc., s. coll., s.n. (BM000647417! ex herb. Clif. no. 447).
C. sepium Lam., Flore Françoise 2: 212 (Lamarck 1779) nom. illegit. superl. – Note: Lamarck (1779)
cites C. betulus L. (Linnaeus 1763: 1416), which includes all citations for C. betulus L. (Linnaeus
1753).
C. compressus Gilib., Exercitia Phytologica 2: 399 (Gilibert 1792) nom. inval., opus utique oppr.
C. ulmifolia Salisb., Prodromus Stirpium in Horto ad Chapel Allerton Vigentium: 392 (Salisbury 1796)
nom. illegit. superl. – Note: Salisbury (1796) cites C. betulus L. (Linnaeus 1763: 1416), which
includes all citations for C. betulus L. (Linnaeus 1753).
C. ulmoides Gray, A Natural Arrangement of British Plants 2: 245 (Gray 1821) nom. illegit. superl. –
Note: Gray (1821) cites C. betulus L. (Linnaeus 1763: 1416), which includes all citations for C. betulus
L. (Linnaeus 1753).
C. austriaca M.Serres, Voyage dans le Tyrol: 320 (Serres 1823) nom. nud. – Original citation: “Les
environs de Brixen [...], je distinguai le carpinus austriaca, tout-à-fait étranger à la Flore du Nord du
Tyrol.” – Associated material: S.d. (P06810513!).
C. carpinizza Host, Flora Austriaca 2: 626 (Host 1831). – C. betulus [unranked] carpinizza (Host)
Dippel, Handbuch der Laubholzkunde 2: 140 (Dippel 1891) [“1892”]. – C. betulus var. carpinizza
(Host) C.K.Schneid., Illustriertes Handbuch der Laubholzkunde 1: 140 (Schneider 1904). –
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HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
C. betulus subsp. carpinizza (Host) O.Schwarz, Mitteilungen der Thüringischen Botanischen
Gesellschaft 1: 96 (Schwarz 1949). – Original citation: “In sylvis Transylvaniae. Kitaibel. Fl. Majo.
Incolae Transylvaniae, valachice loquentes, [...], et Carpinizza vocant.” – Type: not localized (not in
Kitaibel’s herbarium in BP or in W). – Note: the application of this name is somewhat ambiguous.
Host mentions in the description the 3-lobed bracts with smooth margins of the side lobes, typical
for C. betulus. Neilreich (1861: 76), however, mentions that the name “carpinizza” is the vernacular
Walachian name for C. duinensis Scop. (C. orientalis Mill.) and blames Kitaibel to whom Host refers
for mixing that up. Then again, there is a C. orientalis specimen in Kitaibel’s herbarium (XL/84!)
with a label that this plant is called “Kárpánicza”. It was therefore Host, not Kitaibel, misapplying
the name. Schneider (1904) refers the rank variety to Neilreich (1861: 76), but Neilreich did not
publish that combination. Therefore, Schneider’s combination is the irst at this rank.
C. betulus var. integriloba Spach, Annales des sciences naturelles, Botanique ser. 2 16: 252 (Spach
1841a). – Type: Ménagerie, 1833, s. coll., s.n. (lecto-, designated here: P06810518!). – Note: Spach
did not cite any locality or speciic specimen but this collection from Spach’s herbarium bears a note
“Nob.” (ours).
C. betulus var. odontoloba Spach, Annales des sciences naturelles, Botanique ser. 2 16: 252 (Spach
1841a). – Original citation: No detailed data given. – Type: not localized.
C. betulus f. serrata Schur nom. nud. in sched., non Beck (Beck von Mannagetta 1890). – Associated
material: Romania, Siebenbürgen, Bergregion bis 3000’, 1848, P.J.F. Schur 14164 (P06747296!).
Romania, Siebenbürgen, Hermannstadt [Sibiu], 1848, P.J.F. Schur 14166 (P06747370!). – Note:
this name is only known from two specimens, but Schur published many taxa, and this name might
have been published. Should Schur’s name have been validly published before 1904, then it would
have priority over C. betulus var. serrata (Beck) C.K.Schneid. (C. betulus [unranked] serrata Beck,
basion.).
C. intermedia Wierz. ex Rchb., Icones Florae Germanicae et Helveticae 12: 4, pl. DCXXXIII
(Reichenbach 1850). – Original citation: “in Bannatu, am Wege von der Sägemühle gegen Kirscharusch bei Steyersdorf [Anina in Caraș-Severin], [P.P]. Wierzbicki!”. – Type: not localized.
C. betulus var. provincialis Gren. & Godr., Flore de France 3 (1): 121 (Grenier & Godron 1855). –
C. provincialis J.Gay nom. nud. in sched. – Original citation: “Forêts, bois et taillis.” – Type: France
[Provence-Alpes-Côte d’Azur: Var], Pennafort près Draguignan, Jun. 1830, J.H. Perreymond 28
(lecto-, designated here: K!). – Additional original material: France, Draguignan, J.H. Perreymond
s.n. (K!); ibid., 1840, J.H. Perreymond s.n. (P00543785!). – Note: the specimen here designated as
lectotype is mounted with the original note of Gay from 1835, where he describes his new species
C. provincialis. In a note from 1 Dec. 1853 —also attached to the sheet— announcing a letter to
Grenier for the following day, Gay rejects his species as a youthful folly (“péché de ma jeunesse”).
Nevertheless, Grenier took up the name and published is as a variety in his Flore de France.
C. betulus var. obtusifolia G.Kirchn., Arboretum Muscaviense: 667 (Petzold & Kirchner 1864). –
C. betulus f. obtusifolia (G.Kirchn.) H.J.P.Winkl., Das Planzenreich IV 61 (Heft 19): 30 (Winkler
1904). – Original citation: No detailed data given. – Type: not localized. – Note: Petzold & Kirchner
accepted their term “Spielart” in the sense of “varietas” as only rank below the species (1864: 41).
Although they discuss other rank-denoting terms, such as “Unter-Art” (subspecies) and “Form”
(forma) as being widely used, they only accept “Spielarten”. Hence, names below the species rank
in Petzold & Kirchner (1864) are to be treated as varieties.
C. subcordifolia Schur, Enumeratio Plantarum Transsilvaniae: 611 (Schur 1866). – Original citation:
“In Wäldern der Kalkgebirge: auf dem Ecsem-Teteje bei Sz. – Domokos; auf dem Kereszthegy
bei Remete. Jul.” – Type: not localized. – Note: this name is missing in all relevant treatments and
current databases surveyed.
C. nervata Dulac, Flore du Département des Hautes-Pyrénées: 141 (Dulac 1867). – Type: France,
Hautes-Pyrénées, Lascazères, J.J. Corbin s.n. (syn-: BBF?); ibid., Saint-Sever-de-Rustan, J.J.
Corbin s.n. (syn-: BBF?).
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European Journal of Taxonomy 375: 1–52 (2017)
C. ulmifolia St.-Lag., Études des Fleurs ed. 8, 2: 742 (Cariot 1889) nom. illegit. superl. – Note:
Saint-Lager proposed this name to replace C. betulus L.
C. betulus var. haynaldiana Borbás, Oesterreichische Botanische Zeitschrift 39 (6): 234 (Braun
1889). – Original citation: “Im Auwinkel [Zugliget] bei Ofen [Buda].” – Type: Hungary [KözépMagyarország], in silvaticis ad Zugliget (Auwinkel) Budae-Pestini, 15 Jul. 1888, V. de Borbás s.n.
(syn-: BM!); ibid., 27 Jul. 1888, V. de Borbás s.n. (syn-: E!). – Note: this name is missing in all
relevant treatments and current databases surveyed.
C. betulus [unranked] serrata Beck, Flora von Nieder-Österreich: 266 (Beck von Mannagetta
1890). – C. betulus var. serrata (Beck) C.K.Schneid., Illustriertes Handbuch der Laubholzkunde 1:
140 (Schneider 1904). – C. betulus subsp. serrata (Beck) O.Schwarz, Mitteilungen der Thüringischen
Botanischen Gesellschaft 1: 96 (Schwarz 1949). – C. betulus f. serrata (Beck) Georgescu, Flora
Republicii Populare Române 1: 195 (Săvulescu 1952). – Original citation: No detailed data given. –
Type: not localized.
C. betulus var. typica Koehne, Deutsche Dendrologie: 193 (Koehne 1893) nom. inval. [Art. 24.3;
McNeill et al. (2012)].
C. betulus var. angustifolia Błocki ex Zapał., Conspectus Florae Galiciae Criticus 2: 9 (Zapałowicz
1908). – Original citation: “Lesienice pod Lwowem (Błocki), Strzyżów w Jasielskiem (Holzer).” –
Type: [Ukraine, Lviv] Lesienice bei Lemberg, 1886, B. Błocki s.n. (syn-: not localized; isosyn-:
B!, P06747360!, P06747390!, P06809764!, W1985–5965!); [Poland, Województwo podkarpackie]
Jasielski, Strzyżów, Holzer s.n. (syn-: not localized).
C. betulus [unranked] angustifolia Medw., Vestnik Tilisskogo Botanicheskogo Sada 14: 26 (Medwedew
1909) nom. illegit. (later homonym). – C. betulus var. angustifolia (Medw.) Radde-Fom., Trudy
izychno-matematychnogo viddilu. Kiev [Mémoires de la Classe des Sciences Physiques et
Mathématiques de l’Académie des Sciences de l’Ukraine] 15 (1): 80 (Radde-Fomina 1929)
nom. illegit. (later homonym). – C. betulus f. angustifolia (Medw.) Radde-Fom., Trudy izychnomatematychnogo viddilu. Kiev [Mémoires de la Classe des Sciences Physiques et Mathématiques
de l’Académie des Sciences de l’Ukraine] 15 (1): 80 (Radde-Fomina 1929) nom. inval. pro syn. –
Original citation: “найдена въ верхней Аджаріи (ущ. р. Аджарисъ-цкали въ Батумской обл.) ло
Годерзскаго перевала и за нимъ (въ бассейнѣ Куры, Масальскій, герб. Спб. Бот. Сада) а также
въ Терской области на хр. Салатау, между с. Чиркей и Хонцагой, на высотѣ 3000’ (Алекс.,
герб. Ак. Н.)” [Found in Upper Adsharia (Acharis-Tskali river in Batumi province), at Goderdzi
Pass and beyond (in Kura basin; Massalsky, Herb. of the St. Petersburg Botanical Garden). In Terek
region, in Salatau between Czirkei and Honzagoi, alt. 3000’ (Alexeenko, Herb. of the Academy).]. –
Type: not localized. – Note: Medwedew diferentiated three forms of C. betulus in the Caucasus,
but published his C. betulus [unranked] angustifolia explicitly as “var. nov.” Nowadays, the use
of the two diferent ranks in a non-successive order is to be interpreted as unranked (Art. 37.8;
McNeill et al. 2012). Radde-Fomina (1929: 80) accepted Medwedew’s informal use of “form” and
cited Medwedew’s name in the rank forma and gave it the rank as variety afterward, while citing
Medwedew’s form in the synonymy. Irrespective of this, Medwedew’s and Radde-Fomina’s names
are illegitimate due to the priority of Zapałowicz’s name from 1908.
C. betulus var. acuminata K.Reinecke, Mitteilungen des Thüringischen Botanischen Vereins 28: 41
(Reinecke 1911). – Original citation: “Im Glacis der Cyriaksburg b. E. [Erfurt, Germany].” – Type:
not localized.
C. betulus var. parva Radde-Fom., Trudy izychno-matematychnogo viddilu. Kiev [Mémoires de la
Classe des Sciences Physiques et Mathématiques de l’Académie des Sciences de l’Ukraine] 15 (1):
55, tab. VI f, g, e (Radde-Fomina 1929). – Original citation: “Dagestan, Agan-Kale prope Temirchan-Schura [former name of Buynaksk] 2. X. 1916. Budaev [s.n.]!”. – Type: not localized.
C. betulus f. subacuta Domin, Flora Čechoslovenica Exsiccata no. 245 (1931) in sched. – Type: Czech
Republic, Bohemia centralis, gregarie in valle Radotinské údoli prope urbem Praha, solo calcareo,
altitudine circa 250 m a.s.l., 10 Aug. 1931, K. Domin and M. Deyl s.n. (holo-: not localized; iso-: B10
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HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
0752246!, B!, B!, K!, MSB-114979!). – Note: this name is efectively published on specimen labels
accompanying the types (Art. 30.7; McNeill et al. 2012).
C. betulus var. tuzsoni Kárp., Botanikai Közlemények 34: 195 (Kárpáti 1937). – Original citation:
“Während einer Excursion unter der Leitung des Herrn Prof. Dr. J. v. Tuzson machten wir am
Jánoshegy bei Budapest […].”. – Type: not localized. – Note: this is a variety with issured bark.
C. caucasica Grossh., Izvestiya Azerbaidzhanskogo Filiala [Akademii Nauk SSSR] 1940 (5): 34
(Grossheim 1940). – Type: [Russia] Sala-tau, Czirkei × Honzagoi, ± 3000 ft, 28 Jul. 1897, T.
Alexeenko s.n. (holo-: LE). – Note: the holotype might be the same specimen as the one cited as
syntype of C. betulus [unranked] angustifolia Medw.
1a. Carpinus betulus ‘Quercifolia’
C. betulus var. quercifolia hort. ex C.F.Ludw. (Ludwig 1783). – C. quercifolia (C.F.Ludw.) Desf.,
Tableau de l’École de Botanique: 213 (Desfontaines 1804). – C. betulus f. quercifolia hort. ex
K.Koch, Dendrologie 2, 2: 3 (Koch 1873) nom. inval. pro syn. – C. betulus f. quercifolia (C.F.Ludw.)
C.K.Schneid., Illustriertes Handbuch der Laubholzkunde 1: 140 (Schneider 1904). – C. betulus var.
quercifolia (Desf.) Tzvelev, Flora Vostochnoĭ Evropy 11: 91 (Tzvelev 2004) nom. illegit. (later
homonym). – Original citation: not localized. – Type: Paris Botanical Garden [cult.], s. coll., s.n.
(neo-, designated here: P06747373!). – Note: the oak-leaved hornbeam is an early mentioned cultivar
that is usually attributed to Desfontaines (1804: 213). However, the mutation was known before
1783 when Ludwig published an oak-leaved, C. betulus var. quercifolia. The short description by
Ludwig (1783) was actually the English name, under which it was supposedly sold in an English
garden catalogue. Ludwig (1783) cites four English catalogues in the preface and mentions that
he did not translate any name himself. The name ‘Quercifolia’ does not appear in the Loddiges
catalogues (Loddiges 1779, 1783), while the remaining two could not be examined by the present
authors. Therefore, it might be possible that the name was published validly even before. As all
treatments deal with the same taxon, Art. 41.4 (McNeill et al. 2012), applies. The specimen chosen
as neotype is likely to represent what was and still is understood under this name. The end of the
left branch on the neotype has strong ainities to the cultivar ‘Incisa’, showing that these two may
not always be readily distinguished. Already Koch (1873) considered the two names as the same.
However, for now, it deems the present authors best to keep these two names separate.
C. betulus var. heterophylla hort. ex Loudon, Arboretum et Fruticetum Britannicum 3: 2005 (Loudon
1838) nom. inval. pro syn. – Note: this cultivar with an unusual leaf incision is sometimes regarded
as the same as ‘Incisa’, but in contrast to ‘Incisa’ with acute lobes and shallow teeth, the ones in
‘Quercifolia’ are rather obtuse and rounded (cf. Nicholson 1883; Schneider 1904). Koch regarded
C. betulus ‘Quercifolia’ as synonymous to ‘Incisa’ and published the epithet as C. betulus f. incisa
(Aiton) K.Koch (C. betulus var. incisa Aiton, basion.).
1b. Carpinus betulus ‘Incisa’
C. betulus var. incisa Aiton, Hortus Kewensis 3: 362 (Aiton 1789). – C. betulus f. incisa (Aiton) K.Koch,
Dendrologie 2, 2: 3 (Koch 1873). – Type: Kew [cult.], Oct. 1883, s. coll., s.n. (neo-, designated here:
K! ex ligneous herb. Mr. G. Nicholson s.n.). – Note: the specimen designated as neotype here is
mounted together with another specimen marked as “C. betulus var. asplenifolia”.
C. laciniata hort. ex G.Nicholson, The Garden (London 1871–1927) 24: 419 (Nicholson 1883) nom.
inval. pro syn. – Note: this variety, better to be treated as a cultivar, has leaves with a lobulate
margin, triangular lobes, acute apices, and a shallow dentation.
7
European Journal of Taxonomy 375: 1–52 (2017)
1c. Carpinus betulus ‘Variegata’
C. betulus var. variegata Lodd. ex Loudon, Arboretum et Fruticetum Britannicum 3: 2005 (Loudon
1838). – C. betulus f. variegata (Lodd. ex Loudon) H.J.P.Winkl., Das Planzenreich IV 61 (Heft 19):
31 (Winkler 1904). – Original citation: No detailed data given. – Type: not localized. – Note: this
is a cultivar with variegated leaves from Loddiges’ plant catalogue (Loddiges 1836). Winkler cites
neither Loudon nor Loddiges (1836) but Dippel (1891: 140), and according to Art. 41.4. (McNeill
et al. 2012), his name should be considered as a new combination as he is dealing with the same
cultivar. The color of the variegation is unknown, as Loudon does not mention it. Dippel (1891: 149)
recognizes only a single variegated cultivar, irrespective from the color of the variegation, while
Schneider (1904) recognizes a forma albovariegata and a forma aureovariegata.
1d. Carpinus betulus ‘Heterophylla’
C. betulus var. heterophylla G.Kirchn., Arboretum Muscaviense: 667 (Petzold & Kirchner 1864).
– C. betulus f. heterophylla (G.Kirchn.) K.Koch, Dendrologie 2 (2): 3 (Koch 1873). – Original
citation: No detailed data given. – Type: not localized. – Note: Kirchner gave the name “heterophylla”
to plants that have both normal-shaped leaves and deeply serrated leaves, regardless whether the leaf
serration is of the ‘Incisa’ or ‘Quercifolia’ type, which he regarded as synonymous, as did Koch.
As the mutation occurs spontaneously and may be ixed by cuttings, this name ‘Heterophylla’ is
complicated to use. Kirchner refers to “Bth. cat.”, meaning the catalogue of James Booth and Söhne
nursery in Klein-Flottbek, Hamburg, Germany. As there is no reference to Loudon, Kirchner’s name
cannot be regarded as a clear recombination of C. betulus var. heterophylla hort. ex Loudon, which
is an invalid name published as a synonym of C. betulus ‘Quercifolia’. Therefore, Kirchner’s name
is not illegitimate, but rather a later homonym.
1e. Carpinus betulus ‘Pendula’
C. [unranked] pendula Massé, Revue Horticole ser. 4 2: 271 (Massé 1853). – C. betulus var. pendula
(Massé) G.Kirchn., Arboretum Muscaviense: 667 (Petzold & Kirchner 1864). – C. pendula (Massé)
K.Koch, Dendrologie 2 (2): 3 (Koch 1873). – C. betulus f. pendula (Massé) Schelle, Handbuch
der Laubholzbenennung: 48 (Beissner et al. 1903). – Original citation: No detailed data given. –
Type: not localized. – Note: this is a cultivar with weak, overhanging branches. Massé mentions a
“Carpinus pendula” but immediately called it a variety of the common hornbeam. Therefore, the
taxonomic status is unclear upon publication, and the name is to be treated as unranked. Although
neither Koch nor Beissner et al. (1903) explicitly refer to Massé, the names are to be regarded as a
new combination (Art. 41.4; McNeill et al. 2012).
1f. Carpinus betulus ‘Purpurea’
C. betulus f. purpurea K.Koch, Dendrologie 2 (2): 3 (Koch 1873). – C. betulus var. purpurea (K.Koch)
G.Nicholson, The Garden (London 1871–1927) 24: 419 (Nicholson 1883). – Original citation: No
detailed data given. – Type: not localized. – Note: this is a cultivar with initially brownish-red leaves
that turn greenish soon after. Nicholson does not refer to Koch, but intended to describe the same
cultivar, therefore Art. 41.4 (McNeill et al. 2012) applies.
1g. Carpinus betulus ‘Fastigiata’
C. betulus var. fastigiata hort. ex G.Nicholson, The Garden (London 1871–1927) 24: 419 (Nicholson
1883). – C. betulus f. fastigiata (G.Nicholson) Schelle, Handbuch der Laubholzbenennung: 48
(Beissner et al. 1903). – Original citation: No detailed data given. – Type: not localized. – Note: the
8
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
branches of this cultivar are supposed to be “more ascending and the habit altogether more erect.”
Beissner et al. (1903) do not refer to Nicholson, but basically describe the same cultivar, therefore
Art. 41.4 (McNeill et al. 2012) applies.
C. betulus [unranked] pyramidalis Dippel, Handbuch der Laubholzkunde 2: 140 (Dippel 1891) [“1892”]
nom. illegit. superl. – C. betulus f. pyramidalis (Dippel) H.J.P.Winkl., Das Planzenreich IV 61 (Heft
19): 31 (Winkler 1904) nom. illegit. superl. – Note: Dippel (1891) and Winkler (1904) both cite
C. betulus ‘Fastigiata’ as a synonym, therefore their names are superluous.
1h. Carpinus betulus ‘Columnaris’
C. betulus [unranked] columnaris Beissn., Mitteilungen der Deutschen dendrologischen Gesellschaft 8:
136 (Beissner 1899b). – C. betulus f. columnaris (Beissn.) Schelle, Handbuch der Laubholzbenennung:
48 (Beissner et al. 1903). – C. betulus var. columnaris (Beissn.) Bean, Trees and Shrubs, Hardy
in the British Isles 1: 294 (Bean 1914). – Original citation: “In the gardens of Fürst Lobkowitz
[Ferdinand Zdenko von Lobkowitz] at Schloss Eisenberg [Jezeří Castle, Czech Republic]”. – Type:
not localized. – Note: according to Beissner et al. (1903), this cultivar was reported for the irst
time in a garden catalogue of the Späth’sche Baumschule in Berlin, Germany, in 1891, but the
present authors could not verify if that publication contains a valid description. Although neither
Beissner et al. (1903) nor Bean (1914) refer to Beissner (1899b), they all refer to a cultivar with
erect branches forming a slender crown, so their names can be considered as new combinations (Art.
41.4.; McNeill et al. 2012).
1i. Carpinus betulus ‘Pyramidalis’
C. betulus var. pyramidalis hort. ex Bean, Trees and Shrubs, Hardy in the British Isles 1: 294 (Bean
1914). – Original citation: “A ine specimen grows in the Solferino Square at Rouen.”. – Type: not
localized. – Note: this cultivar has erect branches, but not as slender as those of ‘Columnaris’. Bean
does not refer to Dippel (1891) or Winkler (1904), whose names are superluous.
1j. Carpinus betulus ‘Fastigiata Cucullata’
C. betulus f. cucullata hort. ex H.J.P.Winkl., Das Planzenreich IV 61 (Heft 19): 31 (Winkler 1904). –
C. betulus fastigiata cucullata G.Kirchn., Arboretum Muscaviense: 667 (Petzold & Kirchner 1864)
nom. inval. [polynomen as it was not published hyphenated, cf. Art. 20.3, Ex.7, McNeill et al.
(2012)]. – Original citation: No detailed data given. – Type: not localized. – Note: this is a garden
cultivar with pyramidal growth and hooded leaves. The cultivar name ‘Fastigiata Cucullata’ is
the earliest available one (Art. 21.6; Brickell et al. 2009), although not valid under the botanical
Code (Art. 23.6c; McNeill et al. 2012), under which it was independently published as C. betulus f.
cucullata by Winkler (1904).
1k. Carpinus betulus ‘Albovariegata’
C. betulus f. albovariegata C.K.Schneid., Illustriertes Handbuch der Laubholzkunde 1: 140 (Schneider
1904). – Original citation: No detailed data given. – Type: not localized. – Note: it was published as
“albo-variegata”, corrected according to Art. 60.9 (McNeill et al. 2012).
1l. Carpinus betulus ‘Aureovariegata’
C. betulus f. aureovariegata C.K.Schneid., Illustriertes Handbuch der Laubholzkunde 1: 140 (Schneider
1904). – Original citation: No detailed data given. – Type: not localized. – Note: it was published as
“aureo-variegata”, corrected according to Art. 60.9 (McNeill et al. 2012).
9
European Journal of Taxonomy 375: 1–52 (2017)
1m. Carpinus betulus ‘Horizontalis’
C. betulus [unknown rank] horizontalis Simon-Louis, “Preisverw., Herbst” 1902–1903 (SimonLouis 1902). – C. betulus f. horizontalis (Simon-Louis) C.K.Schneid., Illustriertes Handbuch der
Laubholzkunde 1: 140 (Schneider 1904) – C. betulus var. horizontalis (Simon-Louis) Bean, Trees
and Shrubs, Hardy in the British Isles 1: 294 (Bean 1914). – Original citation: No detailed data
given [found by M. Jouin on the Simon-Louis establishments near Metz, France (Bean 1914)]. –
Type: not localized. – Note: this is a cultivar with a lat crown. The present authors do not have the
Simon-Louis catalogue at hand. Depending on whether or not it contains a description and a deinite
rank, Schneider’s name would become valid when he provided both. If the publication by SimonLouis constitutes a valid publication with a deinite rank, either Schneider’s or Bean’s combination
might become superluous.
1n. Carpinus betulus ‘Asplenifolia’
C. betulus var. asplenifolia Bean, Trees and Shrubs, Hardy in the British Isles 1: 294 (Bean 1914). –
Original citation: No detailed data given. – Type: not localized. – Note: cultivar with incised leaves.
The incision depth is described as “the primary teeth large enough to be termed lobes”.
2. Carpinus caroliniana Walter
Flora Caroliniana: 236 (Walter 1788). – Original citation: No detailed data given. – Type: not localized.
C. ostryoides Raf., The Medical Repository ser. 3 2: 333 (Rainesque 1811) nom. nud., non Göppert
(1855). – Note: Göppert’s C. ostryoides is a valid name for a leaf fossil without reference to
Rainesque (1811).
2a. Carpinus caroliniana Walter subsp. caroliniana
Note: It was accepted as subspecies distinct from C. caroliniana subsp. virginiana (Marshall) Furlow in
the Flora of North America (Furlow 1997).
C. americana Michx., Flora Boreali-Americana 2: 201, pl. 7 (Michaux 1803). – Original citation:
“A Canadam ad Floridam.” – Original material: Ameriques septentrionales, A. Michaux s.n.
(P06810890!); ibid., s. coll., s.n. (P06810902!, P06810903!). – Note: Carpinus americana is
regarded as synonymous to C. caroliniana s. lat. in Flora of North America (Furlow 1997).
2b. Carpinus caroliniana subsp. virginiana (Marshall) Furlow
Systematic Botany 12: 429 (Furlow 1987). – C. betulus var. virginiana Marshall, Arbustrum Americanum:
25 (Marshall 1785). – C. virginiana (Marshall) Sudworth, Bulletin of the Torrey Botanical Club 20:
43 (Sudworth 1893) nom. illegit., non Miller (1768). – C. caroliniana var. virginiana (Marshall)
Fernald, Rhodora 37: 425 (Fernald 1935). – Type: s. loc., 16 Sep. 1906, R.W. Woodward s.n.
(neo- , designated by Furlow (1987: 429): NEBC00078970!). – Note: original material in Marshall
Herbarium, but it was destroyed (Furlow 1987).
3. Carpinus chingiana C.C.Yang
Distribution of the Woody Plants in Sichuan [四川树木分布]: 661 (Yang 1997). – Type (according to
IPNI): China, Sichuan, Muli, 2600 m a.s.l., 3 Aug. 1978, Q.-S. Zhao et al. [Zhao Zhenju, Mu Ke-hua,
Yang Yabin/赵清盛, 牟克华, 杨亚滨] 6980 (holo-: SZ00094298, SZ00094299; iso-: CDBI0172181!,
CDBI0172182!, CDBI0172183!). – Note: this name is accepted in the Flora Sichuanica (Editorial
Board of the Flora Sichuanica 2012), but neither listed in the World Checklist (Govaerts & Frodin
10
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
1998) nor in the Flora of China (Li & Skvortsov 1999). The CDBI specimens were labeled by Yang
as “Carpinus chingii”, and the specimen CDBI0172182 bears an annotation slip by Yang with the
unpublished name “Carpinus muliensis”.
4. Carpinus chuniana Hu
Journal of the Arnold Arboretum 13: 334 (Hu 1932). – Type: China, Kwangtung [Guangdong],
Lokchang, 31 May 1929, C.L. Tso [左景烈] 20872 (holo-: PE?; iso-: A00033751!, IBSC0001142,
IBSC0001143, IBSC0001144, NY00253858!).
5. Carpinus cordata Blume
Museum Botanicum Lugduno-Batavum 1 (20): 309 (Blume 1851). – Original citation: “Fisibami
japonice. In Japoniâ a Cl. Viro von Siebold detecta.” – Type: Japan, in Japoniâ, s. coll., s.n. (lecto-,
designated here: L0040896! ex herb. Siebold s.n.; isolecto-?: L0103212).
5a. Carpinus cordata Blume var. cordata
C. erosa Blume, Museum Botanicum Lugduno-Batavum 1 (20): 308 (Blume 1851). – Original citation:
“In Japoniâ.” – Type: Japan, s. coll. (lecto-, designated here: L0040895! ex herb. Siebold s.n.).
C. erosa var. microcarpa Hayashi, 林業試験場研究報告 [Research Report of the Forestry Experiment
Station] 57: 153 (Hayashi 1952). – Type: Japan, Ezo [Hokkaido], Prov. Hiyama, Mt. Gamushi, 8
Aug. 1952, Y. Hayashi s.n. (holo-: TFM).
C. erosa var. velutina Hayashi, 林業試験場研究報告 [Research Report of the Forestry Experiment
Station] 57: 153 (Hayashi 1955). – Type: Japan, Hondo, Prov. Rikuchiu, Yamagata in Kuji, 15 Aug.
1952, N. Karizumi s.n. (holo-: TFM).
Ostrya mandshurica Budischtschew ex Trautv., Trudy Imperatorskago S.-Peterburgskago Botaniceskago
Sada 9: 166 (Trautvetter 1884). – Type: not localized, likely in LE.
C. cordata var. pseudojaponica H.J.P.Winkl., Botanische Jahrbücher für Systematik, Planzengeschichte
und Planzengeographie 50 (Suppl.): 490 (Winkler 1914). – C. erosa var. pseudojaponica Miyabe &
Tatewaki, Bulletin of the Kyushu University Forests 21: 48 (Tatewaki 1953) nom. inval. – C. erosa
var. pseudojaponica (H.J.P.Winkl.) Miyabe & Kudo ex Horikawa, Acta Phytotaxonomica et
Geobotanica 15 (1): 13 (Horikawa 1953) nom. inval. pro syn. – Type: Japan, Insel Yesso, Mororan,
Sep. 1904, U.J. Faurie 5775b (holo-: B†) [on U.J. Faurie 5775 in B marked with b]. – Note: this
name is not mentioned in the World Checklist (Govaerts & Frodin 1998) or in the WCSP (2016),
but it is synonymized under C. cordata in the Flora of Japan (Ohwi 1965). Tatewaki published the
combination without reference to older literature or an own description, it is therefore invalid (Art.
41.3; McNeill et al. 2012). Horikawa referred to the combination as C. erosa var. pseudojaponica
(H.J.P.Winkl.) Miyabe & Kudo in synonymy only. The combination may have been published in
Icones of the essential forest trees of Hokkaido (Miyabe et al. 1920–1932) though, but the present
authors did not have access for veriication
C. cordata var. faurieana H.J.P.Winkl., Botanische Jahrbücher für Systematik, Planzengeschichte und
Planzengeographie 50 (Suppl.): 489 (Winkler 1914). – Type: Japan, Insel Yesso, Sapporo, Jun.
1891, U.J. Faurie 7111 (lecto-, designated here: P06747141!). – Additional type material: Japan,
Iwanai, Sep. 1904, U.J. Faurie 5777 (syn-: A00033752!, BM000580426!). – Note: this name is
not mentioned in the World Checklist (Govaerts & Frodin 1998) or the WCSP (2016). Based on the
subglabrous leaves and the locus classicus being on Hokkaido, it may well represent a distinct taxon.
C. cordata var. robusta H.J.P.Winkl., Botanische Jahrbücher für Systematik, Planzengeschichte und
Planzengeographie 50 (Suppl.): 489 (Winkler 1914). – Type: Japan, Insel Yesso, Mororan, Sep.
1904, U.J. Faurie 5775 (syn-: BM000580424!, E!). – Note: this name is not mentioned in the
World Checklist (Govaerts & Frodin 1998) or the WCSP (2016). As for C. cordata var. faurieana
11
European Journal of Taxonomy 375: 1–52 (2017)
H.J.P.Winkl. it is unlikely that this name is synonymous with the other accepted varieties because
of the relatively subglabrous nature and the locus classicus being on Hokkaido instead of the Asian
mainland.
C. cordata var. winkleri Radde-Fom., Trudy izychno-matematychnogo viddilu. Kiev [Mémoires de la
Classe des Sciences Physiques et Mathématiques de l’Académie des Sciences de l’Ukraine] 15 (1):
70 (Radde-Fomina 1929). – Original citation: “Wladiwostok 1864 Budischczev [s.n.]! «Suchaja
Padi» in viciniis pag. Rakovka 2. IX. 1915, Neczajeva et Gordeev! Orlinoe Gnieso ad marginem
sylvae 17. V. 1907. Palczevsky [s.n.]! Prov Primorskaja, lum Orlinoje Gniesdo IX. 1902 Palczevsky
[s.n.]!”. – Type: not localized.
C. cordata var. brevistachyus S.L.Tung, Bulletin of Botanical Research, Harbin 1 (1‒2): 139 (Tung
1981). – Type: China, Jilin, Linjiang [临江], 700 m a.s.l., 22 Jun. 1962, Y.L. Chou [周以良] 60007
(holo-: NEFI).
5b. Carpinus cordata var. chinensis Franch.
Journal de Botanique [Morot] 13: 202 (Franchet 1899). – C. cordata f. chinensis (Franch.) Nakai, The
Journal of the College of Science, Imperial University of Tokyo, Japan 31: 205 (Nakai 1911). –
C. chinensis (Franch.) C.Pei, Botanical Bulletin of Academia Sinica 2: 223 (Pei 1948). – Type: China,
Sutchuen [Sichuan], in ditione Tchen keou tin [Chengkou County], 1400 m a.s.l., P.G. (Reverend
Père) Farges 14 (lecto-, designated here: P06747063!; isolecto-: L!, P06747058!, P06747059!,
P06747060!). – Note: Carpinus cordata var. chinensis is accepted in the Flora of China (Li &
Skvortsov 1999), but it is segregated as C. chinensis in the Flora Sichuanica (Editorial Board of the
Flora Sichuanica 2012).
5c. Carpinus cordata var. mollis (Rehder) W.C.Cheng ex Y.Chen
Illustrated Manual of Chinese Trees and Shrubs: 163 (Chen 1937). – C. mollis Rehder, Journal of the
Arnold Arboretum 11: 154 (Rehder 1930). – Type: China, Sichuan, Sungpan hsien, side of stream,
17 Aug. 1928, W.P. Fang [方文培] 4245 (holo-: A00033760!; iso-: E00275495!, E00275496!,
K000859923!, NAS00070300, NY00253865!, P01903243!, PE00021928!, PE00021929!). – Note:
Carpinus cordata var. mollis is accepted in Flora of China (Li & Skvortsov 1999), but it is regarded
as a synonym of C. chinensis in the Flora Sichuanica (Editorial Board of the Flora Sichuanica 2012).
6. Carpinus dayongina K.W.Liu & Q.Z.Lin
Bulletin of Botanical Research, Harbin 6 (2): 143 (Liu & Lin 1986). – Type: China, Hunan, Dayong
[now Zhangjiajie], Tianmen shan, 1100 m a.s.l., 28 Jul. 1985, K.W. Liu [Liu Ke-wang] 33359 (holo-:
CSFI; iso-: PE). – Additional type material: ibid., 25 Aug. 1984, Y.T. Xiao [Xiao Yu-tan] 40700
(para-: not localized). – Note: the epithet is sometimes cited as “dayongiana” (Govaerts & Frodin
1998; WCSP 2016), but the authors consistently use “dayongina” (Liu & Lin 1986: 143, 145), and
correctly so (Art. 60 Rec. 60D, McNeill et al. 2012).
7. Carpinus faginea Lindl.
Plantae Asiaticae Rariores 2: 50 (Wallich 1831). – Type: India [Uttar Pradesh], Gurwal, Kamrup
[Kamroop] s.n. (holo-: CGE?).
C. latibracteata Lindl. nom. nud. in sched. – Note: this name is only known from one specimen. It also
bears the notes “Wallich H.I.” and “Lindley 1830”, however, neither Wallich nor Lindley were in
Kumaon in 1830. Likely, it is material from Wallich’s collection (by Robert Blinkworth?), and the
name was written onto the specimen by Lindley in 1830 [India, Kamaon [Kumaon], s. coll., s.n.
(K!)], but never published.
12
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
8. Carpinus fangiana Hu
Journal of the Arnold Arboretum 10: 154 (Hu 1929). – Type: China, Sichuan, Nanchuan Hsien, in
thickets, 1500–1800 m a.s.l., 1 Jun. 1928, W.P. Fang [方文培] 1351 (lecto-, designated by Lin
et al. (2007: 1248): PE00021890; isolecto-: A00033753!, E00275500!, E00275501!, IBSC0367917,
K000859928!, NY00253860!, P06747103!, PE00021891!, PE00021968!). – Additional type material:
ibid., W.P. Fang 1352 (para-: E!, IBSC0367916, K!, P06747102! PE00021888!, PE00021969!). –
Note: accepted in the Flora of China (Li & Skvortsov 1999), in the World Checklist (Govaerts &
Frodin 1998), and in the Flora Sichuanica (Editorial Board of the Flora Sichuanica 2012).
C. wilsoniana Hu, Journal of the Arnold Arboretum 10: 154 (Hu 1929). – Type: China, Sichuan, Mt.
Omei, in thickets, 1675–1800 m a.s.l., 9 Aug. 1928, W.P. Fang 2685 (lecto-, designated here:
PE00021889; isolecto-: A00033772!, E00275499!, E00275528!, IBSC0367915, K000859919!,
NAS00070302!, NY00253869!, P06810764!, PE00021970!).
9. Carpinus fargesiana H.J.P.Winkl.
Botanische Jahrbücher für Systematik, Planzengeschichte und Planzengeographie 50 (Suppl.): 507
(Winkler 1914). – Type: China, Sutchuen [Sichuan], in ditione Tchen keou tin [Chengkou County]
[as “Carpinus yedoensis”], P.G. (Reverend Père) Farges s.n. (lecto-, designated here: P06811425!;
isolecto-: P06811420!). – Additional type material: China, Sichuan, A. Henry 7053 (syn-: not
localized).
9a. Carpinus fargesiana H.J.P.Winkl. var. fargesiana
Note: It was accepted with C. fargesiana var. hwai (see C. fargesiana var. ovalifolia) as another variety
of C. fargesiana in the Flora of China (Li & Skvortsov 1999).
C. daginensis Hu, Acta Phytotaxonomica Sinica 9: 293 (Hu 1964). – Type: China, Sichuan, Dagin
Hsien, Aning, 2900 m a.s.l., 9 May 1958, X. Li [Li Hsien, 李馨] 77351 (holo-: PE00021885!;
iso-: NAS00070301, WUK0252019). – Note: Carpinus daginensis is considered as a synonym
of C. fargesiana var. fargesiana in the Flora of China (Li & Skvortsov 1999) and in the Flora
Sichuanica (Editorial Board of the Flora Sichuanica 2012: 35).
C. fargesiana var. tchouana C.C.Yang, Distribution of the Woody Plants in Sichuan [四川树木分布]:
662 (Yang 1997). – Type: not localized. – Note: this name is only accepted and mentioned in the
Flora Sichuanica (Editorial Board of the Flora Sichuanica 2012), while it is regarded as a synonym
of C. fargesiana var. fargesiana in the WCSP (2016). The present authors did not see the protologue.
9b. Carpinus fargesiana var. ovalifolia (H.J.P.Winkl.) Holstein & Weigend comb. nov.
urn:lsid:ipni.org:names:77167104-1
Basionym: C. turczaninovii var. ovalifolia H.J.P.Winkl., Botanische Jahrbücher für Systematik,
Planzengeschichte und Planzengeographie 50 (Suppl.): 505 (Winkler 1914). – C. hwai Hu &
W.C.Cheng, Bulletin of the Fan Memorial Institute of Biology n.s. 1: 148 (Hu 1948a). – C. fargesiana
var. hwai (Hu & W.C.Cheng) P.C.Li, Flora Reipublicae Popularis Sinicae 21 (2): 82 (Li 1979a)
nom. illegit. superl. – Type: China, Sichuan, No. [North] Wushan, A. Henry 7020 (lecto-, designated
here: P06811189!; isolecto-: BM!, GH00112538!, K000859934!, K000859935!). – Additional type
material: China, “N Huan tou shan”, G. Giraldi s.n. (syn-: not localized); ibid., Sichuan, S Wushan,
A. Henry 7019 (syn-: K000859936!); ibid., distr. Tchen-keou-tin, P.G. [Réverend père] Farges 1273
(syn-: P06811190!, P06811191!, P06811192!, P06811194!, P06811503!). Ibid., P.G. [Réverend
Père] Farges s.n. (syn-: not localized); ibid., Sichuan [locality unreadable], 7500 ft, May 1903,
E.H. Wilson 4489 (syn-, mentioned in appendix to protologue: BM!, IBSC0368215, K000859932!,
13
European Journal of Taxonomy 375: 1–52 (2017)
K000859933!). – Note: Hu (1948b) cited Winkler’s variety for their new species as the only element
(thus, C. hwai is a new name, but only legitimate in the rank of a species). When treating this
taxon as variety, however, again the name “ovalifolia” has priority, which Li apparently overlooked.
The lectotype material was mentioned in the appendix of the protologue, and is thus also original
material. It bears an annotation of Winkler, and duplicates are distributed in several herbaria, so it
seems a good choice for a lectotypiication. This taxon is recognized both in the Flora of China
(Li & Skvortsov 1999) and in the Flora Sichuanica (Editorial Board of the Flora Sichuanica 2012:
35).
10. Carpinus irmifolia (H.J.P.Winkl.) Hu
Bulletin of the Fan Memorial Institute of Biology n.s. 1 (2): 144 (Hu 1948a). – C. turczaninovii var. irmifolia
H.J.P.Winkl., Botanische Jahrbücher für Systematik, Planzengeschichte und Planzengeographie
50 (Suppl.): 505 (Winkler 1914). – C. pubescens var. irmifolia (H.J.P.Winkl.) Hu ex P.C.Li, Flora
Reipublicae Popularis Sinicae 21 (2): 80 (Li 1979a). – Type: China, Kui-Tscheu [Guizhou], Ma-jo,
fr, Sep. 1908, J. Cavalerie 3135 (holo-: E!; iso-: P04815672!). – Note: accepted as C. irmifolia in
the Flora of China (Li & Skvortsov 1999), but listed as a variety in the World Checklist of Fagales
(Govaerts & Frodin 1998).
C. pubescens var. bigiehensis Hu, Acta Phytotaxonomica Sinica 9: 290 (Hu 1964). – Type: China,
Kweichow [Guizhou], Bigieh Hsien, Lin-Ku’Gie-Wo, in open woods on rocky clifs, 1450 m a.s.l.,
13 Sep. 1957, P.H. Yü [Yü Pin-Hwa, 禹平华] 810 (holo-: PE; iso-: IBSC0368211, KUN0590865!,
LBG00053917, WUK206962). – Note: it was listed as a synonym of C. irmifolia in the Flora of
China (Li & Skvortsov 1999).
11. Carpinus hebestroma Yamam.
Supplementa Iconum Plantarum Formosarum 5: 14, ig. 4 (Yamamoto 1932). – Type: R. China, Taiwan,
inter Shinjio et Batakan, 26 Apr. 1917, s. coll. [Y. Yamamoto?], s.n. (lecto-, designated here: TAI
[118773]!). – Additional type material: R. China, Taiwan, inter Batakan et Naitaroko, Apr. 1917, Y.
Yamamoto s.n. (syn-: not localized).
12. Carpinus henryana (H.J.P.Winkl.) H.J.P.Winkl.
Botanische Jahrbücher für Systematik, Planzengeschichte und Planzengeographie 50 (Suppl.): 507
(Winkler 1914). – C. tschonoskii var. henryana H.J.P.Winkl., Das Planzenreich IV 61 (Heft 19):
36 (Winkler 1904). – C. hupeana var. henryana (H.J.P.Winkl.) P.C.Li, Flora Reipublicae Popularis
Sinicae 21 (2): 83 (Li 1979a) nom. illegit. – Type: China, Sichuan, A. Henry 7063 (holo-: B†; lecto-,
designated here: K00859926!; isolecto-: BM!, E00275504!, GH00033755!, P06811429!).
12a. Carpinus henryana (H.J.P.Winkl.) H.J.P.Winkl. var. henryana
C. henryana var. chuana C.C.Yang, Distribution of the Woody Plants in Sichuan [四川树木分布]: 661
(Yang 1997). – Type: not localized. – Note: this name is only accepted and mentioned in the Flora
Sichuanica (Editorial Board of the Flora Sichuanica 2012), while it is regarded as a synonym of
C. henryana var. henryana in the WCSP (2016). The present authors did not see the protologue.
12b. Carpinus henryana var. oblongifolia (Hu) Rushforth
Plantsman 8: 249 (Rushforth 1987). – C. turczaninovii var. oblongifolia Hu, Sunyatsenia 1 (2‒3): 115
(Hu 1933). – C. oblongifolia (Hu) Hu & W.C.Cheng, Bulletin of the Fan Memorial Institute of
Biology n.s. 1: 146 (Hu 1948a). – Type: China, Kiangsu [Jiangsu], Poa-Hwa Shan, 450 m a.s.l., 23
14
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
Jul. 1925, A.N. Steward 1476 (holo-: PE00021938!). – Note: this taxon is considered as a distinct
species in the Flora of China (Li & Skvortsov 1999).
C. huana W.C.Cheng, Contributions from the Biological Laboratory of the Science Society of China,
Botanical Series 9: 68 (Cheng 1933). – Type: China, Zhejiang, [Lin’an City, West Tianmushan], 21
Aug. 1924, W.C. Cheng 5161 (holo-: PE00021905!). – Additional type material: ibid., 15 Aug. 1931,
K. Ling 3519 (para-: not localized). – Note: Carpinus huana is regarded as a synonym of C. hupeana
in the Flora of China (Li & Skvortsov 1999).
C. hupeana Hu, Sunyatsenia 1 (2‒3): 118 (Hu 1933). – Type: China, Hupeh, Liang Sung kou, 1250 m
a.s.l., 9 Sep. 1922, W.Y. Chun et al. 4173 (holo-: PE00021898!). – Note: Carpinus hupeana is accepted
as distinct species in Flora of China (Li & Skvortsov 1999) with C. funiushanensis, C. huana and
C. longipes as synonyms.
C. longipes Hu, Acta Phytotaxonomica Sinica 9: 291 (Hu 1964). – Type: China, Kiangsi [Jiangsi],
Tasiping Shan, Shih-Cha-pu, 15 Jul. 1949, Y.K. Hsiung [Hsi un Yao-Kuo, 熊耀国] 5305 (lecto-,
designated by Lin et al. (2007: 1248): PE00021917!; isolecto-: PE00021915!, PE00021916!). –
Note: Carpinus longipes is regarded as a synonym of a distinct C. hupeana in the Flora of China
(Li & Skvortsov 1999).
C. funiushanensis P.C.Kuo, Flora Tsinlingensis 1 (2): 601 (Instituto Botanico Boreali-occidentali
Academiae Sinicae Edita 1974). – Type: China, Henan, 卢氏, 淇河北面山坡 [Lu-shih Hsien],
920 m a.s.l., 14 Sep. 1958, C.C. Fu [J.Q. Fu, 傅竞秋] 656 (holo-: WUK0407353, WUK0114527;
iso-: IBSC0367962, NAS00070304). – Note: Carpinus funiushanensis is regarded as a synonym of
a distinct C. hupeana in the Flora of China (Li & Skvortsov 1999). Kuo did not indicate a herbarium
for the holotype. As the Flora Tsinlingensis is mainly based on material from WUK, the specimens
there ought to be regarded as holo- and isotype respectively or as one holotype mounted on two
sheets if indicated as one gathering. The specimens from other herbaria are considered as duplicates.
12c. Carpinus henryana var. simplicidentata (Hu) Rushforth
Plantsman 8: 249 (Rushforth 1987). – C. simplicidentata Hu, Bulletin of the Fan Memorial Institute of
Biology n.s. 1: 143 (Hu 1948a). – C. hupeana var. simplicidentata (Hu) P.C.Li, Flora Reipublicae
Popularis Sinicae 21 (2): 83 (Li 1979a). – Type: China, Hubei, near Chin Tai Po, 2150 m a.s.l., 2
Sep. 1922, W.Y. Chun 4175 (lecto-, designated by Lin et al. (2007: 1249): PE00021955!; isolecto-:
PE00021953!, PE00021954!). – Additional type original material: China, Suon Nai Ook, 2150 m
a.s.l., 2 Sep. 1922, W.Y. Chun 4174 (syn-: not localized). – Note: this name is treated in Flora of
China (Li & Skvortsov 1999) as a synonym of C. stipulata (listed as synonym of C. turczaninovii
here).
13. Carpinus insularis N.H.Xia, K.S.Pang & Y.H.Tong
Journal of Tropical and Subtropical Botany 22 (2): 121 (Tong et al. 2014). – Type: China, Hong
Kong, Violet Hill, 22°14´25.30´´ N, 114°11´55.04´´ E, 190 m a.s.l., 21 Aug. 1913, K.Y. Tam s.n.
(holo-: IBSC0770924; iso-: HK0043873). – Note: according to the original authors C. insularis has
similarities to C. hebestroma and C. polyneura.
14. Carpinus japonica Blume
Museum Botanicum Lugduno-Batavum 1 (20): 308 (Blume 1851) – Distegocarpus carpinus Siebold &
Zucc., Abhandlungen der Mathematisch-Physikalischen Klasse der Königlich Bayerischen Akademie
der Wissenschaften 4 (3): 227 (Siebold & Zuccharini 1846). – Distegocarpus carpinoidis Siebold &
Zucc., Abhandlungen der Mathematisch-Physikalischen Klasse der Königlich Bayerischen Akademie
der Wissenschaften 4 (3): 240 (Siebold & Zuccharini 1846) orth. var. (of Distegocarpus carpinus).
– Carpinus carpinus (Siebold & Zucc.) Sarg., Garden and Forest 6: 364 (Sargent 1893) nom. inval.
15
European Journal of Taxonomy 375: 1–52 (2017)
(tautonym). – C. carpinoides (Siebold & Zucc.) Makino, The Botanical Magazine [Tokyo] 26: 391
(Makino 1912). – C. distegocarpus Koidz., The Botanical Magazine [Tokyo] 27: 144 (Koidzumi
1913) nom. illegit. superl. – Type: Japan, s. loc., H. Bürger s.n. (lecto-, designated by Akiyama
et al. (2013: 349): M-0120911!) – Additional original material: Japan, s. loc., H. Bürger s.n. (M0120912!, M-0120914!, M-0120915!, M-0120916!, M-0120917!); ibid., P.F. von Siebold et al. s.n.
(BR!, BR!, GH00033756!, K000859947!, K000859948!, L0175945!, P06811392?!). – Original
material?: s. loc. [via Java], s. coll., s.n. (U1155346!).
C. japonica var. caudata H.J.P.Winkl., Botanische Jahrbücher für Systematik, Planzengeschichte und
Planzengeographie 50 (Suppl.): 488 (Winkler 1914). – Type: Japan, Insel Nippon [Honshu], in den
Ontake-Bergen, Aug. 1905, U.J. Faurie 6641 (lecto-, designated here: BM!).
C. japonica var. cordifolia H.J.P.Winkl., Das Planzenreich IV 61 (Heft 19): 26 (Winkler 1904). –
C. carpinoides var. cordifolia (H.J.P.Winkl.) Makino, The Botanical Magazine [Tokyo] 26: 391
(Makino 1912) nom. illegit. – Type: Japan, Central Japan [Honshu], im Nikko-Gebirge, W.F.K.
Dönitz s.n. (holo-: B†).
C. japonica var. pleioneura H.J.P.Winkl., Botanische Jahrbücher für Systematik, Planzengeschichte und
Planzengeographie 50 (Suppl.): 488 (Winkler 1914). – Type: [Japan] Nikko, Prov. Schimotsuke,
Sep. 1887, s. coll., s.n. (lecto-, designated here: P06811395!) – Additional type material: Japan,
Miyanoschta, O. Warburg 7756 (syn-: B†?); ibid., Yokohama, May and Oct. 1862, C.J. Maximowicz
s.n. (syn-: P06811393!).
15. Carpinus kawakamii Hayata
Icones Plantarum Formosanarum 3: 175, pl. XXXIIIb (Hayata 1913). – Type: R. China, Taiwan, Monte
Morrison [Yushan], Oct. 1907, T. Kawakami and U. Mori 1998 (holo-: TI!). – Note: the type cited
here is the direct basis for the drawing in plate XXXIIIb, and can therefore be interpreted as the
holotype.
15a. Carpinus kawakamii Hayata var. kawakamii
Note: Accepted in the Flora of China (Li & Skvortsov 1999) with a distinct C. kawakamii var.
minutiserrata (Hayata) S.S.Ying, whereas Liao (1996) does not accept an infraspeciic taxon in the
Flora of Taiwan.
C. hogoensis Hayata, Icones Plantarum Formosanarum 6: 62 (Hayata 1916). – Type: R. China, Taiwan,
Musha [Wu sha], Hōgō [Chunyang], 3500 ft, Apr. 1916, B. Hayata s.n. (holo-: not localized; iso-:
TAIF!). – Note: Carpinus hogoensis is regarded as a synonym of C. kawakamii var. kawakamii in
the Flora of China (Li & Skvortsov 1999).
C. sekii Yamam., Supplementa Iconum Plantarum Formosarum 5: 12 (Yamamoto 1932). – Type: R.
China, Taiwan, in monte Daibusan, 3000 ft, 24 May 1918, E. Matsuda s.n. (holo-: TAI [118775]!). –
Note: Carpinus sekii is regarded as a synonym of C. kawakamii var. kawakamii in the Flora of China
(Li & Skvortsov 1999).
C. auriculifera Hayata nom. nud. – Note: C. auriculifera is only known from the specimen collected
in R. China (Ariko-banti, 1 Feb. 1917, E. Matsuda s.n., TAI [035707]!) and is marked as “type”. It
might have been published, but the protologue was not localized. The specimen, however, clearly
belongs to C. kawakamii.
15b. Carpinus kawakamii var. minutiserrata (Hayata) S.S.Ying
Coloured Illustrated Flora of Taiwan 3: 271 (Ying 1988). – C. minutiserrata Hayata, Icones Plantarum
Formosanarum 3: 177 (Hayata 1913). – Type: R. China, Taiwan, Tandaisha, Apr. 1910, U. Mori
s.n. (lecto-, designated here: TI!; isolecto-: TAIF007296!, TAIF007300!). – Note: Liao (1996)
16
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
synonymizes this name in the Flora of Taiwan under C. kawakamii, while it is accepted as separate
species in the Flora of China (Li & Skvortsov 1999).
16. Carpinus kweichowensis Hu
Sinensia 2: 79, ig. 1 (Hu 1931). – Type: China, Guizhou, Chengfeng Hsien, Pa-na, Si-mi-yao, mixed
woods, 24 Oct. 1930, Y. Tsiang [蒋英] 4406 (lecto-, designated here: NAS00070306!; isolecto-:
A00033757!, BM!, E00275503!, K000859925!, N000053496, NAS00070305!, NY00253862!,
PE00021906!, PE00021908!, US00089310!). – Note: the specimen selected here as lectotype is
annotated as holotype, but it was not published as such. It is also the specimen depicted in igure 1
in Hu (1931).
C. austroyunnanensis Hu, Bulletin of the Fan Memorial Institute of Biology n.s. 1 (3): 213 (Hu
1949). – Type: China, Yunnan Province, Xichou, Hsin cheih, 24 Oct. 1947, K.M. Feng 12609
[erroneously published as 13609] (lecto-, designated here: PE00020175!; isolecto-: PE00818041!,
WUK0207172). – Note: the type citation of C. austroyunnanensis is erroneous, as K.M. Feng 13609
(PE00792657) was collected the 23 Nov. 1947 and is a Morinda species, while K.M. Feng 12609
was annotated by Hu as C. austroyunnanensis.
17. Carpinus langaoensis Z.Qiang Lu & J.Quan Liu
Phytotaxa 295 (2): 191 (Lu et al. 2017). – Type: China, Shaanxi, Langao County, Hengxi Township,
108°48´ E, 32°08´ N, 1170 m a.s.l., 28 Jul. 2016, Z.Q. Lu 2016LZQ029 (holo-: LZU; iso-: LZU).
18. Carpinus laxilora (Siebold & Zucc.) Blume
Museum Botanicum Lugduno-Batavum 1 (20): 309 (Blume 1851). – Distegocarpus laxilora Siebold &
Zucc., Abhandlungen der Mathematisch-Physikalischen Klasse der Königlich Bayerischen Akademie
der Wissenschaften 4 (3): 228 (Siebold & Zuccharini 1846). – Type: Japan, s. loc., H. Bürger s.n.
(lecto-, designated by Akiyama et al. (2013: 349): M-0120909!). – Additional original material:
Japan, s. loc., H. Bürger s.n. (M-0120907!, M-0120908!); ibid., s. coll. (L0175947! ex herb. P.F. von
Siebold s.n., L0175948! ex herb. P.F. von Siebold s.n.); ibid., s. coll. (K000859944!, P06811472!,
P06811473!, P06811475!).
C. laxilora f. lacera Hayashi, Journal of Geobotany [北陸の植物] 22: 4 (Hayashi 1974). – Type: not
localized.
C. laxilora var. chartacea H.Lév., Bulletin de la Société botanique de France 51: 424 (Léveillé 1904).
– Original citation: without collecting data. – Type: not localized.
C. laxilora var. pendula Miyoshi, Report of the National Monument Investigation. Plants [天然紀念
物調査報告 植物之部 第 8輯] 8: 45 (Department of the Interior 1928). – C. laxilora f. pendula
(Miyoshi) Sugim., New Keys to Japanese Trees [日本樹木総検索誌]: 117, 458 (Sugimoto 1961). –
Original citation: “Ein einziges Exemplar im Grundstück von Jojuin Tempel, Tochigi Präfektur.”
[A single exemplar on the area of the Jojuin temple, Tochigi prefecture”; 成就院 in Niregimachi,
Kanuma district]. – Type: not localized. – Note: this is tree with hanging branches, better considered
as a cultivar. The tree (most likely a clone) still existed in 2013.
C. laxilora var. macrophylla Nakai, The Botanical Magazine [Tokyo] 45: 112 (Nakai 1931). – C. laxilora
f. macrophylla (Nakai) W.T.Lee, Lineamenta Florae Koreae: 165 (Lee 1996). – Type: South Korea
[Gyeonggi-do], in silvis Kôryô prov. Keiki, Gwangreung, 3 Sep. 1930, T. Nakai 13201 (holo-: TI).
C. laxilora var. obtusisquama Koidz., Acta Phytotaxonomica et Geobotanica 9 (2): 73 (Koidzumi
1940). – Type: South Korea, Chiisan [Mt. Jiri-san], Kokiri, 11 Jun. 1935, G.S. Koidzumi s.n. (holo-:
KYO). – Note: this name is mentioned neither in the World Checklist (Govaerts & Frodin 1998) nor
in the Checklist of Korea (Chang et al. 2014a) nor in the WCSP (2016), synonymization unclear.
17
European Journal of Taxonomy 375: 1–52 (2017)
C. laxilora var. longispica Uyeki, Suigen Gakuho 41: 9 (Uyeki & Lee 1924). – Type: South Korea,
Chyolla australis, mont Chokē (holo-: SNUA†). – Type: South Korea, Jeollanam-do, Suncheon City,
Songkwang-myeon, Seungju-eup, Jukhak-ri, Mt. Jogye-san, near the stream in front of Songkwang
Temple, 30 Jun. 1993, J.I. Jeon and D.J. Ha 10005 (neo-, designated by Chang & Chang (2010:
275): SNUA; isoneo-: KH). – Note: Chang et al. (2014a) accept C. laxilora var. longispica as a
distinct taxon with C. laxilora var. macrothyrsa as its synonym.
C. laxilora var. macrothyrsa Koidz., Acta Phytotaxonomica et Geobotanica 9 (2): 73 (Koidzumi
1940). – Type: South Korea, Mt. Chiisan [Jirisan], Hannyaho, Gawun [Gurae-jun], [Jeollanam-do],
2 Jun. 1935, S. Okamoto s.n. (lecto-, designated by Chang & Chang (2010: 276): KYO). – Note: this
name is regarded as a synonym of C. laxilora var. longispica in Chang et al. (2014a).
C. laxilora var. gosenensis Franch. nom. nud. – Note: this name is written on the specimen only: Japan,
Niigata, Montagnes de Gosen, 28 Jul. 1874, U.J. Faurie s.n. (P06811469!).
C. laxilora var. onoei nom. nud. – Note: this name is written on the specimens only: Japan, [Chūbu],
Ono [Ōno], L. Savatier 2935 (P06811464!); ibid., s. coll., s.n. (P06811462!).
19. Carpinus lipoensis Y.K.Li
Guizhou Science 1983 (2): 20 (Li & Mang 1983). – Type: China, Guizhou, Lipo, in sylvis montium
calcareum, 850 m a.s.l., Y.K. Li [李永康] 9940 (holo-: HGAS? ex “Herb. Inst. Biol. Guizhou”). –
Note: this name is neither listed in the World Checklist (Govaerts & Frodin 1998) nor in the Flora
of China (Li & Skvortsov 1999).
20. Carpinus londoniana H.J.P.Winkl.
Das Planzenreich IV 61 (Heft 19): 32 (Winkler 1904). – Type: China, Yunnan, 4000 ft, A. Henry 11640
(lecto-, designated here: K000859945!; isolecto-: A00033759!, B†, E00275498!, K000859946!,
MO-2140881!, MO-2140882!, NY00253864!).
20a. Carpinus londoniana H.J.P.Winkl. var. londoniana
C. poilanei A.Camus, Bulletin de la Société botanique de France 76: 968 (Camus 1930). – Type:
Vietnam, Annam, Lang-bian, entre Dran et Dung-Lo, E. Poilane 3983 (lecto-, designated here:
P06811169!). – Additional type material: Vietnam, Dalat en face de Manline, F. Evrard 1168 (syn-:
P06811171!).
C. poilanei var. chevalieri A.Camus, Bulletin de la Société botanique de France 76: 969 (Camus 1930). –
Type: Vietnam, Annam, plateau du Langbian, entre Bellevue et Dran, 30 Apr. 1919, A. Chevalier
40445 (lecto-, designated here: P06747035!; isolecto-: P06747037!, P06811163!). – Additional type
material: Vietnam, Belle [Bellevue], 17 Jun. 1921, B. Hayata 707 (syn-: P06811154!); ibid., Dalat, chemin de la montagne de l’éléphant ruisseau de Prenh, 29 Nov. 1924, F. Evrard 1918 (syn-:
P06811160!).
20b. Carpinus londoniana var. lanceolata (Hand.-Mazz.) P.C.Li
Flora Reipublicae Popularis Sinicae 21 (2): 68 (Li 1979a). – C. lanceolata Hand.-Mazz., Oesterreichische
Botanische Zeitschrift 80: 338 (Handel-Mazzetti 1931). – Type: China, Hainan, Ngitse-leng,
G. Fenzel 159 (holo-: not localized). – Additional type material: China, Hainan, secus rivos in media
insula, 300–500 m a.s.l., Oct.–Nov. 1929, G. Fenzel 106 (para-: not localized); ibid., Ngitse-leng, in
glarea, G. Fenzel 164 (para-: not localized).
18
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
20c. Carpinus londoniana var. latifolia P.C.Li
Acta Phytotaxonomica Sinica 17 (1): 87 (Li 1979b) [corrected from “latifolius”]. – Type: China,
Chekiang [Zhejiang], Ning-po [Ningbo (宁波)], s. coll. 1018 (holo-: PE).
20d. Carpinus londoniana var. xiphobracteata P.C.Li
Acta Phytotaxonomica Sinica 17 (1): 87 (Li 1979b). – Type: China, Chekiang [Zhejiang], Yin Hsien
[Yin Xian (鄞县)], G.R. Chen [Chen Gun-rung] 2289 (holo-: PE).
21. Carpinus luochengensis J.Y.Liang
Guihaia 6 (4): 275 (Liang 1986). – Type: China, Guangxi, Luocheng, 28 Jul. 1983, J.Y. Liang K1644
(holo-: IBK00190892!). – Additional type material: ibid., 5 Aug. 1984, F.N. Wei [Wei Fa-Nan, 韦
发南] 1743 (para-: IBK00190885!, IBK00190886!). – Note: not mentioned in the Flora of China
(Li & Skvortsov 1999).
22. Carpinus mengshanensis S.B.Liang & F.Z.Zhao
Bulletin of Botanical Research, Harbin 11 (2): 33 (Liang & Zhao 1991). – Type: China, Shandong,
Pingyi Xian, Meng Shan, 750 m a.s.l., Oct. 1984, F.Z. Zhao 84001 (holo-: SDFS). – Note: not
mentioned in the Flora of China (Li & Skvortsov 1999).
23. Carpinus microphylla Z.C.Chen ex Y.S.Wang & J.P.Huang
Guihaia 5 (1): 15 (Wang & Huang 1985). – Type: China, Guangxi, Tianyang Xian, 750 m a.s.l., 16
May 1964, Z.C. Chen 54089 (holo-: IBK00079486!). – Note: the holotype location is given by the
authors as “HIBG” but according to the Index Herbariorum (Thiers 2016), HIBG is the herbarium
of Hiroshima Botanical Garden. According to the Mandarin text, the holotype is deposited in the
“Guangxi Institute of Botany”, which is IBK.
24. Carpinus mollicoma Hu
Bulletin of the Fan Memorial Institute of Biology n.s. 1 (3): 216 (Hu 1949). – C. polyneura var. mollicoma
(Hu) C.C.Yang, Distribution of the Woody Plants in Sichuan [四川树木分布]: 42 (Yang 1997). –
Type: China, Yunnan, Xichou, Faa-doou, 25 Sep. 1947, K.M. Feng 12053 (lecto-, designated here:
PE00021946!; isolecto-: IBSC0368097, PE00818159!). – Note: Carpinus mollicoma is accepted in
the Flora of China (Li & Skvortsov 1999) and the World Checklist (Govaerts & Frodin 1998). Yang
in Flora Sichuanica (Editorial Board of the Flora Sichuanica 2012: 35) notes that his combination
C. polyneura var. mollicoma (Hu) C.C.Yang is a nom. nud. [comb. inval.?] in the 1997 publication,
but he treats it as a distinct variety in C. polyneura in 2012. If there was no reference to the basionym
in the cited reference, then the new combination was published in 2012.
25. Carpinus monbeigiana Hand.-Mazz.
Anzeiger der oesterreichischen Akademie der Wissenschaften mathematisch-naturwissenschaftliche
Klasse 61: 162 (Handel-Mazzetti 1925). – Original citation: “In silvis dumetisque calide temperatis
inter Dawan et Gwanyilang, legi 3. VII. 1914 (Nr. 3431) et supra Schidsilu, legi 30. VI. 1914 (Nr.
3331) prope urbem Yungbe. Certe in subtropicis ad luvium Mekong leg. Monbeig 1908 ibique
observavi infra vicum Gangpi eandemque verosimiliter ad l. Salwin circa Tschammutong pluries.” –
Type: China, Yunnan, prope urbem Yungbei, in regionis calide temperatae silvis inter vicos Dawan et
Swanyilang, 2400–2600 m a.s.l., 3 Jul. 1914, H.R.E. von Handel-Mazzetti 3431 (lecto-, designated
19
European Journal of Taxonomy 375: 1–52 (2017)
here: WU030928!; isolecto-: E00275494!, K!). – Additional type material: China, Yunnan, prope
urbem Yungbei, in regionis calide temperatae dumetis supra vic. Schidsilu, 2400–2700 m a.s.l., 30
Jun. 1914, H.R.E. von Handel-Mazzetti 3324 (syn-: WU030929!); ibid., am Mekong, 2400‒2600 m
a.s.l., J.T. Monbeig 1908 (syn-: not localized).
C. densispica Hu, Acta Phytotaxonomica Sinica 9: 286 (Hu 1964). – Type: China, [Yunnan], Tehchin
Hsien, Tzichung, Sila, 2500‒2800 m a.s.l., in mixed forest, 19 [in protologue “18”] Jul. 1940, K.M.
Feng 5618 (lecto-, designated here: PE00021886!; isolecto-: PE01062146!). – Additional type
material: China, [Yunnan], Weisi Hsien, Kangpu, at 2300 m a.s.l., in mixed woods, Jul. 1938, C.W.
Wang [王启无] 64343 (para-: PE01062148!).
C. likiangensis Hu, Acta Phytotaxonomica Sinica 9: 287 (Hu 1964). – Type: China, Yunnan, Likiang
Hsien, Snow range, 2700 m a.s.l., in mixed forest, 18 Aug. 1942, K.M. Feng 9055 (lecto-, designated
by Lin et al. (2007: 1248): PE00021913!; isolecto-: PE00021914!). – Additional type material: ibid.,
2400 m a.s.l., 19 Sep. 1955, K.M. Feng 21595 (para-: PE00021912!, PE00818189!).
C. monbeigiana var. weisiensis Hu, Acta Phytotaxonomica Sinica 9: 285 (Hu 1964). – Type: China,
Yunnan, Weisi Hsien [Weixi Lisu Autonomous County], Yehchi, 2040 m a.s.l., 24 Aug. 1956, P.Y.
Mao [Mao Ping-yi/Mao Pin-yih, 毛品一] 197 (lecto-, designated here: PE00021933!; isolecto-:
PE00021934!).
26. Carpinus omeiensis Hu & Fang
Acta Phytotaxonomica Sinica 9: 296 (Hu 1964). – Type: China, Sichuan, Mount Omei [Éméi Shān],
Dashiaochenfeng, 1900 m a.s.l., 7 Oct. 1957, K.H. Yang [Yang Kuang-Hway, 杨光辉] 57490
(lecto-, designated here: PE00021939!; isolecto-: IBSC0368104, KUN0590785!, NAS00070308,
PE00818214!). – Additional type material: China, Guizhou, Teking Hsien, Yaimenko, 1200 m
a.s.l., in dense forest on slope, 15 Aug. 1959, North Kweichow Exped. [黔北队] 1631 (para-:
KUN0590783!, NAS00070309!, PE00818217!, PE00818218!).
27. Carpinus orientalis Mill.
The Gardeners Dictionary ed. 8, no. 3 (Miller 1768). – C. nigra Moench, Verzeichniss ausländischer
Bäume und Stauden: 19 (Moench 1785) nom. illegit. superl. – Type: Oriens, J.P. de Tournefort
s.n. (lecto-, designated here: BM!). – Additional original material: Oriens, J.P. de Tournefort s.n. (P
[Tournefort 5561], LINN no. HS1481.2!, LINN no. HS1481.3!). – Note: the Tournefort specimen
in BM was designated to be lectotype because 1) Miller cited Tournefort’s Corollarium T. 40 in the
protologue, just as given on the label and 2) Miller’s material is deposited in BM (Staleu & Cowan
1976–1988). The other specimen in BM, dating from that time (BM001041899), is a cultivar from
Leiden (“Hort. Bni. [botanici] Boerhaave”). Miller most likely had material from Boerhaave at hand,
but he did not mention cultivated material from Leiden in the protologue. Hence it be or may not be
that this specimen could be considered as original material. There are two C. orientalis specimens in
the Smith Herbarium (LINN-HS 1481.2 and LINN-HS 1481.3). One (LINN-HS 1481.3) refers only
to Tournefort’s Institutiones Rei Herbariae (Tournefort 1700) and the other one to both Tournefort’s
publications (Tournefort 1700, 1703) but not to the actual page number in contrast to the specimen in
BM. Additionally, Miller did not refer to the Institutiones Rei Herbariae nor do the LINN specimens
match the description since they have (sub-)acute leaf apices, while Miller explicitely refers to
obtuse apices. Hence, we do not believe that Miller saw the material in the Smith Herbarium for his
description. The lectotypiication with the Tournefort specimen in Paris by Olshanskyi (2014: 68)
is not efective because the author forgot the phrase “designated here” or an equivalent (Art. 7.10,
McNeill et al. 2012). We deviate from his suggestion because Miller’s herbarium is in fact now
located in BM (Staleu & Cowan 1976–1988), hence lectotypes of his names should preferably be
placed there.
20
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
27a. Carpinus orientalis Mill. var. orientalis
C. duinensis Scop., Flora Carniolica ed. 2, 2: 243 (Scopoli 1772). – Original citation: “circa Duinum”
[Duino]. – Type: not localized.
C. minor Pall., Bemerkungen auf einer Reise in die Südlichen Statthalterschaften des Russischen Reichs
2: 95 (Pallas 1801). – Original citation: “Sobald man den Belbek [a river], etwa zwölf Werste von
Bachtschisarai [Bakhchysarai] durchfahren hat.” [This is a locality on Crimea, and the plant must
have been collected soon after Easter 1793]. – Type: not localized.
C. betulus var. edentula Heuf., Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien 8:
196 (Heufel 1858). – C. edentula Kit., Descriptiones et Icones Plantarum Rariorum Hungariae 2:
XXXII (Waldstein & Kitaibel 1803‒1805) nom. nud. – Original citation: “Smyrmium et Banatum
inhabitant.” – Type: In Comitatu Syrmiensi et in sylvis Banatus atque Serviae, P. Kitaibel A57 (lecto-,
designated here: BP-XL/144!; isolecto-: B-W 17700–030!). – Additional original material: s. loc.,
A. Wolny s.n. (BP-XL/86! ex herb. P. Kitaibel). – Note: Carpinus edentula is a nom. nud. appearing
irst in Kitaibel’s diary in 1804 (cf. Mólnar 2015). Spach (1842: 221) separates this name in a key,
and refers to a “Carpinus edentula Roch. Bann.”. Rochel (1828: 26) just listed this name as having
been communicated by Kitaibel. Steudel (1840: 300) published this name again, but only mentions
the perennial life form and the origin, so his publication cannot be considered as valid, and Spach
(1842: 221) published it as synonym to an invalid C. betulus var. β. In 1858, Heufel recognized
Kitaibel’s name as a variety of C. betulus and amended it with a description thus validating the name.
C. orientalis f. umbraculifera (Beissn.) Schelle, Handbuch der Laubholzbenennung: 49 (Beissner
et al. 1903). – C. orientalis [unranked] umbraculifera Beissn., Mitteilungen der Deutschen
dendrologischen Gesellschaft 8: 133 (Beissner 1899a). – C. betulus [unranked] umbraculifera hort.
ex Beissn., Mitteilungen der Deutschen dendrologischen Gesellschaft 8: 133 (Beissner 1899a) nom.
inval. (not accepted by the author). – Original citation: “Schloss Eisenberg ist herrlich oben am dicht
bewaldeten Erzgebirge gelegen, im tiefer liegenden Park”. – Type: not localized. – Note: Beissner
mentioned a cultivar with a spherical crown that has been called C. betulus umbraculifera, but he
supposed it to be a C. orientalis. A deinite rank was given in 1903.
C. orientalis f. calcarea Radde-Fom., Trudy izychno-matematychnogo viddilu. Kiev [Mémoires de la
Classe des Sciences Physiques et Mathématiques de l’Académie des Sciences de l’Ukraine] 15 (1):
95 (Radde-Fomina 1929). – Original citation: “Prov. Czernomorskaja, distr. Novorossijsk, ad
capitem Dob, prope pag. Kabardinka. 30 VI. 1912, [D.I.] Sosnowsky!” – Type: not localized.
C. orientalis var. coronata Radde-Fom., Trudy izychno-matematychnogo viddilu. Kiev [Mémoires de la
Classe des Sciences Physiques et Mathématiques de l’Académie des Sciences de l’Ukraine] 15 (1):
107 (Radde-Fomina 1929). – C. coronata Schischk. nom. nud. – Original citation: “Anatolien,
Trapezund, unterer Teil des Deïrmen-dere-Tales; Gebüsch westlichen Abhängen, 12 Juni 1917.” –
Type: not localized (LE?).
C. orientalis var. gibbosa Radde-Fom., Trudy izychno-matematychnogo viddilu. Kiev [Mémoires de la
Classe des Sciences Physiques et Mathématiques de l’Académie des Sciences de l’Ukraine] 15 (1):
107 (Radde-Fomina 1929). – C. ovata Schischk. ex Radde-Fom., Trudy izychno-matematychnogo
viddilu. Kiev [Mémoires de la Classe des Sciences Physiques et Mathématiques de l’Académie des
Sciences de l’Ukraine] 15 (1): 107 (Radde-Fomina 1929) nom. inval. pro syn. – Original citation:
“Anatolien, Sandshak Trapezund, Schlucht Gadshevara, zwischen Platana und Fol; Abhänge
mit Gebüsch. 27. Juni 1917.” – Type: not localized (LE?). – Note: Carpinus ovata was a name
suggested by Schischkin on a herbarium specimen that he had sent to Radde-Fomina for revision.
She suggested that this plant should be called “C. orientalis var. gibbosa” instead of being described
as a new species.
C. orientalis f. grandifolia Radde-Fom., Trudy izychno-matematychnogo viddilu. Kiev [Mémoires
de la Classe des Sciences Physiques et Mathématiques de l’Académie des Sciences de l’Ukraine]
21
European Journal of Taxonomy 375: 1–52 (2017)
15 (1): 94 (Radde-Fomina 1929). – Original citation: “Tauria ad viam e Simpheropol in Aluschtam”
[Alushta], VIII. 1925, leg. N. Dubowik!” – Type: not localized.
C. paucicostata Schischk. ex Radde-Fom., Trudy izychno-matematychnogo viddilu. Kiev [Mémoires
de la Classe des Sciences Physiques et Mathématiques de l’Académie des Sciences de l’Ukraine]
15 (1): 107 (Radde-Fomina 1929) nom. inval. pro syn. – Note: Radde-Fomina regarded Schischkin’s
plant as a mere C. orientalis. However, she indicated that he intended to describe the name later on.
This name is therefore provisional and invalid here. The present authors were unable to locate a
later, validating publication.
C. orientalis f. banatica Kárp., Botanikai Közlemények 34: 194 (Kárpáti 1937). – Original citation: “Im
Banat bei Herkulesbad” [Băile Herculane, Romania]. – Type: not localized. – Note: this is supposed
to be a narrow-leaved form.
27b. Carpinus orientalis subsp. macrocarpa (Willk.) Browicz
Flora Iranica 97: 2 (Browicz 1972). – C. orientalis var. macrocarpa Willk., Forstliche Flora von
Deutschland und Oesterreich ed. 2: 368 (Willkomm 1887). – C. macrocarpa (Willk.) H.J.P.Winkl.,
Das Planzenreich IV 61 (Heft 19): 38 (Winkler 1904). – C. orientalis f. macrocarpa (Willk.) RaddeFom., Trudy izychno-matematychnogo viddilu. Kiev [Mémoires de la Classe des Sciences Physiques
et Mathématiques de l’Académie des Sciences de l’Ukraine] 15 (1): 90 (Radde-Fomina 1929) nom.
nud. pro syn. – Original citation: “In Turkomanien hat [G.S.] Karelin”. – Type: not localized. – Note:
The published combination C. orientalis f. macrocarpa as a synonym was a misunderstanding by
Radde-Fomina. She believed that Medwedew (1909: 32) regarded the taxon as a forma. However,
he used the term “form” informally only, because “form” appears in only the text (Medwedew
1909: 32) while “var. macrocarpa” is written explicitly in the appendix (Medwedew 1909: 41).
Medwedew’s name is unranked, but would have been a later homonym anyway.
28. Carpinus paohsingensis W.Y.Hsia
Contributions from the Institute of Botany, National Academy of Peiping 2: 179 (Hsia 1934). – Type:
China, Sichuan, Paohsing-hsien (Mupin) [Baoxing], 1960 m a.s.l., 8 Jul. 1933, T.H. Tu 4356 (lecto-,
designated here: PE00021940!; isolecto-: IBK00079502!, IBSC0368268, PE00021941!). – Note:
Carpinus paohsingensis is accepted in the World Checklist (Govaerts & Frodin 1998), but accepted
neither in the Flora of China (Li & Skvortsov 1999) nor by Chang et al. (2014a) but listed as
synonymous with C. tschonoskii.
C. falcatibracteata Hu, Acta Phytotaxonomica Sinica 9: 297 (Hu 1964). – C. tschonoskii var.
falcatibracteata (Hu) P.C.Li, Flora Reipublicae Popularis Sinicae 21 (2): 85 (Li 1979a). – Type:
China, Sichuan, Hunghwa to Wali, 1900 m a.s.l., 14 Jul. 1959, Szechuan Econ. Bot. Liangshan
Exp. [四川经济植物(凉山)调查队] 1219 (lecto-, designated here: PE00021887!; isolecto-:
CDBI0172170!, PE00802589!). – Additional type material: China, Sichuan, Le-Po-Hsien, on
open mountain slope, 1950 m a.s.l., 8 Aug. 1934, T.T. Yu [俞德浚] 3639 (para-: IBSC0368265,
IBSC0368266, IBSC0368269, PE00802576!, WUK0330666); ibid., Liang Ho-Ko, in forest, 1700 m
a.s.l., 28 Sep. 1930, F.T. Wang 22597 (para-: PE00802565!); ibid., Kiangsi [Jiangxi], Shui Swe
Hsien, Mofu Shan, along gully, 10 Sep. 1947, Y.K. Hsiung [Hsiung Yao-Kuo/熊耀国] 5830 (para-:
LBG00001770!, PE00802561!, PE00802562!, PE00802563!). – Note: this name is neither accepted
in the World Checklist (Govaerts & Frodin 1998) nor in the Flora of China (Li & Skvortsov 1999)
nor by Chang et al. (2014a). It is regarded in the irst treatment as a synonym of C. paohsingensis,
which itself is treated as synonymous with C. tschonoskii in the latter two treatments. In the Flora
Sichuanica (Editorial Board of the Flora Sichuanica 2012), it is accepted pro parte minore as distinct
variety of C. tschonoskii, while the rest ought to be synonymous with C. tschonoskii and with
C. paohsingensis as full synonym of C. tschonoskii.
22
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
C. obovatifolia Hu, Acta Phytotaxonomica Sinica 9: 289 (Hu 1964). – Type: China, Yunnan, Tsenyih
Hsien, Shiao-Ma-La in forest, 1954, Y.H. Li [Li Yen-hei, 李延辉] 150 (lecto-, designated by Lin et al.
(2007: 1249): PE00021937!; isolecto-: PE00021936!). – Note: this name is listed as synonymous
with C. tschonoskii in the Flora of China (Li & Skvortsov 1999), in the Flora Sichuanica (Editorial
Board of the Flora Sichuanica 2012), and by Chang et al. (2014a) for the Flora of Korea, while it is
synonymous with C. paohsingensis in the World Checklist (Govaerts & Frodin 1998).
29. Carpinus polyneura Franch.
Journal de Botanique [Morot] 13: 202 (Franchet 1899). – C. turczaninovii var. polyneura (Franch.)
H.J.P.Winkl., Das Planzenreich IV 61 (Heft 19): 38 (Winkler 1904). – Type: China, Sutchuen
[Sichuan], in ditione Tchen keou tin [Chengkou County], P.G. (Reverend Père) Farges s.n. (lecto-,
designated here: P06811144!; isolecto-: P06811145!?, P06811146!). – Note: Carpinus polyneura is
accepted as a species, with the two varieties (with C. polyneura var. sungpanensis) that are listed
here, in the World Checklist (Govaerts & Frodin 1998) and in the Flora Sichuanica (Editorial Board
of the Flora Sichuanica 2012), but they are accepted as two distinct species in the Flora of China
(Li & Skvortsov 1999).
29a. Carpinus polyneura Franch. var. polyneura
C. handelii Rehder, Journal of the Arnold Arboretum 1: 59 (Rehder 1919). – Type: China, Hunan, inter
urbes Linling (Yungchoufu) e Sinning in silvis collium supra vicum Tjentiesse, 400 m a.s.l., 14
Aug. 1917, H.R.E. von Handel-Mazzetti 421 (holo-: A00033754!; iso-: WU 30926!). – Type: China,
Hunan, in silva infra Tungdjiapai prope minas Hsikwangchan, dist. Hsinhwa, 550 m a.s.l., 20 May
1918, H.R.E. von Handel-Mazzetti 11884 [“534”] (para-: E!, P06811414!, WU 30927!). – Note:
this name is listed as a synonym of C. polyneura in the World Checklist (Govaerts & Frodin 1998),
in the Flora of China (Li & Skvortsov 1999), and in the Flora Sichuanica (Editorial Board of the
Flora Sichuanica 2012). Rehder cites the type as “421”, which is a crossed-out number written with
a pencil only on the specimen in A. It is therefore the specimen that Rehder explicitly cited and type,
as the WU duplicate does not contain that number.
C. polyneura var. glandulosopunctata C.J.Qi, Acta Phytotaxonomica Sinica 22 (6): 494 (Qi 1984). –
C. glandulosopunctata (C.J.Qi) C.J.Qi, Bulletin of Botanical Research, Harbin 20 (1): 3 (Lin & Qi
2000) [“glanduloso-punctata”]. – Type: China, Hunan, Ningxiang, on hillside, 200 m a.s.l., 10 Oct.
1982, Y.T. Xiao [Xiao Yu-tan] 40004 (holo-: CSFI; iso-: PE). – Note: this name is listed as a synonym
of C. polyneura in the World Checklist (Govaerts & Frodin 1998), but it is not mentioned otherwise.
C. polyneura var. wilsoniana H.J.P.Winkl., Botanische Jahrbücher für Systematik, Planzengeschichte
und Planzengeographie 50 (Suppl.): 506 (Winkler 1914). – Type: China, [Sichuan,] am Berge
Omi [Éméi Shān], May 1904, E.H. Wilson 5791 (lecto-, designated here: K000859929!; isolecto-:
A00033763!, BM!). – Additional type material: China, Hupeh, E.H. Wilson 1170 p.p. (syn-: W). –
Note: This name is mentioned neither in the World Checklist (Govaerts & Frodin 1998) nor in the
Flora of China (Li & Skvortsov 1999) nor in the Flora Sichuanica (Editorial Board of the Flora
Sichuanica 2012).
29b. Carpinus polyneura var. sungpanensis (W.Y.Hsia) P.C.Li
Flora Reipublicae Popularis Sinicae 21 (2): 86 (Li 1979a). – C. sungpanensis W.Y.Hsia, Contributions
from the Institute of Botany, National Academy of Peiping 2: 180 (Hsia 1934). – C. tschonoskii var.
sungpanensis (W.Y.Hsia) C.C.Yang, Distribution of the Woody Plants in Sichuan [四川树木分布]:
42 (Yang 1997). – Type: China, Sichuan, Sungpan-Hsien, 2050 m a.s.l., 16 Sep. 1932, T.T. Yü 2580
(lecto-, designated here: PE01062157!; isolecto-: IBSC0368164, IBSC0368182, PE01062158!). –
Note: the taxon is accepted as a distinct species in Flora of China (Li & Skvortsov 1999), but treated
23
European Journal of Taxonomy 375: 1–52 (2017)
as a variety of C. polyneura in the World Checklist (Govaerts & Frodin 1998) and in the Flora
Sichuanica (Editorial Board of the Flora Sichuanica 2012).
29c. Carpinus polyneura var. tsunyihensis (Hu) P.C.Li
Flora Reipublicae Popularis Sinicae 21 (2): 88 (Li 1979a). – C. tsunyihensis Hu, Acta Phytotaxonomica
Sinica 9: 296 (Hu 1964). – Type: China, Kweichow [Guizhou], Tsunyih Hsien, Nanchien Shan, on
hilly slope, 900‒1050 m a.s.l., 17 Aug. 1956, Szechuan-Kweichow Exp. 973 (holo-: PE00818388!). –
Note: this taxon is accepted as a distinct species in the Flora of China (Li & Skvortsov 1999), but it
is considered as a variety of C. polyneura in the World Checklist (Govaerts & Frodin 1998) and in
the Flora Sichuanica (Editorial Board of the Flora Sichuanica 2012).
30. Carpinus pubescens Burkill
Journal of the Linnean Society, Botany 26 (178): 502 (Forbes & Hemsley 1899) [“1890”]. – Type:
China, Yunnan, Mi-lê district, A. Henry 9929 (holo-: K000859939!; iso-: A00033764!, E00275493!,
NY00253866!, PE00934232!).
C. seemeniana Diels, Botanische Jahrbücher für Systematik, Planzengeschichte und Planzengeographie
29: 279 (Diels 1901). – C. pubescens var. seemeniana (Diels) Hu, Acta Phytotaxonomica Sinica 9:
290 (Hu 1964). – Type: China, S Nan ch’uan, Shan tzu p’ing, Wald, Aug., C. Bock and A. von
Rosthorn [“BvR”] 294 (lecto-, designated here: GZU-Rosthorn 000261231!; isolecto-: A00033767!,
B†, K000859940!).
C. pinfaensis H.Lév. & Vaniot, Bulletin de la Société botanique de France 52: 142 (Léveillé 1905). – Type:
China, Kouy-tchéou, Pin-fa, 28 Mar. 1903, J. Cavalerie 1011 (holo-: E00275492!; iso-: A00077469!).
C. austrosinensis Hu, Sinensia 2: 87, ig. 4 (Hu 1931). – Type: China, Kweichow [Guizhou], Anlung, in
lightly shaded mixed woods, 620 m a.s.l., 22 Oct. 1930, Y. Tsiang [蒋英] 7470 (holo-: not localized;
iso-: A00033748!, BM000580520!, E00275505!, IBSC0368184, K000859927!, N000053493,
NY00253859!, P06747032!, PE00020176!, PE00020177!, US00089311!). – Note: the type shown
in Hu (1931: ig. 4) is not among the specimens seen.
C. kweitingensis Hu, Sinensia 2: 82, ig. 2 (Hu 1931). – C. pubescens var. kweitingensis (Hu) Hu, Acta
Phytotaxonomica Sinica 9: 290 (Hu 1964). – Type: China, Guizhou, Guiding/Kweiting: Pingfa, light
shaded woods, 400‒550 m a.s.l., 29 Jun. 1930, Y. Tsiang [蒋英] 5440 (holo-: not localized; iso-:
A00077466!, A00077467!, E00275502!, IBSC0001132, M-0265493!, NAS00070311!, P06811206!,
PE00021904!). – Note: the type shown in Hu (1931: ig. 2) is not among the specimens seen.
C. tsiangiana Hu, Sinensia 2: 90, ig. 5 (Hu 1931). – Type: China, Guizhou, Kwangcheng, open hillsides,
23 Jul. 1930, Y. Tsiang [蒋英] 8610 (lecto-, designated here: NAS00070318!; isolecto-: A00033770!,
BM000580521!, E00275491!, K000859920!, NAS00070312!, NAS00070313!, NAS00070314!,
NY00253867!, PE00021964!, US00089309!). – Note: the specimen chosen as lectotype here is the
one depicted in Hu (1931: ig. 5).
C. tungtzeensis Hu, Sinensia 2: 85, ig. 3 (Hu 1931). – Type: China, Guizhou, Tungtze. 450 m a.s.l. (1476.4
ft.), 26 May 1930, Y. Tsiang [蒋英] 5111 (syn-: NAS00070319, NAS00070320, NAS00070321;
isosyn-: BM000580523!, K000859921!, N000053499, NY00253868!, PE00021966!, PE00021967!,
US00089308!).
C. kweiyangensis Hu, Bulletin of the Fan Memorial Institute of Biology n.s. 1 (2): 187 (Hu 1948b). –
Type: China, Kweichow [Guizhou], Kweiyang [贵阳市, Guiyang], Chen-lin Shan, 10 Oct. 1937,
P.C. Tsoong [钟补勤] 1118 (holo-: PE00021948!).
C. lancilimba Hu, Bulletin of the Fan Memorial Institute of Biology n.s. 1 (2): 185 (Hu 1948b). – Type:
China, Kweichow [Guizhou], Tse-ching Hsien [Zhijin Co., 织金县], [Dongshan, 县东山], 28 Oct.
1938, P.C. Tsoong [钟补勤] 1879 (lecto-, designated here: PE00021909!; isolecto-: PE00021910!).
24
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
C. pilosinucula Hu, Bulletin of the Fan Memorial Institute of Biology n.s. 1 (2): 142 (Hu 1948a). – Type:
China, Guizhou, Anlong Co., 15 Jul. 1940, P.C. Tsoong [钟补勤] 1536 (holo-: PE00021943!).
C. pingpienensis Hu, Bulletin of the Fan Memorial Institute of Biology n.s. 1 (2): 188 (Hu 1948b). –
Type: China, Yunnan, Ping-pien Hsien, 1500 m a.s.l., 18 Jul. 1934, H.T. Tsai [Tsai Hsi-Tao, 蔡希陶]
60994 (holo-: PE00021944!; iso-: IBSC0368199, LBG00053976!, WUK0046644).
C. tsoongiana Hu, Bulletin of the Fan Memorial Institute of Biology n.s. 1 (2): 186 (Hu 1948b). – Type:
China, Kweichow [Guizhou], Yungchun Hsien, on rocky slope along river [贵州省, 从江县, Guizhou
Province from River County], 29 Jul. 1937, P.C. Tsoong [钟补勤] 1084 (holo-: PE00021965!).
C. wangii Hu & W.C.Cheng, Bulletin of the Fan Memorial Institute of Biology n.s. 1 (2): 147 (Hu
1948a). – Type: China, Yunnan, Foo-ning, Lung-mai, on rocky hill, 1000 m a.s.l., C.W. Wang [王启
无] 89104 (holo-: not localized; iso-: KUN0590973!).
C. marlipoensis Hu, Bulletin of the Fan Memorial Institute of Biology n.s. 1 (3): 215 (Hu 1949). – Type:
China, Yunnan, Malipo County, Pan-chia-chu, 31 Oct. 1947, K.M. Feng 12619 (lecto-, designated
here: PE00021921!; isolecto-: PE00021922!, WUK0207179).
C. marlipoensis var. angustifolia Hu, Acta Phytotaxonomica Sinica 9: 288 (Hu 1964). – Type: China,
Yunnan, Faadou, 1400 m a.s.l., in mixed forests on rocky hills, 24 May 1943, P.C. Tsoong and K.Z.
Kuang [钟补勤、匡可任] 426 (lecto-, designated here: PE00021925!; isolecto-: IBSC0368204,
PE00021923!, PE00021924!).
C. parva Hu, Acta Phytotaxonomica Sinica 9: 292 (Hu 1964). – Type: China, SE Yunnan: Sichour
Hsien, in shrubbery on cretaceous clif, 1600 m a.s.l., 17 Oct. 1958, H.T. Tsai [Tsai Hsi-Tao, 蔡希
陶] 58–8559 (holo-: PE00021942!).
C. pingpienensis var. lichuanensis W.C.Cheng, 科学 [Science] 32 (8): 246 (Cheng 1950) nom. nud.? –
C. pingpienensis var. lichuanensis W.C.Cheng, 湖北植物志 [Flora Hupehensis] 1: 95 (Hubei Institute
of Botany 1976) nom. nud. – C. pingpienensis var. hupehensis W.C.Cheng nom. nud. – Note: The
name “C. pingpienensis var. hupehensis” only appears on the specimens PE00021911!, A00033762
and K000859922! (China, [Hubei,] Li-Chuan, Hsio-Ho, Long-Tawn-Pu, in forest, 4100 ft, 13
Sep. 1948, W.C. Cheng and C.T. Hwa 950) and appears to be provisional. Cheng annotated the PE
specimen in 1960 as “Carpinus lichuanensis Hu” and as “Carpinus pingpienensis var. lichuanensis”
in 1967. Tang (1987) cited the name C. pingpienensis var. lichuanensis as a “n.n.” (nom. nud.) as
does Chen (2015). The present authors did not ind the protologue for validation. Synonymization
follows Tang (2007) as the name is not mentioned in the Flora of China (Li & Skvortsov 1999).
C. whilungpaensis W.C.Cheng, 科学 [Science] 32 (8): 246 (Cheng 1950) nom. nud.? – Note: Tang
(2007) cited this name as a “n.n.” [nomen nudum?] found on a specimen of W.C. Cheng and C.T.
Hwa 866 (duplicate without that note: K!). The present authors did not ind the protologue, and the
name is not cited in the Flora of China (Li & Skvortsov 1999).
31. Carpinus purpurinervis Hu
Acta Phytotaxonomica Sinica 9: 293 (Hu 1964). – Type: China, Kwangsi [Guangxi], Du-an, in forests,
2 Jul. 1957, Y.K. Li [Li Yin-kuen/李荫昆] P01567 (holo-: PE; iso-: IBK00190887!, IBK00190888!,
IBK00190889!, IBK00190890!, IBK00190891!, IBSC0368220).
32. Carpinus putoensis W.C.Cheng
Contributions from the Biological Laboratory of the Science Society of China, Botanical Series 8: 72
(Cheng 1932). – Type: China, Zhejiang, [普陀区, 佛顶山/Putuo District, Fodingshan], 15 May
1930, K.K. Tsoong [钟观光] 94 (lecto-, designated here: PE00021950!; isolecto-: PE00021949!,
PE00021951!).
25
European Journal of Taxonomy 375: 1–52 (2017)
33. Carpinus rankanensis Hayata
Icones Plantarum Formosanarum 6: 63, ig. 8 (Hayata 1916). – Type: R. China, [Taiwan, Yilan County],
Rankanzan [Lan-K’an Shan, a mountain range NE of Nanao], 4000 ft, 12 May 1916, B. Hayata s.n.
(lecto-, designated here: TI!; isolecto-: TAIF007302!, TAIF007303!, TAIF007304!).
33a. Carpinus rankanensis Hayata var. rankanensis
33b. Carpinus rankanensis var. matsudae Yamam.
Supplementa Iconum Plantarum Formosarum 5: 15, ig. 5 (Yamamoto 1932). – Type: R. China, Taiwan,
Ariko-banti, 8 Jul. 1917, E. Matuda [also called Matsuda] 11 (holo-: TAI118774!; iso-: TI!). – Note:
this variety is accepted as a distinct variety of C. rankanensis in Flora of China (Li & Skvortsov
1999) but not by Liao (1996).
34. Carpinus rupestris A.Camus
Bulletin de la Société botanique de France 76: 966 (Camus 1930). – Type: s. loc., s. coll., eventually
J. Cavalerie 4560 (lecto-, designated here: P01903242!; isolecto-: P01903240!, P01903241!). –
Additional type material: China, Kouy-tchéou [Guizhou], Houang-tia-pa [Houang-ts’ao-pa],
grands rochers, J. Cavalerie 4560 (syn-: A00033766!, E00240621!, K!); ibid., [Guizhou], Hiny-hien [missionary station in SW of Guizhou, NW of Xingyi city, Hsien-i-fu, 興義府], rochers,
J. Cavalerie s.n. (syn-: P01903239!). – Note: the specimen designated here as lectotype bears an
annotation by Camus, so it is original material, but it lacks (as do the isolectotypes) an original label
by Cavalerie. The specimen P01903242 bears a written note on an extracted page with the protologue
that “Cavalerie 4560” is apparently missing in the Paris herbarium. Possibly, the original label
became detached and is lost. The specimens P01903240–2 are not duplicates of the other syntype
P01903239, which has male catkins and narrower leaves. In addition, P01903241 and P01903242
contain infrutescences.
35. Carpinus × schuschaensis H.J.P.Winkl.
Das Planzenreich IV 61 (Heft 19): 32 (Winkler 1904). – C. betulus [unranked] schuschaensis (H.J.P.
Winkl.) Medw., Vestnik Tilisskogo Botanicheskogo Sada 14: 33 (Medwedew 1909). – C. betulus
f. schuschaensis (H.J.P.Winkl.) Radde-Fom., Trudy izychno-matematychnogo viddilu. Kiev
[Mémoires de la Classe des Sciences Physiques et Mathématiques de l’Académie des Sciences de
l’Ukraine] 15 (1): 80 (Radde-Fomina 1929) comb. inval. pro syn. – Type: Karabagh, Schuscha,
R.F. Hohenacker s.n. (syn-: B†, G-Boiss., W). – Note: Radde-Fomina published the combination
C. betulus f. schuschaensis in synonymy, because she believed that Medwedew understood the name
as a forma. However, Medwedew used the term “form” informally, as he used form (Medwedew
1909: 33) and variety (Medwedew 1909: 41) in parallel. His combinations are therefore unranked
(Art. 37.8; McNeill et al. 2012). Biologically, this is a hybrid between C. betulus and C. orientalis.
C. × grosseserrata H.J.P.Winkl., Das Planzenreich IV 61 (Heft 19): 40 (Winkler 1904). – Type: Iran,
bei Radkan, F.A. Buhse 1043 (holo-: B†).
C. × hybrida H.J.P.Winkl., Das Planzenreich IV 61 (Heft 19): 40 (Winkler 1904). – Type: Transkaukasien,
R.F. Hohenacker (syn-: B† and/or LE). Karabagh, s. coll., s.n. (syn-: B† and/or LE).
C. × oxycarpa H.J.P.Winkl., Das Planzenreich IV 61 (Heft 19): 31 (Winkler 1904). – C. betulus [unranked]
oxycarpa (H.J.P. Winkl.) Medw., Vestnik Tilisskogo Botanicheskogo Sada 14: 27 (Medwedew
1909). – C. betulus f. oxycarpa (H.J.P. Winkl.) Radde-Fom., Trudy izychno-matematychnogo
viddilu. Kiev [Mémoires de la Classe des Sciences Physiques et Mathématiques de l’Académie des
Sciences de l’Ukraine] 15 (1): 83 (Radde-Fomina 1929) comb. inval. pro syn. – Type: Karabagh,
26
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
Schuscha, R.F. Hohenacker, U.i. 1838 (syn-: B†, G-Boiss.?, LE, Herb. Moritz Winkler, W†). –
Note: Olga Radde-Fomina’s combination in synonymy is a misunderstanding. She believed that
Medwedew published a forma, but he used the term “form” informally, which is apparent in the
index of his treatment, where he explicitly notes “var. oxycarpa (Winkl.) Medw.” (Medwedew
1909: 41). Medwedew’s combination is nevertheless to be regarded as unranked (Art. 37.8, McNeill
et al. 2012). In contrast to Winkler’s note (Winkler 1904), duplicates are not present in “Breslau”
(Wrocław; BRSL or WRSL, pers. comm. K. Świerkosz).
C. × oxycarpa nothovar. betuloides H.J.P.Winkl., Das Planzenreich IV 61 (Heft 19): 32 (Winkler
1904). – Type: Karabagh, bei Schuscha, R.F. Hohenacker, U.i. 1838 (syn-: B†, G, LE).
C. × geokczaica Radde-Fom., Trudy izychno-matematychnogo viddilu. Kiev [Mémoires de la Classe des
Sciences Physiques et Mathématiques de l’Académie des Sciences de l’Ukraine] 15 (1): 89 (RaddeFomina 1929). – Original citation: “Prov. Baku distr. Geokczai, in frutecetis siccis in ascensu as
pagum Vaenk 1700’. 31 VII. 1899. Alexeenko (Herb. Acad. Petrop.) Prov. Baku distr. Schemacha,
prope pagum Müdshi, in decliviis meridionalibus, in papidosis calcareis. 4000’. 30. VII. 1900.
Alexeenko (Herb. Acad. Petrop.)”. – Type: not localized.
36. Carpinus shensiensis Hu
Bulletin of the Fan Memorial Institute of Biology n.s. 1 (2): 145 (Hu 1948a). – Type: China, Shensi
[Shaanxi], Yuan Kwan T’ang, mixed with Celtis, 15 Oct. 1935, Y.Y. Pai 2891 (lecto-, designated
here: PE00021952!; isolecto-: NAS00070315, WUK0072726). – Additional type material: China,
Shaanxi, Hsing Lung Sze, 15 Oct. 1935, Y.Y. Pai 2860 (para-: PE00802529!, WUK0072723,
WUK0170781).
C. shensiensis var. paucineura S.C.Qu & K.Y.Wang, Bulletin of Botanical Research, Harbin 8 (4): 111
(Qu & Wang 1988). – Type: China, Shaanxi, Zhenping Xian, Feihe Xiang, Laoyachung, 1800 m
a.s.l., Aug. 1986, K.Y. Wang 747 (holo-: NWFC). – Additional type material: ibid., 1200 m a.s.l., K.Y.
Wang 740 (para-: not localized).
37. Carpinus shimenensis C.J.Qi
Journal of Nanjing Technological College of Forest Products 1981 (3): 123 (Qi 1981). – Type: China,
Hunan, Shimen district, Huping shan, 2050 m a.s.l., P.C. Cai [Cai Pingcheng] 20241 (holo-:
CSFI017382; iso-: NF). – Note: this name is accepted in the World Checklist (Govaerts & Frodin
1998), but it is not mentioned in the Flora of China (Li & Skvortsov 1999).
38. Carpinus tientaiensis W.C.Cheng
Contributions from the Biological Laboratory of the Science Society of China, Botanical Series 8: 135
(Cheng 1932). – C. laxilora var. tientaiensis (W.C.Cheng) Hu, Sunyatsenia 1: 112 (Hu 1933). –
Type: China, Zhejiang, Tiantai, Tien Tai Shan, open woods, 2800 ft, 12 Aug. 1927, Y.L. Keng [耿
以礼] 1065 (lecto-, designated by Lin & Sun (2007: 178): PE00021959!; isolecto-: A00033765!,
PE00021935!). – Additional type material: China, Zhejiang, Tiantai, Tientai Shan, in wood partially
shaded, 3500 ft, 10 May 1924, R.C. Ching 1547 (syn-: PE00021958!). – Note: the isolectotype from
PE lacks a specimen label, but it has a collection label that matches the lectotype collection label
exactly. It is therefore very likely a duplicate. However, it is annotated as “Carpinus nobilis Hu
type”, a name whose publication status is unknown to the present authors.
39. Carpinus tropicalis (Donn.Sm.) Lundell
Lloydia 2: 79 (Lundell 1939). – C. americana var. tropicalis Donn.Sm., Botanical Gazette 15: 28 (Smith
1890). – C. caroliniana var. tropicalis (Donn.Sm.) Standl., Contributions from the United States
27
European Journal of Taxonomy 375: 1–52 (2017)
National Herbarium 23 (1): 169 (Standley 1920). – Type: Guatemala, Alta Verapaz, Chicoyonito,
4300 ft, Apr. 1889, J. Donnell Smith in Donn. Sm. Exs. Pl. cit. 1667 (lecto-, designated by Furlow
(1987: 431): US00089312!; isolecto-: GH00033747!, K000512590!). – Additional type material:
Guatemala, Alta Verapaz, bei Coban Tactic Purulá, 1250‒1500 m a.s.l., May 1882, F.C. Lehmann Pl.
Guatemal., Costaric. Columbianae 1446 (syn-: BM000956101!, K!); ibid., Cumbre de Sta. Lucia,
Jan. 1878, K.G. Bernoulli and R. Cario Fl. Guatemal. 2606 (syn-: GOET!, K!); ibid., Las Majadas,
May 1878, K.G. Bernoulli and R. Cario Fl. Guatemal. 2607 (syn-: GOET!, K!).
39a. Carpinus tropicalis (Donn.Sm.) Lundell var. tropicalis
39b. Carpinus tropicalis var. mexicana Furlow
Systematic Botany 12 (3): 431 (Furlow 1987). – Type: Mexico [México], [Jalisco], Municipio de Gómez
Farias, Tamaulipas, camino al Rancho del Cielo, 1150 m a.s.l., 8 Aug. 1969, H. Puig 5159 (holo-:
ENCB). – Additional type material: Mexico [México], Cerro de Corona, Zacualpan, 2‒3 May 1954,
E. Matuda et al. 30712 (para-: BH); ibid., Municipio de Ocuilan, cerca del Rio Chalma, Ahuehuete,
19 Sep. 1962, May Nah AM-48 (para-: ENCB, MICH); ibid., 5 km al SW de Sultipec, sobre el
camino a Amatepec, 2350 m a.s.l., 18 Feb. 1979, J. Rzedowski 36059 (para-: ENCB, MICH); ibid.,
[Michoacán], Municipio de Pueblo Nuevo, ladera S del Volcán Tancitaro, 2000 m a.s.l., 19 Mar.
1977, L.M.V. de Puga and S. Carvajal H. 10134 (para-: ENCB); ibid., Nuevo León, Dulces Nombres,
13 Jul. 1948, F.G. Meyer and D.J. Rogers 2752 (para-: BM000956117!, E!, GH); ibid., Nuevo León,
Municipio de Montemorelos, trail from La Trinidad to Sierra de la Cebolla, 20 Aug. 1939, C.H.
Muller 2867 (para-: GH).
40. Carpinus tsaiana Hu
Bulletin of the Fan Memorial Institute of Biology n.s. 1 (2): 141 (Hu 1948a). – Type: China, Yunnan,
Ping-pien Hsien [屏边苗族自治县, 水塘坡, Pingbian Miao Autonomous County], 7 Jul. 1934,
H.T. Tsai 62398 (lecto-, designated here: PE00021960!; isolecto-: PE00021961!, PE00021962!). –
Additional type material: ibid., 3 Jun. 1934, H.T. Tsai 62191 (para-: LBG00053971, PE00802549!);
ibid., Yunnan, Si-chou Hsien, Far-doe [西畴县, 法斗], 12 Dec. 1939, C.W. Wang [王启无] and Y. Li
[刘瑛] 85686 (para-: PE00021918!, PE00021919!, PE00021920!, PE00802552!) [partly misspelled
as 2 Dec. and 65686 but all with same label data]. – Note: Carpinus tsaiana Hu is accepted in the
Flora of China (Li & Skvortsov 1999). The lectotype designated here is not the unpublished one,
labeled as holotype. We deviate from the original intention, because the intended “holotype” has lost
the fruits of its infructescence, therefore we choose a more complete specimen.
C. sichourensis Hu, Bulletin of the Fan Memorial Institute of Biology n.s. 1 (3): 214 (Hu 1949). – Type:
China, Yunnan, Xichou [西畴县], Ma-chia, 14 Oct. 1947, K.M. Feng 12516 (holo-: PE00802497!).
41. Carpinus tschonoskii Maxim.
Bulletin de l’Academie imperiale des sciences de St.-Pétersbourg 27: 534 (Maximowicz 1881b). – Type:
Japan, [Kanagawa], Hakone, 1864, S. Tschonoski s.n. (syn-: LE; isosyn-: BM!, BR!, GH00033769
p.p.!, K000859942!, L! M-0265492!, P06809892!, P06811215!, P06809882!?, P06811396!?,
P06809866!?, PE00021963 p.p.!, WU!); ibid., Fudzi-yama [Fuji yama], 1864, S. Tschonoski s.n.
(syn-: LE; isosyn-: BM000580518!, GH00033769 p.p.!, K000859941!, P06811217!, PE00021963
p.p.!).
C. yedoensis Maxim., Bulletin de l’Academie imperiale des sciences de St.-Pétersbourg 27: 535
(Maximowicz 1881b). – Type: Japan, [Tokio,] cult. in Yedo [Edo, Tokio] Garden, Nov. 1862, C.J.
Maximowicz s.n. (syn-: not localized); ibid., L. Savatier 1172 (syn-: not localized).
28
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
C. fauriei Nakai, The Botanical Magazine [Tokyo] 26: 325 (Nakai 1912). – Type: South Korea, [Jeju],
Quelpaert, in silvis, Oct. 1907, E.-J. Taquet 587 (syn-: TI, KYO); ibid., Aug. 1911, T. Mori 32 (syn-:
TI); ibid., Aug. 1911, T. Mori 33 (syn-: TI).
C. eximia Nakai, Repertorium Specierum Novarum Regni Vegetabilis 13: 249 (Nakai 1914). –
C. tschonoskii var. eximia (Nakai) Hatus., Bulletin of the Kyushu Imperial University Forests 5:
48 (Hatushima 1934). – Type: South Korea, [Jeollanam-do], secus vias et in silvis circa templum
Senonji pede montem Chirisan, 280 m a.s.l., 15 Jul. 1913, T. Nakai 11 (holo-: TI). – Note: this name
is accepted as a distinct species in the World Checklist (Govaerts & Frodin 1998), but listed as a
synonym of C. tschonoskii in Chang et al. (2014a) and the WCSP (2016).
C. tschonoskii var. jablonskyi H.J.P.Winkl., Botanische Jahrbücher für Systematik, Planzengeschichte
und Planzengeographie 50 (Suppl.): 500 (Winkler 1914). – Type: Japan, prov. Kotzuke, Bamba
und Nogurizawa, 29 Jul. 1894, M. Shirai s.n. (syn-: not localized); ibid., 1 Aug. 1894, M. Shirai s.n.
(syn-: not localized). – Note: this name is mentioned neither in the World Checklist (Govaerts &
Frodin 1998) nor in the WCSP (2016) nor in the Flora of Japan (Ohwi 1965). Chang et al. (2014a)
list it as synonymous with C. tschonoskii.
C. tschonoskii var. serratiauriculata H.J.P.Winkl., Botanische Jahrbücher für Systematik,
Planzengeschichte und Planzengeographie 50 (Suppl.): 500 (Winkler 1914). – Type: Japan,
[Tokyo,] Jedo, L. Savatier 1172 (syn-: not localized); ibid., L. Savatier s.n. (syn-: not localized).
Ibid., 1874, L. Savatier s.n. (syn-: P06811219!, P06810761!). – Note: this name is mentioned neither
in the World Checklist (Govaerts & Frodin 1998) nor in the WCSP (2016) nor in the Flora of Japan
(Ohwi 1965). Chang et al. (2014a) list it as synonymous with C. tschonoskii.
C. tschonoskii var. subintegra H.J.P.Winkl., Botanische Jahrbücher für Systematik, Planzengeschichte
und Planzengeographie 50 (Suppl.): 501 (Winkler 1914). – Type: South Korea, [Jeju], Insel Quelpart,
Hallaisan [Hallasan mt.], Jul. 1907, U.J. Faurie 1535 (holo-: not localized; iso-: A00033735!, BM!,
E00388827!, P06809858!). – Note: this name is not mentioned in the World Checklist (Govaerts &
Frodin 1998), while Chang et al. (2014a) list it as synonymous with C. tschonoskii.
C. tschonoskii var. brevicalycina Nakai ex Kawamoto, Handbook of Korea-Manchurian Forestry: 92
(Kawamoto 1939). – Type: not localized. – Note: this name is not mentioned in the World Checklist
(Govaerts & Frodin 1998) nor in the WCSP (2016). Jeon & Chang (1997) consider it as a synonymous
with C. tschonoskii, Chang et al. (2014a) list it as a invalid name. The present authors did not see the
protologue, so validity of the name is unclear.
C. coreensis Koidz., Acta Phytotaxonomica et Geobotanica 9 (2): 73 (Koidzumi 1940). – Original
citation: “Carpinus fargesiana (non Winkl.) Nakai Fl. Sylv. Korea. II (1915). […] Korea: mt. Chiisan,
Hannyaho, Kokiri”. – Type: not localized. – Note: Jeon & Chang (1997) and Chang et al. (2014a)
consider this name to be synonymous with C. tschonoskii, as it is listed in the World Checklist
(Govaerts & Frodin 1998).
C. tschonoskii f. pendula Hayashi, 林業試験場研究報告 [Research Report of the Forestry Experiment
Station] 125: 72 (Hayashi 1960). – Type: Japan, Hondo, prov. Mushi, Kokubunji, 10 May 1958, Y.
Hayashi s.n. (holo-: TFM). – Note: this name is not mentioned in the World Checklist (Govaerts &
Frodin 1998) or in the Flora of Japan (Ohwi 1965). Most likely, this is to be regarded as a cultivar
with hanging branches rather than as a botanical taxon.
C. mianningensis T.P.Yi, Bulletin of Botanical Research, Harbin 12 (4): 335 (Yi 1992). – Type: China,
Sichuan, Mianning County, Lingshan, 18 Jul. 1991, T.P. Yi [易同培] 91170 (holo-: SIFS; iso-:
PE00934231!). – Note: this name is accepted in the Flora Sichuanica (Editorial Board of the Flora
Sichuanica 2012), but it is listed as synonymous with C. tschonoskii in the Flora of China (Li &
Skvortsov 1999).
29
European Journal of Taxonomy 375: 1–52 (2017)
41. Carpinus turczaninovii Hance
Journal of the Linnean Society, Botany 10 (44): 203 (Hance 1868). – Type: China, in collibus
ad occidentem urbis Peking [Beijing] sitis, Aug. 1865, S.W. Williams 12681 (holo-:
K000859937!). – Additional original material: China, in collinis Pekinensibus, Aug. 1865, S.W.
Williams 12681 (BM000580390!). – Additional original material?: China, in ditione Pekinensis,
Aug. 1866, S.W. Williams 12681 (GH00112537!, LE). – Note: several gatherings are distributed
with the number 12681, all belonging to the same species. Winkler (1914: 502) already mentions
the admixed nature of the specimens with the same “collection” number. Hance cited “collibus ad
occidentem urbis Peking, m. Augusto 1865,” which is only written on the specimen K000859937,
which is therefore the holotype. However, the other specimens seen by the present authors were
distributed from Hance’s collection, so they are original material.
C. paxii H.J.P.Winkl., Das Planzenreich IV 61 (Heft 19): 35 (Winkler 1904). – Type: China, Peking, Tsetai-ssu, H. Wawra von Fernsee 1067 (syn-: B†, W†; isosyn-: A00033761!). – Note: Winkler (1914:
502) acknowledged C.K. Schneider’s comment that his C. paxii is synonymous to C. turczaninovii.
This name is not mentioned in the Flora of China (Li & Skvortsov 1999), but it is listed as synonym
of C. turczaninovii in the Flora of Japan (Ohwi 1965) and in the Flora Sichuanica (Editorial Board
of the Flora Sichuanica 2012).
C. stipulata H.J.P.Winkl., Das Planzenreich IV 61 (Heft 19): 35 (Winkler 1904). – C. turczaninovii var.
stipulata (H.J.P.Winkl.) H.J.P.Winkl., Botanische Jahrbücher für Systematik, Planzengeschichte und
Planzengeographie 50 (Suppl.): 505 (Winkler 1914). – Type: China, Schensi [Shaanxi], C-China,
im Tsin-ling-shan-Gebirge, 10 Jul. 1900, G. [Giuseppe, but also called Joseph; Reverend] Giraldi
7267 (lecto-, designated here: K000859930!; isolecto-: A00077468!, B†). – Note: this name is listed
as a synonym of C. turczaninovii in the Flora of Japan (Ohwi 1965), but it is accepted as a distinct
species in the Flora of China (Li & Skvortsov 1999).
C. tanakaeana Makino, The Botanical Magazine [Tokyo] 28: 32 (Makino 1914). – Type: Japan,
Tokushima, prov. Tosa, Mt. Yokogura (on limestone), T. Makino s.n. (holo-: MAK046499!).
C. turczaninovii var. makinoi H.J.P.Winkl., Botanische Jahrbücher für Systematik, Planzengeschichte
und Planzengeographie 50 (Suppl.): 505 (Winkler 1914). – Type: Japan, prov. Tosa, 1889, T.
Makino 277 (syn-: not localized); ibid., s. loc., M. Shirai s.n. (syn-: not localized). – Note: this name
is not listed in the World Checklist (Govaerts & Frodin 1998) or in the WCSP (2016), but listed as a
synonym of C. turczaninovii in the Flora of Japan (Ohwi 1965).
C. coreana Nakai, The Botanical Magazine [Tokyo] 40: 162 (Nakai 1926). – C. turczaninovii var. coreana
(Nakai) W.T.Lee, Lineamenta Florae Koreae 166 (Lee 1996). – Type: South Korea, [Jeollanam-do],
in monte Kannonzan insl. Wangto, 20 Jun. 1913, T. Nakai 581 (holo-: TI). – Additional type material:
South Korea, [Incheon], prov. Keiki, Chemulpo [Jemulpo], in horto residentis germanici, Sep.
1906, U.J. Faurie 202 [cited as E. Taquet 202] (para-: BM000580428, E00275514!, P05591545!);
ibid., [Jeollanam-do,], prov. Zennan, monte Yudalsan [mt.], Mokpo [city near Yudalsan], T. Nakai
9377 (para-: not localized); ibid., [Jeollanam-do], in silvis Genkeimen, T. Nakai 9378 (para-:
not localized); ibid., [Jeollanam-do], in monte insl. Chito, T. Nakai 9379 (para-: not localized);
ibid., [Jeollanam-do], in silvis montis Semsatsuzan insl. Chinto [Jindo], T. Nakai 9381 (para-: not
localized); ibid., [Jeollanam-do], in silvis insl. Baikato, T. Ishidoya and Chung 3382 (para-: not
localized); ibid., [Jeollanam-do], in monte Monganzan insl. Daikokuzanto, T. Ishidoya and Chung
3384 (para-: not localized); ibid., [Jeju,] Quelpaert [Jeju-do]: in silvis, E. Taquet 6220 (para-: not
localized); ibid., [Jeollanam-do], Kantoku, T. Nakai 1439 (para-: not localized); Japan, [Nagasaki],
Tsusima [Tsushima] in monte Sumo-shiratake, T. Nakai s.n. (para-: not localized). – Note: Carpinus
coreana is cited as a distinct species in the World Checklist (Govaerts & Frodin 1998), but listed
as a synonym of C. turczaninovii in WCSP (2016). Nakai described his new species by using the
diferences of some Korean specimens that had been discussed by Winkler (1914) as being similar
to C. turczaninovii, but not typical. Sun et al. (2011) used three accessions in their phylogeny and
30
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
found one to be almost inseparable from C. turczaninovii [Lee s.n. (SKK); Yoo & Wen (2002)],
and two being closely related to C. fangiana and C. japonica. Although the three accessions could
not be examined, it is likely that the irst one concerns C. coreana. For the two other accessions no
specimen is cited, but they were discussed by Sun et al. (2011) as difering by “long hairs on the
mid vein, inlorescences, and leaf margin veins”. The placement in the phylogeny, however, rather
suggests that those specimens belong to the Distegocarpus group, and hence might be misidentiied
C. japonica or C. cordata.
C. coreana var. major Nakai, The Botanical Magazine [Tokyo] 40: 162 (Nakai 1926). – Type: South
Korea, [Hwanghae-namdo], Yooshin, prov. Kokei, T.H. Chung [Chung-tyai-hyon] s.n. (holo-: TI). –
Note: according to the diagnosis, the large leaf and bract size rather suggest stronger ainities to
C. tschonoskii than to C. turczaninovii, however this remains speculative without having examined
the type. Jeon & Chang (1997), Chang et al. (2014a) and the WCSP (2016) consider this to be a
synonym of C. turczaninovii.
C. chowii Hu, Journal of the Arnold Arboretum 13: 333 (Hu 1932). – Type: China, Hopei, Fang-shan,
500 m a.s.l., 5 Oct. 1931, H.F. Chow 41730 (lecto-, designated here: PE00021883!; isolecto-:
A00033750!, PE00021884!, PE00802985!).
C. coreana var. multilora Nakai ex Kawamoto, Handbook of Korea-Manchurian Forestry: 92
(Kawamoto 1939). – Type: not localized. – Note: this name is neither mentioned in the World Checklist
(Govaerts & Frodin 1998) nor in the Flora of China (Li & Skvortsov 1999) or the WCSP (2016).
Jeon & Chang (1997) consider is to be synonymous with C. turczaninovii, Chang et al. (2014a) list
it as invalid name. The present authors did not have the protologue at hand for veriication.
C. turczaninovii var. arguta Uyeki, Woody-Plants and their Distributions in Tŷsen: 16 (Uyeki 1940). –
Type: not localized. – Note: this name is not mentioned in the World Checklist (Govaerts & Frodin
1998) or the WCSP (2016), but Chang et al. (2014a) list it as a invalid name. The present authors did
not see the protologue to conirm this.
C. turczaninovii var. chungnanensis P.C.Kuo, Flora Tsinlingensis 1 (2): 601 (Instituto Botanico
Boreali-occidentali Academiae Sinicae Edita 1974). – Type: China, Shaanxi, Chang-an Hsien
[终南山, Chung-nan-shan; Zhongnanshan], 1700 m a.s.l., 15 Jun. 1933, H.W. Kung 2838 (holo-:
WUK0000756, WUK0000764). – Note: if necessary, the lectotype should be chosen from one of
the two cited specimens, because the Flora Tsinlingensis is mainly based on material from WUK.
As there is no image of the specimens available to the present authors, the designation of a lectotype
here does not seem to be appropriate.
43. Carpinus viminea Lindl.
Plantae Asiaticae Rariores 2: 4 (Wallich 1831). – Type: India, in Kamaon, R. Blinkworth in N. Wallich
list no. 2800b (lecto-, designated here: K001117116!; isolecto-: BM000580526!). – Additional type
material: Nepal, in montibus Napaliae, N. Wallich list no. 2800a (syn-: BM000580527!, E00301177!,
E00301178!, K001117115!, K000859917!, K000859918!, M!); India, Sirmore [Sirmaur], W.A.
Webb s.n. (syn-: not localized). – Note: it is accepted in Flora of China (Li & Skvortsov 1999) with
C. viminea var. chiukiangensis as a distinct variety. The Kew specimen K000859918 bears a note
“Napalia Wallich 1830” but Wallich was in Nepal in 1821 (Wallich list no. 2800a). Most likely, the
number has to be regarded as the year when the specimen was received at Kew.
C. laxilora var. macrostachya Oliv., Icones Plantarum 20 (4): tab. 1989 (Oliver 1891). – C. macrostachya
(Oliv.) Koidz., Acta Phytotaxonomica et Geobotanica 9 (2): 73 (Koidzumi 1940). – Type: China,
Hupeh, North Patung, A. Henry 7013 (lecto-, designated here: K000859943!; isolecto-: BM!, E!,
P06811517!). – Note: this name is treated as a synonym of C. viminea var. viminea in the Flora of
China (Li & Skvortsov 1999) and the Flora Sichuanica (Editorial Board of the Flora Sichuanica
2012).
31
European Journal of Taxonomy 375: 1–52 (2017)
C. laxilora var. davidii Franch., Journal de Botanique [Morot] 13: 203 (Franchet 1899). – C. davidii
(Franch.) C.K.Schneid., Illustriertes Handbuch der Laubholzkunde 2: 893 (Schneider 1912). –
Type: China, China centralis, prov. Kiukiang, in montibus Ly chan, Sep. 1868 [Reverend Père,
J.P.] A. David 750 (lecto-, designated here: P06811502!; isolecto-: P06811505!, P06811506!,
P06811507!). – Note: this name is treated as a synonym of C. viminea var. viminea in the Flora of
China (Li & Skvortsov 1999) and the Flora Sichuanica (Editorial Board of the Flora Sichuanica
2012).
C. fargesii Franch., Journal de Botanique [Morot] 13: 202 (Franchet 1899). – C. laxilora var. fargesii
(Franch.) Burkill, Journal of the Linnean Society, Botany 26 (178): 501 (Forbes & Hemsley 1899). –
Type: China, Sutchuen [Sichuan], in ditione Tchen keou tin [Chengkou County], P.G. [Reverend
Père] Farges 699 (holo-: P06811515!; iso-: NY00253861!). – Note: this name is treated as a
synonym of C. viminea var. viminea in the Flora of China (Li & Skvortsov 1999) and the Flora
Sichuanica (Editorial Board of the Flora Sichuanica 2012).
C. kenpukwan Koidz., Acta Phytotaxonomica et Geobotanica 9 (2): 74 (Koidzumi 1940). – Type:
[China, China centralis] Hu & Chun (1927: 15, tab. 15) [tabula, sub C. laxilora (Siebold & Zucc.)
Blume]. – Note: this name is mentioned neither in the Flora of China (Li & Skvortsov 1999) nor in
the Flora Sichuanica (Editorial Board of the Flora Sichuanica 2012).
C. tehchingensis Hu, Acta Phytotaxonomica Sinica 9: 283 (Hu 1964). – Type: China, Yunnan, Tehchin
Hsien, Sila, in mixed woods, 2700‒2900 m a.s.l., 18 Jul. 1944, K.M. Feng [冯国楣] 5593 (holo-:
PE00021957!). – Note: this name is also synonymous with C. viminea var. viminea according to
the Flora of China (Li & Skvortsov 1999) and the Flora Sichuanica (Editorial Board of the Flora
Sichuanica 2012).
C. viminea var. chiukiangensis Hu, Acta Phytotaxonomica Sinica 9: 282 (Hu 1964). – Type: China,
Yunnan, Chiukiang valley, in wood, 1950 m a.s.l., 31 Jul. 1938, T.T. Yü 19531 (lecto-, designated
here: PE00802369!; isolecto-: A00033771!, E!, PE00802338!, PE00802368!). – Note: this name is
accepted as distinct variety of C. viminea in Flora of China (Li & Skvortsov 1999).
C. fargesii var. latifolia S.Y.Wang & C.L.Chang, Journal of the Henan Agricultural College 14 (2): 6
(Ding et al. 1980). – Type: China, Henan, Luanchuan (栾川), 12 Aug. 1978, S.Y. Wang et al. [王遂
义等] 78 0346 (holo-: HEAC).
Invalid names
C. betulus var. marmoratus de Vos, Beredeneerd woordenboek der voornaamste heesters en coniferen, in
Nederland gekweekt: 21 (Vos 1867) nom. nud. – Note: the name by de Vos lacks a valid description,
so it is to be ignored under the ICN rules (McNeill et al. 2012) but available as a cultivar name
(Art. 21.6. ICNCB, Brickell et al. 2009): C. betulus ‘Marmorata’. According to Krüssmann (1951),
the name was also published in a Späth garden catalogue, but these were not at hand to the present
authors for the veriication of the validity.
C. albopilosa Hu nom. nud. – Note: no publication of this name is known, name only written on the
specimen PE00020172! (China, Sichuan, Leibo County, 1934, W.Y. Chun 3942).
C. calcarea W.C.Cheng nom. nud. – Note: this name is only written on a specimen (China, Metasequoia
Area, Hubei-Sichuan, 1948 m a.s.l., W.C. Cheng and C.T. Hwa 866: A00033749!, K!). A duplicate in
HIB (HIB0156026) is stored as Ostrya japonica. The specimen was also cited under the apparently
invalid name C. whilungpaensis W.C.Cheng.
C. cambodica L.Pierre nom. nud. – Note: apparently an unpublished name. The specimens are stored in
P herbarium, P06811149!, P06811167! and P06810780! (Vietnam, crescit ad montem Chiao shan in
Cochinchina, Feb. 1877, L. Pierre 3300), under C. poilanei A.Camus (synonymous to C. londoniana
H.J.P.Winkl. var. londoniana).
32
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
C. changhwaensis Hu nom. nud. – Note: no publication of this name known, only written on the specimen
PE00020179! (China, Zhejiang, Linan, 8 May 1929, s. coll. 363).
C. japonica var. auricula Maxim. nom. nud. – Note: this name is only listed on written specimens
BM000580481!, BR! and K! (Japan, Yokohama, 1862, C.J. Maximowicz s.n.).
C. laifengensis Wuzhi nom. nud. – Note: apparently an unpublished herbarium name, mentioned in
Chen (2015).
C. mexicana Griseb. nom. nud. – Note: this name is only written on a specimen in GOET! (Mexico, W.
Schafner Pl. Mexicanae no. 14) and on two specimens in P, P06810806! (Mexico, W. Schafner Pl.
Mexicanae no. 14) and P06810805! (Mexico, in montanis mexique Orizaba, M.B. [M. Botteri?]).
Despite having the same name, but at a diferent rank, Furlow (1987) does not mention the apparently
unpublished Grisebach name.
C. microbracteata Hu nom. nud. – Note: there is no publication with this name known to the present
authors; it is only written on the specimens BM! and PE00818273! (China, Sichuan, Baoxing,
1900 m a.s.l., 11 Sep. 1936, K.L. Chu [曲桂龄] 3824).
C. tatarinowii Hance nom. nud. – Note: there is no publication with this name known to the present
authors; it is only written on the specimens P06811225! and P06811224! (China, ad Pekinum
[giulles], P.G. [Reverend Père] Farges s.n.).
Checklist of Ostrya
Order Magnoliales Juss. ex Bercht. & J.Presl
Family Betulaceae Gray
Subfamily Coryloideae Hook.f.
Tribe Carpineae A.DC.
Ostrya Scop.
Flora Carniolica: 414 (Scopoli 1760) nom. cons. – Type: Ostrya carpinifolia Scop. – Note: the name was
conserved against Ostrya J.Hill (Hill 1757: 513) [“1756”], which is a publication of a superluous
name for Carpinus L.
1. Ostrya carpinifolia Scop.
Flora Carniolica ed. 2, 2: 244 (Scopoli 1772). – O. carpinifolia var. genuina Fliche, Bulletin de la
Société botanique de France 35: 166 (Fliche 1888) nom. inval. (although not explicitly cited but
in context including the type, see Art. 24.3, McNeill et al. 2012). – O. italica subsp. carpinifolia
(Scop.) H.J.P.Winkl., Das Planzenreich IV 61 (Heft 19): 22 (Winkler 1904). – Carpinus ostrya L.,
Species Plantarum: 998 (Linnaeus 1753) p.p. – Original citation: “Italia, Virginia.” – Type: s. loc.,
s. coll. (lecto-, designated by Iamonico & Reveal (2012: 866): LINN! ex herb. Linn. no. 1131.4). –
Additional original material: S.d. (BM000647418! ex herb. Cliford 447, Carpinus 2).
O. vulgaris Willd., Species Plantarum ed. 4, 4 (1): 469 (Willdenow 1805) nom. illegit. superl. – Note:
Willdenow cites O. carpinifolia as synonym, so his name is superluous.
O. italica P.Micheli ex Spach, Annales des sciences naturelles, Botanique ser. 2 16: 246 (Spach
1841b) nom. illegit. superl. – O. italica Scop., Flora Carniolica: 414 (Scopoli 1760) nom. inval.
(polynomen), pro syn. – Carpinus italica Steud., Nomenclator Botanicus ed. 2, 1: 300 (Steudel
1841) nom. inval. pro syn. – O. ostrya var. italica (P.Micheli ex Spach) C.K.Schneid., Illustriertes
33
European Journal of Taxonomy 375: 1–52 (2017)
Handbuch der Laubholzkunde 1: 143 (Schneider 1904) nom. inval. (tautonym). – Note: Ostrya
italica Scop. is rather a misinterpretation as Scopoli discusses only genera and as he did not adopt
Linnaeus’ binomials in this work. Scopoli’s name might better be read as polynomial “Ostrya Italica,
Carpini folio liongiore &? breviore.” that he took from Micheli (1729: 223) as one of the names used
for the genus Ostrya. Spach accepted O. italica in 1841, but he cited O. carpinifolia in synonymy.
One year later, Spach refers to the species as O. vulgaris Willd. Steudel refers to an unpublished
name, or used by Scopoli, “Carpinus italica” (or mistaken by Steudel) as a synonym of O. vulgaris.
O. cylindrica Friv., A’Magyar Tudós Társaság Évkönyvei 3 (3): 163 (Frivaldszky von Frivald 1837)
nom. nud. – Note: the name also appears on a specimen (WU!) from Macedonia.
O. carpinifolia var. alpestris Ces., Saggio su la Geograia botanica e su la Flora della Lombardia: 61
(Cesati 1844). – Type: not localized (RO?). – Note: There is a specimen by Cesati from Varenna,
Lombardy, Italy (P06751764!). It was annotated by him as “Ostrya carpinifolia β collina mihi”.
Eventually, Cesati changed his mind about the epithet upon publication of his new variety.
O. ladelcii Sanguin., Flora Romanae Prodromus Alter: 818 (Sanguinetti 1864). – Original citation: “In
sylvaticis Latii, Frascati, Albano.” – Type: not localized (RO?).
O. carpinifolia var. corsica Fliche, Bulletin de la Société botanique de France 35: 166 (Fliche 1888). –
Type: France, Corsica, forêt communale de Pietrosa, May 1885, P. Fliche s.n. (syn-: not localized).
France, Corsica, forêt de Tova, May 1885, P. Fliche s.n. (syn-: not localized); ibid., forêt de Bavella,
Mathieu s.n. (syn-: not localized); ibid., plateau de Nebbio, Mathieu s.n. (syn-: not localized).
O. carpinifolia var. lanceolata Kem.-Nath., Trudy Tbilisskogo Botaniceskogo Instituta 2: 131 (KemulariaNathadze 1937). – Type: Georgia, prov. Kutais, distr. Letschchum, fauces l. Ladjanura loc. Orpiri,
prope pag. Atscharis-chidi ad rupes calcareas, 22 May 1929, L.M. Kemularia-Nathadze s.n. (syn-:
not localized); ibid., 19 Oct. 1932, L.M. Kemularia-Nathadze s.n. (syn-: not localized).
2. Ostrya chinensis I.M.Turner
Annales Botanici Fennici 51: 308 (Turner 2014). – O. multinervis Rehder, Journal of the Arnold
Arboretum 19: 71, pl. 217 (1938) nom. illegit., non Ettingshausen (1868) (later homonym). – Type:
China, Hunan, Ma-ling-tung, Sinning Hsien, in mixed forest on slope, 650 m a.s.l., 13 Oct. 1935, C.S.
Fan and Y.Y. Li 605 (holo-: A00033789!; iso-: BM001191567!, NAS00286610!, L!, P06811043!).
3. Ostrya japonica Sarg.
The Silva of North America 9: 32 (Sargent 1896). – O. ostrya var. japonica (Sarg.) C.K.Schneid.,
Illustriertes Handbuch der Laubholzkunde 1: 143 (Schneider 1904) nom. inval. (tautonym). – Type:
Japan, Hokkaido, Hill near Sapporo, 18 Sep. 1892, C.S. Sargent s.n. (syn-: GH00055441!); ibid.,
Oiwagi, 1861, C.J. Maximowicz s.n. (syn-: BM!, BR!, K000859914!, L!, M-0265487!, P06809338!,
P06811038!, W1886–1595!); ibid., Nambo, 1865, S. Tschonoski s.n. (syn-: K0000859913!,
P06811037!); ibid., in sylvis frondosis circa lacum Konoma (syn-: not localized); cult. in Arnold
Arboretum from seeds sent from Japan by H. Mayr (syn-: not localized). – Note: the name
O. japonica Sarg. was only accepted by the author in the 1896 publication. Before, Sargent (1893,
1894) initially expressed certainty on the distinctness “In the forests of Yezo I felt no doubt of its
speciic distinctness […] all, things considered, it is, perhaps, best to consider the Japanese tree as
speciically distinct”. However, just in the next sentence, he weakened his opinion stating “Only
after it has been grown here during many years side by side with the American species will it be
possible to reach any opinion on this subject worthy of much consideration. If it proves to be distinct
it should bear the name of Ostrya Japonica”. According to Art. 36.1. b (McNeill et al. 2012), the two
early publications (Sargent 1893, 1894) cannot be considered as a valid publication of O. japonica.
In 1896, however, Sargent accepted the Japanese collections as fully distinct and formalized the
name.
34
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
O. carpinifolia f. japonica Schelle, Handbuch der Laubholzbenennung: 49 (Beissner et al. 1903) nom.
nud.
O. liana Hu, Journal of the Arnold Arboretum 11: 49 (Hu 1930). – Type: China, Chihli, eastern tomb,
15 Sep. 1929, C.F. Li timber no. 3 [ield no. 3, Herbarium number 15] (lecto-, designated here:
PE00021986!; isolecto-: A00033788!).
O. japonica var. homochaeta Honda, The Botanical Magazine [Tokyo] 47: 433 (Honda 1933). – Type:
Japan, Honshu, Nikko, prov. Shimotsuke, 1889, M. Shirai s.n. (holo-: TI). – Additional type material:
ibid., 1931, H. Itoo s.n. (para-: not localized); ibid., prov. Bicchu, Takahashi, 1915, G. Koidzumi s.n.
(para-: not localized); ibid., Hokkaido, in monte Moiwa, circ. Sapporo, 1893 G. Yamada (para-: not
localized); South Korea, [Seogwipo,] in monte Hallaisan, ins. Saishuutoo, 1917, E.J. Taquet 4439
(para-: E00388817); ibid., [Jeju,] ins. Saishuto [Jeju-do, Quelpaert], 1917, T. Nakai 4905 (para-: not
localized); ibid., [Haenam], in monte Daikoji, prov. Zenra-nandoo, 1921, T. Nakai 9384 (para-: not
localized).
O. virginica var. japonica Sarg., Garden and Forest 6: 383 (Sargent 1893) nom. nud. – Note: Sargent
wrote that Maximowicz (1881a) had named the Japanese Ostrya collections as var. japonica, but
there is no such name mentioned in the cited Maximowicz paper.
4. Ostrya knowltonii Coville
Garden and Forest 7: 114 (Coville 1894). – O. carpinifolia f. knowltonii (Coville) Schelle, Handbuch der
Laubholzbenennung: 49 (Beissner et al. 1903). – Type: USA, Arizona, Yavapai Co., Grand Canon of
the Colorado River, 10 Jul. 1892, J.W. Toumey 272 (holo-: US00130392!; iso-: A00263686!).
O. baileyi Rose, Contributions from the United States National Herbarium 8 (4): 293 (Rose 1905). –
Type: USA, Texas, Guadalupe Mountains, 19 Aug. 1901, V. Bailey 435 (holo-: US00089317!).
O. chisosensis Correll, Wrightia 3 (7): 128 (Correll 1965). – O. knowltonii subsp. chisosensis (Correll)
A.E.Murray, Kalmia 12: 22 (Murray 1982). – O. knowltonii var. chisosensis (Correll) A.E.Murray,
Kalmia 13: 10 (Murray 1983). – Ostrya virginiana var. chisosensis (Correll) Henrickson ex
A.M.Powell, Trees and Shrubs of the Trans-Pecos and Adjacent Areas: 75 (Powell 1998). –
Type: USA, Texas, Brewster Co., Chisos Mts, Big Bend NP, base of north-facing ledges, Emory
Peak, 17 Jun. 1964, D.S. Correll 29733 (holo-: LL00370396!; iso-: C10009751!, GH00033786!,
LL00370395!, MEXU00122440!, MEXU00711117!, MICH1192171!, NCU00000363!, TEX,
US00089316!). – Additional type material: USA, Texas, high shady crevice in rock bluf, near
Nail place, Chisos Mts., 21 Aug. 1915, M.S. Young s.n. (para-: SRSC3716!, TEX00378112); ibid.,
north of Crown Mt, Chisos Mts, 1 Jul. 1937, B.H. Warnock 21633 (para-: SRSC3715!, SRSC3721!,
TEX00378110); ibid., igneous soil on Emory-Boot trail, Big Bend NP, Chisos Mts, 7500 ft, 8 Jun.
1952, B.H. Warnock 10478 (para-: LL00378111, SRSC3718!); ibid., along a trail from Boot Spring
and Basin, Chisos Mts, north side of emory Peak, Big Bend NP, 15 Jul. 1955, B.H. Warnock 12726
(para-: LL00378109, SRSC3714!). – Note: Ostrya chisosensis was recognized in Flora of North
America (Furlow 1997). However, its taxonomic status has been challenged recently (McCauley &
Paces 2015).
5. Ostrya rehderiana Chun
Journal of the Arnold Arboretum 8 (1): 29 (Chun 1927). – Type: China, Tien Moh Shan, 400 m a.s.l.,
fairly common in open woods, 2 Oct. 1925, R.C. Ching 3385 (lecto-, designated here: A00033790!;
isolecto-: N, US00089314!).
6. Ostrya trichocarpa D.Fang & Y.S.Wang
Guihaia 3 (3): 189 (Fang & Wang 1983). – Type: China, Guangxi, Napo County (那坡县), 百都公社,
弄化, 1260 m a.s.l., 石山 [rocky hill], 23 Apr. 1981, D. Fang, T.-H. Tan and Z.-G. Wang [方鼎, 覃
35
European Journal of Taxonomy 375: 1–52 (2017)
德海, 王振刚] 22412 (holo-: GXMI050301). – Additional type material: ibid., D. Fang, M.-X. Lai
et al. 25379 (para-: GXMI); ibid., Guanxi Zhuang Auton. Region, Jinxi County, 780 m a.s.l., 17 Nov.
1962, Z.-J. Li and H.-Q Li 1650 (para-: IBK00079836!, IBK00079837!).
7. Ostrya virginiana (Mill.) K.Koch
Dendrologie 2 (2): 6 (Koch 1873). – Carpinus virginiana Mill., The Gardeners Dictionary ed. 8, no.
6 (Miller 1768). – Zugilus virginica Raf., Florula Ludoviciana: 159 (Rainesque 1817) orth. var.,
nom. illegit. superl. – O. ostrya var. virginiana (Mill.) C.K.Schneid., Illustriertes Handbuch der
Laubholzkunde 1: 143 (Schneider 1904) nom. inval. (tautonym). – Type: [Cult. garden specimen],
1740, Chelsea Physic Garden 938 (lecto-, designated by Reveal (1990): BM001041898!). – Note: in
his protologue, Miller only cited a Plukenet (1692) drawing, which does not correspond well to what
is today considered as O. virginiana. Reveal (1990) found a matching typotype (H.S. 95:143, BMSL) that turned out to be O. carpinifolia Scop., and three specimens from that era that Miller might
have had seen (but without actual evidence) of what is now considered O. virginiana. Nevertheless,
the proposal to conserve the name with a new type was rejected, and it was decided that one of the
specimens should serve as lectotype (Brummitt 1993).
7a. Ostrya virginiana (Mill.) K.Koch subsp. virginiana
Carpinus virginica Münchh., Der Hausvater 5 (1): 120 (Münchhausen 1770) orth. var. – O. carpinifolia
var. virginica (Münchh.) Fliche, Bulletin de la Société botanique de France 35: 166 (Fliche 1888). –
Note: Münchhausen misspelled the name given by Miller.
Carpinus trilora Moench, Methodus: 694 (Moench 1794). – Type: not localized. – Note: the protologue
does not state any locality, and Moench’s herbarium is destroyed (Staleu & Cowan 1976–1988).
O. americana F.Michx., Histoire des Arbres Forestiers de l’Amérique Septentrionale 3 (1): 54
(Michaux 1812) nom. inval. – Associated material: A Canada ad Carolinam montosam, s. coll., s.n.
(P06810885!). – Note: while Michaux briely mentions an “Ostrya americana”, he apparently does
not consider it to be a species diferent from “Carpinus ostrya” (Ostrya carpinifolia). The name is
not mentioned in the index either, so the name is rather to be interpreted as meaning the plants from
America. The associated material by Michaux also suggests that.
O. virginica var. eglandulosa Spach, Annales des sciences naturelles, Botanique ser. 2 16: 246 (Spach
1841b). – Type: Cult. at Trianon Garden, 1833, s. coll., s.n. (lecto-, designated here: P06811525!). –
Note: the specimen designated as lectotype here bears a note by Spach as author “Nob.” after the
name.
O. virginica var. glandulosa Spach, Annales des sciences naturelles, Botanique ser. 2 16: 246 (Spach
1841b). – O. virginiana var. glandulosa (Spach) Sarg., Botanical Gazette 67: 216 (Sargent 1919). –
O. virginiana f. glandulosa (Spach) J.F.Macbr., Publications of the Field Museum of Natural History,
botanical series 4 (7): 192 (MacBride 1929). – Type: Cult. at h. P. [hortus Parisiensis], 1833, s. coll.,
s.n. (lecto-, designated here: P06809319!). – Note: the specimen designated as lectotype here bears
a note by Spach as author “Nob.” after the name.
O. ostrya (L.) MacMill., The Metaspermae of the Minnesota Valley: 287 (MacMillan 1892) nom. inval.
(tautonym), p.p. – Note: MacMillan referred only to the Virginian part of Linnaeus’ Carpinus ostrya
L., but he produced a tautonym, which is not valid according to Art. 23.4 of the ICN (McNeill et al.
2012).
O. virginiana var. lasia Fernald, Rhodora 38: 414 (Fernald 1936). – O. virginiana subsp. lasia (Fernald)
A.E.Murray, Kalmia 12: 22 (Murray 1982). – Type: USA, Florida, Columbia Co., Lake City, 11‒19
Jul. 1895, G.V. Nash 2158 (holo-: GH00033787!; iso-: E00870598!, K!, P06810598!, P06810607!,
US00089313!, WU!). – Additional type material: USA, Virginia, Henrico Co., Richmond, 5 May
1894, J.R. Churchill s.n. (para-: not localized); ibid., Princess Anne Co., rich dry woods, Little Neck,
36
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
M.L. Fernald et al. 4627 (para-: not localized); ibid., Norfolk Co., dry rich woods, east of Gertie,
M.L. Fernald et al. 4628 (para-: not localized).
7b. Ostrya virginiana subsp. guatemalensis (H.J.P.Winkl.) A.E.Murray
Kalmia 13: 10 (Murray1983). – O. italica var. guatemalensis H.J.P.Winkl., Das Planzenreich IV
61 (Heft 19): 22 (Winkler 1904). – O. guatemalensis (H.J.P.Winkl.) Rose, Contributions from
the United States National Herbarium 8 (4): 292 (Rose 1905). – O. virginiana var. guatemalensis
(H.J.P.Winkl.) J.F.MacBr., Publications of the Field Museum of Natural History, Botanical Series
4 (7): 193 (MacBride 1929). – Type: Guatemala, Volcán de Fuego, 6400 ft, 7 Aug. 1873, O. Salvin
s.n. (lecto-, designated here: W1886–8378!; isolecto-: K!). – Additional type material: Guatemala,
J. Donnell Smith in Donn. Sm. Pl. Guatem. 2470 (syn-: K!); ibid., San Miguel Uspantán, Depart.
Quiché, 6000 ft, Apr. 1892, E.T. Heyde and E. Lux in Donn. Sm. Pl. Guatem. 2910 (syn-: K!,
M-0265489!, MO-188762!, WU!); “Guatemala und Costa Rica” (Winkler 1904), J. von Warszewicz
55 (syn-: not localized). Mexico, Orizaba, Aug. 1853, F. Müller 1526 (syn-: W0065339!); Mexico,
Jalapa, 23 May 1899, C.G. Pringle 8188 (syn-: BM000993542!, BR0000005269754!, E00870592!,
F, GOET!, K!, M-0265488!, MANCH!, P06810561!, P06810571!, P06810594!, PH00020034!,
US00898166, UVMVT221269!, W1900–3036!, WU!). – Note: the lectotype was chosen to be the
Salvin specimen in W because it is the only Guatemalan specimen that bears a note by Winkler. The
Pringle 8188 specimen in W also bears a Winkler note but Rose separated it to his O. mexicana
Rose. Choosing this specimen as lectotype would make plenty of isolectotypes, but a new name
would become necessary in case that the Mexican and the Guatemalan specimens turn out to be
diferent after all.
O. mexicana Rose, Contributions from the United States National Herbarium 8 (4): 292 (Rose 1905).
– Type: Mexico, Guerrero, Omilteme, 25 May 1903, E.W. Nelson 7050 (lecto-, designated here:
US00089315!; isolecto-: NY00253871!, NY00253872!). – Additional type material: Mexico,
Orizaba, M. Botteri 284 (syn- : GOET!, US00898164); ibid., Jalapa, 23 May 1899, C.G. Pringle
8188 (syn-: BM000993542!, BR0000005269754!, E00870592!, F, GOET!, K!, M-0265488!,
MANCH!, P06810561!, P06810571!, P06810594!, PH00020034!, US00898166, UVMVT221269!,
W1900–3036!, WU!).
8. Ostrya yunnanensis W.K.Hu
Acta Phytotaxonomica Sinica 17 (1): 87 (Li 1979b). – Type: China, Yunnan, Lu-chuan Hsien, in thin
forest, 2600 m a.s.l., 5 Dec. 1938, P.Y. Mao [Mao Pin-yi] 1935 (lecto-, designated here: PE00021988!;
isolecto-: KUN446238, KUN446239, PE00734260!, PE00734261!).
Discussion
Our inding that 78 basionyms are missing in major loristic treatments, 46 (plus 3 likely too late to be
included) in the World Checklist and the bibliography of Fagales (Govaerts & Frodin 1998), and 32
in the WCSP (2016) alone is thought-provoking. Most of the basionyms that have not been treated in
the checklists are below species level and none was considered as accepted upon publication. Floristic
treatments draw a diferent picture though, although it must be noted that full synonymy is often not given
for stylistic reasons. However, missing four accepted species in the Flora of China should be alarming
given its importance as a major tool for species identiication. Most of the basionyms that have been
missed, in general, are likely explained due to the unavailability of the protologues, mainly published
in literature from the former Soviet Union, China, and Japan. This is also most likely the explanation
for the overlooked species in the Flora of China. The fact that IPNI did not record infraspeciic names
before 1971 contributes to the gap. The old-fashioned concept of formalizing minor morphological
diferences in new taxon names (Endersby 2009) led to a multitude of names that needs to be revised.
37
European Journal of Taxonomy 375: 1–52 (2017)
Page (2016) coined the term “dark texts” for literature that is not digitally available. However, that
term can be expanded to all literature that is unavailable, even if the obstacle is formed by language or
the nescience or unavailability of the hard-copy literature. Limited access to material in collections far
away from the natural origin, and limited access to, or knowledge of existing literature were – and still
are – pitfalls for taxonomy. With new taxa being described constantly, overlooked names are a source
of error contributing to the instability of names. This is demonstrated by the fact that the name Ostrya
multinervis had already been published for a fossil species (Turner 2014) or by the description of a new
Ostryopsis intermedia B.Tian & J.Q.Liu although a valid basionym, Corylus davidii var. cinerascens
Franch. was already available (Holstein & Weigend 2016). The immense eforts to digitize literature and
herbaria during the last years are an enormous help for taxonomic work (Knapp et al. 2002; Wheeler
2004, Godfray 2007; Turner 2014), but there is a need for 1) an expansion to publications in non-Latin
script, and 2) from a structured review of old literature. The former, then, would proit if the script were
not only readable, but also markable for automatic translation. This is of special relevance for texts
before 1935 for the question of validity (Art. 39.1, McNeill et al. 2012).
Overlooking names due to limited access to literature or nescience is critical not only for the sake of
completeness, but because it may lead to overlooking potentially endangered taxa and underestimating
biodiversity. Taxon diferences are hypotheses that are tested traditionally using morphological
characters, and rejection of such hypotheses ends in synonymization of the applied names (Gaston &
Mound 1993; Holstein & Luebert 2017). Uncritical adoption of synonymizations might ease taxonomic
and loristic work at irst, but may well be the source of incorrect subsequent taxonomical decisions.
When synonymizations were made, material availability may have been limited or type material may
not have been accessible. Critical revision including new data is therefore important (Wheeler 2004).
Furthermore, adopting synonymy and ignoring infraspeciic taxa appears to be justiied, but this is only
true if species are regarded as a homogeneous entity. Those who rely on the recognition of heterogeneity
within species, such as population geneticists, plant breeders or phytochemists may beg to difer.
Overlooking infraspeciic taxa should therefore not be belittled (Nixon & Wheeler 1990).
The percentage of non-cultivar synonyms in relation to accepted basionyms is 66.8% in Carpinus
and 62% in Ostrya, both being on the lower spectrum of the numbers found by Scotland & Wortley
(2003) for a range of other taxa. However, the number of synonyms between the accepted taxon names
greatly difers among the Carpinus taxa. While another 14 names represent cultivars of C. betulus,
many synonymous basionyms belong to wide-spread species. Some of them were likely created due
to limited access to material, thus overemphasizing the diferences, e.g., in the treatments of Winkler
(1904, 1914) and partly of Hu (1929, 1930, 1931, 1932, 1933). Although Iwasaki et al. (2010) report
only a few instances of hybridization between C. cordata and C. japonica, and between C. tschonoskii
and C. laxilora, some of the synonyms, also in C. turzaninovii, might be explained by hybridization,
or introgression. The ive names under C. × schuschaensis may represent either natural variability or
diferent degrees of introgressive hybridization between C. betulus and C. orientalis, which had already
been suggested in the Flora of the USSR (Bobrov 1936). A good number of synonyms, however, was
described in regional treatments or even outside a wider taxonomic treatment. An exaggerated focus on
local taxa ignoring the species outside of the respective province or country borders carries the risk of
overemphasizing local variability and describing more wide-spread taxa repeatedly (Mabberley 1991;
Scotland & Wortley 2003; Michelangeli 2005). The aim to produce loras (target 1 of the GSPC), which
usually focus on regional to national scales (Funk 2006), might thus create problems, since the whole
scope of a species is ignored.
The IUCN Red List (International Union for Conservation of Nature and Natural Resources; status
from 29 March 2016) lists 27 species of Carpinus with 13 of them classiied as “data deicient” and 2,
C. putoensis and C. tientaiensis, as “critically endangered”. Shaw et al. (2014) also regard C. hebestroma
38
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
as critically endangered, whereas it is listed as “not assessed” online. For Ostrya, the Red List gives data
for only six species with O. chisosensis and O. rehderiana listed as critically endangered. As there are
at least 42 accepted species plus 17 infraspeciic taxa of, partly, poorly known distribution in Carpinus
alone, the conservation status is clearly very insuiciently documented for this genus. Changing this
situation may not be easy, since the acceptance status difers among the current treatments and checklists
in 21 of the Carpinus names, with four species missing in the Flora of China. The stability of taxon status
and recognizability of taxa are, however, crucial for conservation, comparison, and monitoring (Mace
2004). The fact that revisionary work is important for conservation has also been shown elsewhere, for
example in Juncaceae and Potamogetonaceae (Kirschner & Kaplan 2002) or in Cotula (Powell et al.
2014). Target 2 of the Global Strategy for Plant Conservation (GSPC) of the Convention of Biological
Diversity (CBD, https://www.cbd.int) aims at an assessment of the conservation status of all known
plant species. Meeting target 2, however, not only requires an assessment of the taxonomic status as-is,
but also critical revisions.
One of the crucial tasks of taxonomic revision, being the localization and identiication of type material
is tackled here for Carpinus and Ostrya. This is, however, not always straight-forward, and collections
are often wide-spread among diferent herbaria. The valuable information on where potential type
collections are located in Taxonomic Literature II (Staleu & Cowan 1976–1988) help to prioritize
herbarium visits, but may leave many collections being ignored for a long time. We found that many
types of most (in case of Ostrya even all) taxa are already available in silico, but many of them have not
been marked as type. Completely (or nearly so) digitized collections, such as the ones of the Muséum
national d’Histoire naturelle (Paris), Naturalis (L, U, WAG), and the Chinese Virtual Herbarium (PE
and partly others) were therefore very convenient. Digitizing type specimens may be a priority and ease
taxonomic work quickly (Berents et al. 2010), but complete digitization is inevitable. Our inding of the
unrecognized type of C. orientalis in BM among the specimens in the general collection may serve as
a good example, apart from our result that all surveyed herbaria contained hitherto unrecognized type
material. Digital collections have been conirmed as a useful low-cost tool (Steudel et al. 2012), but they
also save time and help preserving specimens thanks to the reduced risks of transportation and handling
damage (Friis 2017). However, information on the history of the collectors and collections, usually
the expertise from curatorial staf, is a not to be underestimated asset. Premature decisions, such as
inappropriate lecto- or neotypiications due to only partly digitized collections may become problematic,
just as ongoing taxonomic changes or novelties due to unavailability of (even more modern) literature
and physical material. We therefore consider open and easy access to fully-digitized collections to be not
only cost-efective in the long-term but also a prerequisite for correct taxonomic work, especially since
digital collections are being used more and more.
Conclusion
Despite the great eforts of digitization, we still ind yet undiscovered taxon names in Carpinus and
Ostrya, and even recent loristic treatments miss taxa. This is especially critical if publications are
hardly accessible and in languages that are not widely understood, which then may explain the limited
accessibility in libraries. The loss of names, and therefore hypotheses for taxa, due to oversight of littleknown publications, language barriers, and uncritical synonymization, contributes to future taxonomic
instability or overlooked biodiversity. Regional treatments ignoring taxa from neighbouring countries
might do more harm than good by adding more (superluous) names. The urgency to inventory biodiversity
is without dispute, but it should not go without critical revisionary work. Digital collections turn out to
be very valuable for this work, but with many types not recognized as such, partial digitization might
give the dangerous illusion of taxonomic work already completed, where it has not even really started.
39
European Journal of Taxonomy 375: 1–52 (2017)
Acknowledgements
We are grateful to the various literature and herbarium digitization projects that enable the easy access
to the resources, and to the curators and staf of B, BM, BR, E, GOET, K, LIV, M, MANCH, MSB, W,
and WU for making the collections available. We also thank Adam Boratynski, Hans-Joachim Esser,
Tilo Henning, Qi Lin, Zurab Manvelidze, Sergei L. Mosyakin, Alexander Sukhorukov, Genevieve
K. Walden, and the library staf of B, BM, and K for their help in obtaining literature. Thanks are
given to Federico Luebert for discussions on taxonomy, Tianyun Liu for translation help from Chinese
language, to two anonymous reviewers and to Alejandro Quintanar Sánchez for additions and helpful
comments. This research received support (visit to E collection) from the SYNTHESYS Project
(http://www.synthesys.info/), which is inanced by the European Community Research Infrastructure
Action under the FP7 “Capacities” Program.
References
Aiton W. 1789. Hortus Kewensis. Vol. 3. Printed for George Nicol, Bookseller to his majesty, Pall Mall,
London. https://doi.org/10.5962/bhl.title.4504
Akiyama S., Thijsse G., Esser H.-J. & Ohba H. 2013. Siebold and Zuccarini’s type specimens and
original material from Japan, Part 2. Angiosperms. Dicotyledoneae 1. Journal of Japanese Botany 88:
346‒377.
Ashburner K., McAllister H.A. & Rix M. 2013. The genus Betula: a taxonomic revision of birches. Kew
Publishing, Kew.
Bean W.J. 1914. Trees and Shrubs, Hardy in the British Isles. Vol. 1. J. Murray, London.
https://doi.org/10.5962/bhl.title.45879
Beck von Mannagetta G. 1890. Flora von Nieder-Österreich. Vol. 1. C. Gerold’s Sohn, Wien.
Beissner L. 1899a. Besuch der fürstl. Lobkowitz’schen Besitzung Eisenberg. Mitteilungen der Deutschen
dendrologischen Gesellschaft 8: 132–134.
Beissner L. 1899b. Besuch zu Muskau. Mitteilungen der Deutschen dendrologischen Gesellschaft 8:
134–137.
Beissner L., Schelle E. & Zabel H. 1903. Handbuch der Laubholzbenennung. Paul Parey, Berlin.
Available from: https://archive.org/details/Laubholzbenennung1903 [accessed: 1 Jul. 2015].
Berents P., Hamer M. & Chavan V. 2010 Towards demand-driven publishing: approaches to the
prioritization of digitization of natural history collection data. Biodiversity Informatics 7: 47–52.
https://doi.org/10.17161/bi.v7i2.3990
Blume C.L. 1851. Museum Botanicum Lugduno-Batavum. Vol. 1 (20): 305‒320. E.J. Bril, Leiden.
https://doi.org/10.5962/bhl.title.274
Bobrov E.G. 1936. Coryleae. In: Komarov V.L. (ed) Flora of the U.S.S.R. Vol. 5: 253–268. Izdatel’stvo
Akademii Nauk SSSR.
Braun H. 1889. Flora von Oesterreich-Ungarn. D. Ungarn. Oesterreichische Botanische Zeitschrift
39 (6): 233‒236.
Brickell C.D., Alexander C., David J.C., Hetterscheid W.L.A., Leslie A.C., Malecot V., Jin X. & Cubey
J.J. 2009. International Code of nomenclature for cultivated plants. Scripta horticulturae 10: 1–184.
Britton N.L. & Brown A. 1913. An Illustrated Flora of the Northern United States ed. 2. Vol. 1. Charles
Scribner’s sons, New York. https://doi.org/10.5962/bhl.title.55504
40
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
Browicz K. 1972. 97: Corylaceae. In: Rechinger K.H. (ed.) Flora Iranica. Akademische Druck - und
Verlagsanstalt, Graz.
Brummitt R.K. 1993. Report of the Committee for Spermatophyta: 39. Taxon 42 (4): 873‒879.
https://doi.org/10.2307/1223273
Camus A. 1930 [“1929”]. Espèces asiatiques nouvelles du genre Carpinus. Bulletin de la Société
botanique de France 76: 966‒969.
Cariot A. 1889. Études des Fleurs ed. 8. Vol. 2. Lyon.
Cesati V. 1844. Saggio su la Geograia Botanica e su la Flora della Lombardia. Tipograia Bernardoni,
Milano. Available from: https://books.google.de/books?id=Hc4ZAAAAYAAJ [accessed: 25 Nov. 2017].
Chang C.-S. & Chang K.S. 2010. Typiication of Corylopsis coreana (Hamamelidaceae) and Carpinus
laxilora var. longispica (Betulaceae). Journal of Japanese Botany 85: 270‒276.
Chang C.-S., Kim H. & Chang K.S. 2014a. Provisional checklist of Vascular plants for the Korea
Peninsula Flora (KPF). Designpost, Pajo.
Chang C.-S., Kim H. & Chang K.S. 2014b. A catalogue of Vascular Plant type specimens from Korea.
Designpost, Pajo.
Chen J.P. 2015. Flora of Hubei. Available from:
http://www.nature-museum.net/BioBook/HuBeiFlora/1/home.html [accessed 1 Jul. 2015].
Chen Y. 1937. Illustrated Manual of Chinese Trees and Shrubs. Agricultural Association of China,
Nanjing.
Cheng W.C. 1932. Two new ligneous plants from Chekiang. Contributions from the Biological
Laboratory of the Science Society of China, Botanical Series 8: 72‒76.
Cheng W.C. 1933. An enumeration of Vascular Plants from Chekiang, II. Contributions from the
Biological Laboratory of the Science Society of China, Botanical Series 9 (1): 59‒91.
Cheng W.C. 1950. 中国树木新种. 科学 [Science] 32 (8): 246–249.
Chun W.Y. 1927. New species and new combinations of Chinese plants. Journal of the Arnold Arboretum
8 (1): 19‒22.
Correll D.S. 1965. Some additions and corrections to the lora of Texas. Wrightia 3 (7): 126‒140.
Coville F.V. 1894. New or little-known plants. Ostrya knowltonii, a new species of hop hornbeams.
Garden and Forest 7: 114‒115.
Department of the Interior (ed.). 1928. Report of the National Monument Investigation. Plants
8 [天然紀念物調査報告 植物之部 第 8輯]. Tōkōshoin [刀江書院], Tokyo. Available from:
http://dl.ndl.go.jp/info:ndljp/pid/1139904 [accessed: 7 Jul. 2015].
Desfontaines R.L. 1804. Tableau
https://doi.org/10.5962/bhl.title.13828
de
l’École
de
Botanique.
J.A.
Brosson,
Paris.
Diels L. 1901. Die Flora von Central-China. Botanische Jahrbücher für Systematik, Planzengeschichte
und Planzengeographie 29: 169‒659.
Ding B.C., Wang S.Y. & Chao T.B. 1980. 河南新植物. Journal of the Henan Agricultural College
14 (2): 1‒10.
Dippel L. 1891 [“1892”]. Handbuch der Laubholzkunde. Vol. 2. Paul Parey, Berlin.
https://doi.org/10.5962/bhl.title.12978
41
European Journal of Taxonomy 375: 1–52 (2017)
Dodoens R. 1616. Remberti Dodonaei mechliniensis medici caesarei Stirpium historiae pemptades
sexsive libri xxx ed. 2. Balthasar et Joannus Morretos, Antwerp. https://doi.org/10.5962/bhl.title.7120
Dulac J. 1867. Flore du Département
https://doi.org/10.5962/bhl.title.10724
des
Hautes-Pyrénées.
F.
Savy,
Paris.
Editorial Board of the Flora Sichuanica. 2012. Flora Sichuanica. Vol. 21. Sichuan Publishing Group,
China.
Endersby J. 2009. Lumpers and splitters: Darwin, Hooker, and the search for order. Science 326: 1496–
1499. https://doi.org/10.1126/science.1165915
Ettingshausen C. 1868. Die fossile Flora der älteren Braunkohleformation der Wetterau. Sitzungsberichte
der Kaiserlichen Akademie der Wissenschaften, Wien, Mathematisch-Naturwissenschaftliche Classe
Abteilung 1, 57: 807–893.
Fang D. & Wang Y.S. 1983. A new species of Ostrya (Corylaceae) from Guangxi. Guihaia 3 (3): 189‒191.
Fernald M.L. 1935. Midsummer vascular plants of Virginia. Rhodora 37: 423‒454.
Fernald M.L. 1936. Plants from the outer coastal plain of Virginia. Rhodora 38: 414‒452.
Fliche P. 1888. Sur les formes du genre Ostrya. Bulletin de la Société botanique de France 35: 160‒172.
Forbes F.B. & Hemsley W.B. 1899 [“1890”]. An enumeration of all the plants known from China
proper, Formosa, Hainan, Corea, the Luchu Archipelago, and the Island of Hongkong, together
with their distribution and synonymy. Journal of the Linnean Society, Botany 26 (178): 1‒592.
https://doi.org/10.1111/j.1095-8339.1899.tb01642.x
Franchet A. 1899. Plantarum sinensium ecloge tertia. Journal de Botanique [Morot] 13 (7): 198‒208.
Friis I. 2017. Old tropical botanical collections: how to improve their availability, comprehensibility and
use in modern taxonomy? Webbia 72 (1): 5–16. https://doi.org/10.1080/00837792.2017.1301708
Frivaldszky von Frivald E. 1837. Balkány vidéki természettudományi utazás [Natural history journey
taken in Balkan region]. A’Magyar Tudós Társaság Évkönyvei 3 (3): 156–184. Available from:
http://opacplus.bsb-muenchen.de/title/7526457/ft/bsb10500528?page=638 [accessed: 1 Jul. 2015].
Funk V. 2006. Floras: a model for biodiversity studies or a thing of the past? Taxon 55 (3): 581–588.
Furlow J.J. 1987. The Carpinus caroliniana complex in North America. II. Systematics. Systematic
Botany 12 (3): 416‒434. https://doi.org/10.2307/2419267
Furlow J.J. 1997. Betulaceae. In: Flora of North America Editorial Committee (ed.) Flora of North
America: North of Mexico. Vol. 3. Oxford University Press, New York.
Gaston K.J. & Mound L.A. 1993. Taxonomy, hypothesis testing and the biodiversity crisis. Proceedings
of the Royal Society of London B 251: 139–142. https://doi.org/10.1098/rspb.1993.0020
Gilibert
J.E.
1792.
Exercitia
Phytologica.
J.B.
Delamollière,
Available from: http://catalogue.bnf.fr/ark:/12148/cb37275795s [accessed: 1 Jul. 2015].
Godfray H.C.J. 2007. Linnaeus
https://doi.org/10.1038/446259a
in
the
information
age.
Nature
446:
Lyon.
259–260.
Göppert
H.R.
1855.
Die
tertiäre
Flora
von
Schossnitz
in
Schlesien.
Heyn’sche
Buchhandlung,
Görlitz.
Available
from:
http://www.mdz-nbn-resolving.de/urn/resolver.pl?urn=urn:nbn:de:bvb:12-bsb10226015-1 [accessed: 1
Jul. 2015].
42
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
Govaerts R. & Frodin D.G. 1998. World checklist and bibliography of Fagales (Betulaceae, Corylaceae,
Fagaceae and Ticodendraceae). Royal Botanic Gardens, Kew.
Gray S.F. 1821. A Natural Arrangement
https://doi.org/10.5962/bhl.title.43804
of
British
Plants.
Vol.
2.
London.
Grenier J.C.M. & Godron D.A. 1855. Flore de France. Vol. 3 (1). J.B. Baillière, Paris.
https://doi.org/10.5962/bhl.title.6635
Grimm G.W. & Renner S.S. 2013. Harvesting Betulaceae sequences from GenBank to generate
a new chronogram for the family. Botanical Journal of the Linnean Society 172: 465–477.
https://doi.org/10.1111/boj.12065
Grossheim A.A. 1940. K sistematike drevesnykh porod Kavkaza. Izvestiya Azerbaidzhanskogo Filiala
[Akademii Nauk SSSR] 1940 (5): 32‒38.
Hance H.F. 1868 [“1869”]. On the Fagus Castanea of Loureiro’s ’Flora Cochinchinensis’ with
descriptions of two new Chinese Corylaceae. Journal of the Linnean Society, Botany 10 (44): 199‒203.
https://doi.org/10.1111/j.1095-8339.1868.tb00437.x
Handel-Mazzetti H.R.E. von 1925 [“1924”]. Plantae novae Sinensis, diagnosibus brevibus descriptae
a Dre. Henr. Handel-Mazzetti (29. Fortsetzung). Anzeiger der Öesterreichischen Akademie der
Wissenschaften Mathematisch-Naturwissenschaftliche Klasse 61: 162‒170.
Handel-Mazzetti H.R.E. von 1931. Kleine Beiträge zur Kenntnis der Flora von China I. Oesterreichische
Botanische Zeitschrift 80: 337‒343.
Hatushima S. 1934. Preliminary reports on the lowering plants and ferns collected in the Nansen
Experimental Forest of Kyushu Imperial University. Bulletin of the Kyushu Imperial University Forests
5: 1‒280.
Hayashi Y. 1952. 日本産樹木新報知(I). 林業試験場研究報告 57: 151‒157.
Hayashi Y. 1955. 日本産樹木新報知(II). 林業試験場研究報告 77: 25‒34.
Hayashi Y. 1960. 日本産樹木新報知(4). 林業試験場研究報告 125: 67‒78.
Hayashi Y. 1974. 日本産植物新報知. Journal of Geobotany [北陸の植物] 22: 4‒6.
Hayata B. 1913. Icones Plantarum Formosanarum. Vol. 3: 1‒197. Bureau of Productive Industry,
Formosa.
Hayata B. 1916. Icones Plantarum Formosanarum. Vol. 6: 1‒163. Bureau of Productive Industry,
Formosa.
Heufel J.A. 1858. Enumeration plantarum Banatus Temesiensis. Verhandlungen der ZoologischBotanischen Gesellschaft in Wien 8: 39–240.
Hill J. 1757 [“1756”]. The British Herbal. London.
Holstein N. & Luebert F. 2017. Taxonomy: stable taxon boundaries. Nature 548: 158.
https://doi.org/10.1038/548158d
Holstein N. & Weigend M. 2016. An annotated checklist and lectotypiication of Ostryopsis Decne.
(Betulaceae), with notes on the numbering system in the collections of P.J.M. Delavay. Phytotaxa
282 (3): 235–238. https://doi.org/10.11646/phytotaxa.282.3.9
Honda M. 1933. Nuntia ad Floram Japoniae XXI. The Botanical Magazine [Tokyo] 47: 433‒437.
Horikawa T. 1953. Key to the species of Carpinus of Japan and Corea. Acta Phytotaxonomica et
Geobotanica 15 (1): 12–13. https://doi.org/10.18942/bunruichiri.KJ00002992590
43
European Journal of Taxonomy 375: 1–52 (2017)
Host
N.T.
1831.
Flora
austriaca.
Vol.
2.
Frider,
Beck,
Vienna.
Available from: https://books.google.de/books?id=PO8UAAAAYAAJ [accessed: 1 Jul. 2015].
Hsia W.Y. 1934. Two new Carpinus from Szechuan. Contributions from the Institute of Botany, National
Academy of Peiping 2: 179‒180.
Hu H.H. 1929. Two new species of Carpinus from Szechuan. Journal of the Arnold Arboretum 10:
154‒156.
Hu H.H. 1930. Notulae systematicae ad loram sinensem. Journal of the Arnold Arboretum 11: 48‒50.
Hu H.H. 1931. Plantae Tsiangianae Corylaceae. Sinensia 2: 79‒93.
Hu H.H. 1932. Notulae systematicae ad Floram Sinensem IV. Journal of the Arnold Arboretum 13:
333‒336.
Hu H.H. 1933. A review of the genus Carpinus in China. Sunyatsenia 1 (2‒3): 103‒120.
Hu H.H. 1948a. Notulae Systematicae ad Floram Sinensem XI. Bulletin of the Fan Memorial Institute
of Biology n.s. 1 (2): 141‒152.
Hu H.H. 1948b. On four new species of Carpinus from South-Western China. Bulletin of the Fan
Memorial Institute of Biology n.s. 1 (2): 185‒189.
Hu H.H. 1949. Four new species of Carpinus from Yunnan. Bulletin of the Fan Memorial Institute of
Biology n.s. 1 (3): 213‒218.
Hu H.H. 1964. The materials on the monography of gen. Carpinus Linn. of China. Acta Phytotaxonomica
Sinica 9 (3): 281‒298.
Hu H.H. & Chun W.Y. 1927. Icones Plantarum Sinicarum. Vol. 1. Commercial Press, Shanghai.
Hubei Institute of Botany (eds) 1976. 湖北植物志 [Flora Hupehensis]. Vol. 1. 湖北人民出版社 [Hubei
People’s Publishing House], Wuhan.
Iamonico D. & Reveal J.L. 2012. Lectotypiication of the Linnean name Carpinus ostrya L. (Betulaceae).
Taxon 61 (4): 866.
Instituto Botanico Boreali-occidentali Academiae Sinicae Edita (ed.). 1974. Flora Tsinlingensis. Vol.
1 (2). Chinese Academy of Sciences, Beijing.
International Union for Conservation of Nature and Natural Resources. 2016. The IUCN Red List of
threatened species. Available from: http://www.iucnredlist.org/ [accessed 29 Mar. 2016].
Iwasaki T., Tono A., Aoki K., Seo A. & Murakami N. 2010. Phylogeography of Carpinus japonica
and Carpinus tschonoskii (Betulaceae) growing in Japanese deciduous broad-leaved forests,
based on chloroplast DNA variation. Acta Phytotaxonomica et Geobotanica 61 (1): 1–20.
https://doi.org/10.18942/apg.KJ00006537150
Jeon J.I. & Chang C.-S. 1997. Reconsideration of Carpinus L. (Betulaceae) of Korea primarily based on
quantitative characters. Korean Journal of Plant Taxonomy 27 (2): 157‒187.
Kárpáti Z. 1937. Dendrológiai jegyzetek. I. Két új gyertyánfa-változat. Botanikai Közlemények 34 (5–
6): 194–196.
Kawamoto T. 1939. Handbook of Korea-Manchurian Forestry. Forest Experiment Station, Korea.
Kemularia-Nathadze L.M. 1937. New data on the lora of Georgia (USSR). Trudy Tbilisskogo
Botaniceskogo Instituta 2: 131‒138.
Kirschner J. & Kaplan Ž. 2002. Monographs in relation to global Red Lists. Taxon 51 (1): 155–158.
https://doi.org/10.2307/1554973
44
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
Knapp S., Bateman R.M., Chalmers N.R., Humphries C.J., Rainbow P.S., Smith A.B., Taylor P.D.,
Vane-Wright R.I. & Wilkinson M. 2002. Taxonomy needs evolution, not revolution. Nature 419: 559.
https://doi.org/10.1038/419559a
Koch K.H.E. 1873. Dendrologie; Bäume, Sträucher und Halbsträucher, welche in Mittel- und NordEuropa im Freien kultivirt werden. Vol. 2. F. Enke, Erlangen. https://doi.org/10.5962/bhl.title.20459
Koehne
B.A.E.
1893.
Deutsche
Dendrologie.
Ferdinand
Enke,
Stuttgart.
Available from: http://bibdigital.rjb.csic.es/ing/Libro.php?Libro=3517&Pagina=1 [accessed: 1 Jul.
2015].
Koidzumi G. 1913. Spicilegium Betulacearum Japonicarum novarum vel minus cognitarum. The
Botanical Magazine [Tokyo] 27: 143‒149. https://doi.org/10.15281/jplantres1887.27.319_143
Koidzumi G. 1940. Contributiones ad Cognitionem Florae Asiae Orientalis. Acta Phytotaxonomica et
Geobotanica 9 (2): 71‒81. https://doi.org/10.18942/bunruichiri.KJ00002594589
Krüssmann G. 1951. Die Laubgehölze. Paul Parey, Berlin.
Lamarck J.B.A.P.M. de. 1779.
https://doi.org/10.5962/bhl.title.9461
Flore
Françoise.
Vol.
2.
Imprimérie
Royale,
Paris.
Lee W.T. 1996. Lineamenta Florae Koreae. Academy Publisher, Seoul.
Léveillé M.H. 1904. Nouveautés chinoises, coréennes et japonaises. Bulletin de la Société botanique de
France 51: 422‒424.
Léveillé M.H. 1905. Quelques Amentacées nouvelles d’Extrême-Orient. Bulletin de la Société botanique
de France 52: 141‒143.
Li J.-H. 2008. Sequences of low-copy nuclear gene support the monophyly of Ostrya and
paraphyly of Carpinus (Betulaceae). Journal of Systematics and Evolution 46 (3): 333–340.
https://doi.org/10.3724/SP.J.1002.2008.08026
Li P.C. 1979a. Betulaceae. In: Wang W.T. (ed.) Flora Reipublicae Popularis Sinicae. Vol. 27: 37‒59.
Science Press, Beijing.
Li P.C. 1979b. New taxa of Betulaceae from China. Acta Phytotaxonomica Sinica 17 (1): 87‒91.
Li P.C. & Skvortsov A.K. 1999. Betulaceae. In: Wu Z.Y. & Raven P.H. (eds) Flora of China. Vol. 4:
286‒301. Science Press, Beijing, and Missouri Botanical Garden Press, St. Louis.
Li Y.K. & Mang X.M. 1983. Three new tree species from Guizhou province [贵州树木的三个新种].
Guizhou Science 1983 (2): 18‒24.
Liang J.Y. 1986. A new species of Carpinus from Guangxi. Guihaia 6 (4): 275‒276.
Liang S.B. & Zhao F.Z. 1991. A new species of Carpinus from Shandong. Bulletin of Botanical Research,
Harbin 11 (2): 33‒34.
Liao J.C. 1996. Betulaceae. In: Editorial Committee of the Flora of Taiwan (ed.) Flora of Taiwan. Vol.
2: 44‒50. Editorial Committee of the Flora of Taiwan, Taipei.
Lin Q. & Qi C.J. 2000. 湖南木本植物新分类群. Bulletin of Botanical Research, Harbin 20 (1): 1–6.
Lin Q. & Sun Q. 2007. Lectotypiication of two names in Betulaceae. Acta Botanica Boreali-Occidentalia
Sinica 27 (2): 177–178.
Lin Q., Sun Q., Sun M. & Li H.L. 2007. Lectotypiications of twenty-eight names of Chinese taxa in
Angiospermae. Acta Botany Boreal-Occident Sinica 27 (6): 1247‒1255.
45
European Journal of Taxonomy 375: 1–52 (2017)
Linnaeus
C.
1753.
Species
https://doi.org/10.5962/bhl.title.669
Laurentius
Plantarum.
Linnaeus C. 1763. Species Plantarum
https://doi.org/10.5962/bhl.title.11179
ed.
2.
Salvius,
Laurentius
Salvius,
Stockholm.
Stockholm.
Liu K.W. & Lin Q.Z. 1986. A new species of Carpinus from Hunan. Bulletin of Botanical Research,
Harbin 6 (2): 143‒145.
Loddiges J.C. 1779. A Catalogue of plants and seeds. C. Heydinger, London.
Loddiges J.C. 1783. Catalogue of plants ed. 2. London.
Loddiges W. 1836. Catalogue of plants ed. 16. W. Wilson, London.
Loudon J.C. 1838. Arboretum et Fruticetum Britannicum. Vol. 3. Longman, Orme, Brown, Green, and
Longmans, London. https://doi.org/10.5962/bhl.title.37723
Lu Z.-Q., Liu S.-Y., Yang X.-Y., Liang Q.-L., Yang Y.-Z., Zhang D., Milne R. & Liu J.-Q. 2017.
Carpinus langaoensis (Betulaceae), a new hornbeam species from the Daba Mountains in Shaanxi,
China. Phytotaxa 295 (2): 185–193. https://doi.org/10.11646/phytotaxa.295.2.6
Ludwig C.F. 1783. Die neuere wilde Baumzucht. Müller, Leipzig. Available from:
http://www.mdz-nbn-resolving.de/urn/resolver.pl?urn=urn:nbn:de:bvb:12-bsb10293944-2 [accessed: 1
Jul. 2015].
Lundell C.L. 1939. Studies of Mexican and Central American Plants VII. Lloydia 2: 73‒108.
Mabberley D.J. 1991. The problem of older names. In: Hawksworth D.L. (ed.) Improving the stability
of names: needs and options: 123–134. Koeltz Scientiic Books, Königstein.
MacBride J.F. 1929. Variants of Ostrya and a Texas Tephrosia. Publications of the Field Museum of
Natural History, Botanical Series 4 (7): 192‒193.
MacMillan C. 1892. The Metaspermae
https://doi.org/10.5962/bhl.title.10852
of
the
Minnesota
Valley.
Minneapolis.
Mace G.M. 2004. The role of taxonomy in species conservation. Philosophical Transactions of the
Royal Society B 359: 711–719. https://doi.org/10.1098/rstb.2003.1454
Makino T. 1912. Observations on the Flora of Japan. The Botanical Magazine [Tokyo] 26: 384‒399.
https://doi.org/10.15281/jplantres1887.26.312_384
Makino T. 1914. Observations on the Flora of Japan. The Botanical Magazine [Tokyo] 28: 31‒36.
https://doi.org/10.15281/jplantres1887.28.326_31
Marshall
H.
1785.
Arbustrum
https://doi.org/10.5962/bhl.title.68506
Americanum.
Joseph
Crukshank,
Philadelphia.
Massé A. 1853. Nouveautés horticoles, d’origine française, obtenues par les semis. Revue Horticole ser.
4 2: 269–272.
Maximowicz C.J. 1881a. Diagnoses plantarum novarum asiaticarum IV. Mélanges biologiques tirés du
Bulletin de l’Académie impériale des sciences de St. Pétersbourg 11: 155‒350.
Maximowicz C.J. 1881b. Diagnoses plantarum novarum asiaticarum IV. Bulletin de l’Academie
impériale des sciences de St.-Pétersbourg 27: 425‒460.
McCauley R.A. & Paces M.N. 2015. A preliminary analysis of phylogeography and species
diversiication of Ostrya (Betulaceae) in the southwestern US and Mexico. Published abstracts, Botany
2015. Edmonton, Alberta, Canada.
46
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
McNeill J., Barrie F.R., Buck W.R., Demoulin V., Greuter W., Hawksworth D.L., Herendeen P.S., Knapp
S., Marhold K., Prado J., Prud’homme van Reine W.F., Smith G.F., Wiersema J.H. & Turland N.J. 2012.
International Code of Nomenclature for algae, fungi, and plants (Melbourne Code). Koeltz Scientiic
Books, Koenigstein.
Medwedew J.S. 1909. Главнейшие Ильмовые и Сережчатые растения Кавказа. Vestnik Tilisskogo
Botanicheskogo Sada 14: 1‒42.
Michaux A. 1803. Flora Boreali-Americana. Vol. 2. Carolus Grapelet, Paris and Strasburg.
https://doi.org/10.5962/bhl.title.330
Michaux F.-A. 1812. Histoire des Arbres Forestiers de l’Amérique Septentrionale. Vol. 3 (1). Imprimerie
de L. Haussmann et d’Hautel, Paris.
Michelangeli F.A. 2005. Tococa (Melastomataceae). Flora Neotropica 98: 1–114.
Micheli
P.A.
1729.
Nova
plantarum
https://doi.org/10.5962/bhl.title.49380
genera.
Bernardo
Paperini,
Florence.
Miller P. 1768. The Gardeners Dictionary ed. 8. John and Francis Rivington, London.
https://doi.org/10.5962/bhl.title.541
Miyabe K., Kudo Y., Suzaki C. 1920–1932. Icones of the essential forest trees of Hokkaido. Hokkaido
Government, Sapporo.
Moench C. 1785. Verzeichniss ausländischer Bäume und Stauden. Frankfurt et Leipzig. Available from:
http://www.mdz-nbn-resolving.de/urn/resolver.pl?urn=urn:nbn:de:bvb:12-bsb10297735-6 [accessed: 1
Jul. 2015].
Moench
C.
1794.
Methodus.
https://doi.org/10.5962/bhl.title.304
Oicina
nova
libraria
academiae,
Marburg.
Mólnar V.A. 2015. Kitaibel. Egy magyar tudós élete című könyv. Debreceni Egyetem TTK Növénytani
Tanszék, Debrecen.
Münchhausen O. von 1770. Der Hausvater. Vol. 5 (1). Verlag Seel. Nic. Försters und Sohns Erben HofBuchhandlung, Hanover. https://doi.org/10.5962/bhl.title.13511
Murray A.E. 1982. Notae Spermatophytae. Kalmia 12: 18‒27.
Murray A.E. 1983. Notae Spermatophytae. No. 2. Kalmia 13: 3‒11.
Nakai T. 1911. Flora Koreana. Pars secunda. The Journal of the College of Science, Imperial University
of Tokyo, Japan 31: 1‒573.
Nakai T. 1912. Notulae ad Plantas Japoniae et Coreae VIII. The Botanical Magazine [Tokyo] 26:
321‒328. https://doi.org/10.15281/jplantres1887.26.311_321
Nakai T. 1914. Plantae novae Coreanae et Japonicae. I. Repertorium Specierum Novarum Regni
Vegetabilis 13: 243‒250.
Nakai T. 1915. Præcursores ad loram sylvaticam Koreanam II. The Botanical Magazine [Tokyo] 29:
33–47. https://doi.org/10.15281/jplantres1887.29.340_35
Nakai T. 1926. Notulae ad Plantas Japoniae & Koreaea XXXI. The Botanical Magazine [Tokyo] 40:
161‒171. https://doi.org/10.15281/jplantres1887.40.161
Nakai T. 1931. Notulae ad Plantas Japoniae & Koreaea XL. The Botanical Magazine [Tokyo] 45: 91‒137.
https://doi.org/10.15281/jplantres1887.45.91
47
European Journal of Taxonomy 375: 1–52 (2017)
Neilreich
A.
1861.
Nachträge
zu
Maly’s
Enumeratio
plantarum
phanerogamicarum imperii austriaci universi. Braumüller, Wien. Available from:
http://www.mdz-nbn-resolving.de/urn/resolver.pl?urn=urn:nbn:de:bvb:12-bsb10302442-6 [accessed: 1
Jul. 2015].
Nicholson G. 1883. The hornbeam. The Garden (London 1871–1927) 24: 418‒420.
Nixon K.C. & Wheeler Q.D. 1990. An ampliication of the phylogenetic species concept. Cladistics 6:
211–223. https://doi.org/10.1111/j.1096-0031.1990.tb00541.x
Ohwi J. 1965. Flora of Japan [English version]. Smithonian Institution, Washington D.C.
Oliver D. 1891. Icones Plantarum. Vol. 20 (4). Williams and Norgate, London and Edinburgh, R.
Friedländer und Sohn, Berlin.
Olshanskyi I.G. 2014. Підродина Coryloideae J.D. Hooker (Betulaceae) у флорі України [The
subfamily Coryloideae J.D. Hooker (Betulaceae) in the lora of Ukraine]. Вісник ЛНУ імені Тараса
Шевченка Біологічні науки 12 (295): 62‒74.
Pallas P.S. 1801. Bemerkungen auf einer Reise in die Südlichen Statthalterschaften des Russischen
Reichs. Vol. 2. Gottfried Martin, Leipzig. https://doi.org/10.5962/bhl.title.48469
Page R. 2016. DNA barcoding and taxonomy: dark taxa and dark texts. Philosophical Transactions of
the Royal Society B 371: 20150334. https://doi.org/10.1098/rstb.2015.0334
Palmer M.W., Wade G.L. & Neal P. 1995. Standards for the writing of loras. BioScience 45 (5): 339–
345. https://doi.org/10.2307/1312495
Pei C. 1948. Flowering plants of Northwestern China II. Botanical Bulletin of Academia Sinica 2:
215‒227.
Petzold E.C.A. & Kirchner G. 1864. Arboretum Muscaviense. W. Opetz, Gotha. Available from:
https://books.google.de/books?id=a5UCAAAAYAAJ [accessed: 1 Jul. 2015].
Plukenet
L.
1692.
Phytographia.
Vol.
3.
London.
Available
http://bibdigital.rjb.csic.es/ing/Libro.php?Libro=4827&Pagina=1 [accessed: 1 Jul. 2015].
from:
Powell A.M. 1998. Trees and Shrubs of the Trans-Pecos and Adjacent Areas. University of Texas Press,
Austin, Texas.
Powell R.F., Boatwright J.S. & Magee A.R. 2014. A taxonomic revision of the Cotula coronopifolia
group (Asteraceae) and implications for the conservation statuses of the species. South African Journal
of Botany 93: 105–117. https://doi.org/10.1016/j.sajb.2014.03.008
Qi C.J. 1981. Three new woody species from Hunan Province. Journal of Nanjing Technological College
of Forest Products 1981 (3): 123‒126.
Qi C.J. 1984. New taxa from Hunan. Acta Phytotaxonomica Sinica 22 (6): 493‒494.
Qu S.Z. & Wang K.Y. 1988. A new variety of Carpinus shensiensis Hu from Shaanxi. Bulletin of
Botanical Research, Harbin 8 (4): 111‒112.
Radde-Fomina O. 1929. До питання систематики роду Carpinus в межах C.P.C.P. Trudy izychnomatematychnogo viddilu Kiev [Mémoires de la Classe des Sciences Physiques et Mathématiques de
l’Académie des Sciences de l’Ukraine] 15 (1): 51‒107.
Rainesque C.S. 1811. An essay on the exotic plants, mostly European, which have been naturalized,
and now grow spontaneously in the Middle States of North America. The Medical Repository ser. 3 2:
330‒345.
48
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
Rainesque C.S. 1817. Florula Ludoviciana. C. Wiley and Co., New York. Available from:
http://gallica.bnf.fr/ark:/12148/bpt6k973548 [accessed: 1 Jul. 2015].
Rehder A. 1919. New species, varieties and combinations from the herbarium and the collections of the
Arnold Arboretum. Journal of the Arnold Arboretum 1: 44‒60.
Rehder A. 1930. New species, varieties and combinations from the herbarium and the collections of the
Arnold Arboretum. Journal of the Arnold Arboretum 11: 153‒168.
Rehder A. 1938. New species, varieties and combinations from the herbarium and the collections of the
Arnold Arboretum. Journal of the Arnold Arboretum 19: 71‒87.
Reichenbach H.G.L. 1850. Icones Florae Germanicae et Helveticae. Vol. 12. Fridericus Hofmeister,
Lispiae. https://doi.org/10.5962/bhl.title.6353
Reinecke K.L. 1911. Neue Beiträge zur Kenntnis der Flora von Thüringen. Mitteilungen des Thüringischen
Botanischen Vereins 28: 36‒43.
Reveal J.L. 1990. (979) Proposal to conserve the name and type of Carpinus virginiana P. Miller
(Betulaceae), the basionym of Ostrya virginiana (P. Miller) K. Koch. Taxon 39 (2): 357‒359.
https://doi.org/10.2307/1223075
Rochel A. 1828. Plantae Banatus rariores. Ludovici Landerer de Füskút, Pest. Available from:
https://books.google.de/books?id=Iw9JAAAAcAAJ [accessed: 1 Jul. 2015].
Rose J.N. 1905. Studies of Mexican and Central American plants—No. 4. Contributions from the United
States National Herbarium 8 (4): 281‒339.
Rushforth K. 1987. Keys to Carpinus and Ostrya. Plantsman 8: 249‒256.
Salisbury R.A. 1796. Prodromus Stirpium in Horto ad Chapel Allerton Vigentium. London.
https://doi.org/10.5962/bhl.title.427
Sanguinetti P. 1864. Flora Romanae Prodromus Alter. Rome. https://doi.org/10.5962/bhl.title.9982
Sargent C.S. 1893. Notes on the Forest Flora of Japan.—XXI. Garden and Forest 6: 383‒385.
Sargent C.S. 1894. Forest lora of Japan. Houghton, Milin and Company, Boston and New York.
https://doi.org/10.5962/bhl.title.18184
Sargent C.S. 1896. The Silva of North America. Vol. 9. Houghton, Milin and Company, Boston and
New York. https://doi.org/10.5962/bhl.title.14668
Sargent C.S. 1919. Notes on North American trees. IV. Botanical Gazette 67: 208‒242.
Săvulescu T. 1952. Flora Republicii Populare Române. Vol. 1. Academiei republicii populare Române,
București.
Schneider C.K. 1904. Illustriertes Handbuch der Laubholzkunde. Vol. 1. Gustav Fischer, Jena.
https://doi.org/10.5962/bhl.title.194
Schneider C.K. 1912. Illustriertes Handbuch der Laubholzkunde. Vol. 2. Gustav Fischer, Jena.
https://doi.org/10.5962/bhl.title.194
Schur P.J.F. 1866. Enumeratio
https://doi.org/10.5962/bhl.title.9958
Plantarum
Transsilvaniae.
Wilhelm
Braumüller,
Wien.
Schwarz O.K.A. 1949. Beiträge zur Nomenklatur und Systematik der mitteleuropäischen Flora.
Mitteilungen der Thüringischen Botanischen Gesellschaft 1: 82–119.
Scopoli
J.A.
1760.
Flora
https://doi.org/10.5962/bhl.title.6681
Carniolica.
49
Johann
Thomas
Trattner,
Vienna.
European Journal of Taxonomy 375: 1–52 (2017)
Scopoli J.A. 1772. Flora Carniolica ed. 2. Vol. 2. Johann Paulus Krauss, Vienna.
Scotland R.W. & Wortley A.H. 2003. How many species of seed plants are there? Taxon 52: 101–104.
https://doi.org/10.2307/1224723
Serres P.M.T. de. 1823. Voyage dans le Tyrol. Vol. 2. Nepveu, Paris. Available from:
https://books.google.de/books?id=3AxfAAAAcAAJ [accessed: 1 Jul. 2015].
Shaw K., Stritch L., Rivers M., Roy S., Wilson B. & Govaerts R. 2014. The Red List of Betulaceae.
Botanic Gardens Conservation International, Richmond.
Siebold P.F. von & Zuccharini J.G. 1846. Florae japonicae familiae naturales, adjectis generum et
specierum exemplis selectis. Sectio prima. Plantae dicotyledoneae polypetalae. Abhandlungen der
Mathematisch-Physikalischen Klasse der Königlich Bayerischen Akademie der Wissenschaften 4 (3):
109‒240.
Simon-Louis [frères]. 1902. Preisverw., Herbst. 1902–1903. Metz.
Smith J.D. 1890. Undescribed plants from Guatemala VIII. Botanical Gazette 15 (2): 27‒29.
Spach É. 1841a. Notes sur les Carpinus. Annales des sciences naturelles, Botanique ser. 2 16: 248–254.
Spach É. 1841b. Notes sur les Ostrya. Annales des sciences naturelles, Botanique ser. 2 16: 243–248.
Spach É. 1842. Histoire naturelle des végétaux: Phanérogames. Vol. 11. Librairie encyclopédique de
Roret, Paris. https://doi.org/10.5962/bhl.title.44839
Staleu F.A. & Cowan R.S. 1976–1988. Taxonomic literature II. Bohn, Scheltema et Holkeman, Dr. W.
Junk b.v., Publishers, Utrecht, Antwerpen, The Hague, Netherlands, Boston, USA.
Standley P.C. 1920. Trees and shrubs of Mexico. Contributions from the United States National
Herbarium 23 (1): 1‒170.
Steudel E.G. von. 1841. Nomenclator Botanicus ed. 2. Vol. 1. J.G. Cotta, Tübingen.
https://doi.org/10.5962/bhl.title.655
Steudel B., Kessler M. & Nyfeler R. 2012. Using the internet to aid in discovery of unrecognized type
material. Phytotaxa 62: 13–24. https://doi.org/10.11646/phytotaxa.62.1.4
Sudworth G.B. 1893. On legitimate authorship of certain binomials with other notes on nomenclature.
Bulletin of the Torrey Botanical Club 20: 40‒46.
Sugimoto J. 1961. New Keys of Japanese Trees [日本樹木総検索誌]. Rokugatsusha Publishing [六月
社], Osaka.
Sun Y.L., Wang D., Lee H.B., Park W.G., Kwon, O.W. & Hong, S.K. 2011. Phylogeny of Korean
Hornbeam (Carpinus turczaninovii) based on nuclear ribosomal ITS sequence. African Journal of
Biotechnology 10 (76): 17435–17442. https://doi.org/10.5897/AJB11.1337
Tang G. 2007. A revision of “The Metasequoia lora and its phytogeographic signiicance” [对《水杉
区系及其植物地理学意义》一文的订正]. Journal of Nanjing Forestry University (Natural Science)
1987 (1): 88‒104.
Tatewaki M. 1953. The primary survey of the vegetation of the Hokkaido Experiment Forest, Kyushu
University. Bulletin of the Kyushu University Forests 21: 1–60.
Thiers B. 2016. Index Herbariorum: A global directory of public herbaria and associated staf. New York
Botanical Garden’s Virtual Herbarium. Available from: http://sweetgum.nybg.org/science/ih/ [accessed
22 Jul. 2016].
50
HOLSTEIN N. & WEIGEND M., Taxonomic checklist of Carpinus and Ostrya
Tong Y.H., Pang K.S. & Xia N.H. 2014. Carpinus insularis (Betulaceae), a new species
from Hong Kong, China. Journal of Tropical and Subtropical Botany 22 (2): 121‒124.
https://doi.org/10.3969/j.issn.1005-3395.2014.02.002
Tournefort J.P. de. 1700. Institutiones Rei Herbariae. Vol. 1. Paris. https://doi.org/10.5962/bhl.title.713
Tournefort J.P. de. 1703. Corollarium institutionum rei herbariae. Paris. Available from:
http://gallica.bnf.fr/ark:/12148/bpt6k97536w [accessed: 1 Jul. 2015].
Trautvetter E.R. von. 1884. Incrementa lorae phaenogamae rossicae. Fasc. III. Trudy Imperatorskago
S.-Peterburgskago Botaniceskago Sada 9: 1‒220.
Tung S.L. 1981. Study on Betulaceae of North-Eastern China [东北桦木科植物的研究]. Bulletin of
Botanical Research, Harbin 1 (1‒2): 123‒151.
Turner I.M. 2014. Names of extant Angiosperm species that are illegitimate homonyms of fossils.
Annales Botanici Fennici 51: 305–317. https://doi.org/10.5735/085.051.0506
Tutin T.G. & Walters S.M. 1993. Corylaceae. In: Tutin T.G., Burges N.A., Chater A.O., Edmondson
J.R., Heywood V.H., Moore D.M., Valentine D.H., Walters S.M. & Webb D.A. (eds) Flora Europaea 1:
70–71. Cambridge University Press, Cambridge.
Tzvelev N.N. 2004. Flora Vostochnoĭ Evropy. Vol. 11. KMK Scientiic Press, St. Petersburg & Moscow.
Uyeki H. 1940. Woody-Plants and their Distributions in Tŷsen. Agricultural and Forestry College,
Suigen.
Uyeki H. & Lee J.-O. 1924. Forest distribution pattern around the 35°N latitude in Chosen. Suigen
Gakuho 41: 3‒13.
Vos C. de 1867. Beredeneerd woordenboek der voornaamste heesters en coniferen, in Nederland
gekweekt. J.B. Wolters, Groningen. https://doi.org/10.5962/bhl.title.112499
Waldstein F.P.A. & Kitaibel P. 1803‒1805. Descriptiones et Icones Plantarum Rariorum Hungariae. Vol. 2.
Mattias Andreas Schmidt, Vienna. Available from: http://bibdigital.rjb.csic.es/ing/Libro.php?Libro=1918
[accessed: 1 Jul. 2015].
Wallich N. 1831. Plantae Asiaticae
https://doi.org/10.5962/bhl.title.468
Rariores.
Vol.
2.
Treuttel
and
Würtz,
London.
Walter T. 1788. Flora Caroliniana. J. Fraser, London. https://doi.org/10.5962/bhl.title.9458
Wang, Y.S. & Huang, J.P. 1985. New species of Carpinus from Guangxi, China. Guihaia 5 (1): 15‒16.
WCSP. 2016. World Checklist of Selected Plant Families. Facilitated by the Royal Botanic Gardens,
Kew. Available from: http://apps.kew.org/wcsp/ [accessed 8 Jul. 2016].
Wheeler Q.D. 2004. Taxonomic triage and the poverty of phylogeny. Philosophical Transactions of the
Royal Society of London B 359: 571–583. https://doi.org/10.1098/rstb.2003.1452
Willdenow C.L. 1805. Species
https://doi.org/10.5962/bhl.title.727
Plantarum
ed.
4.
Vol.
4
(1).
G.C.
Nauk,
Berlin.
Willkomm H.M. 1887. Forstliche Flora von Deutschland und Oesterreich ed. 2. C.F. Winter’sche
Verlagshandlung, Leipzig and Heidelberg.
Winkler H.J.P. 1904. Betulaceae. In: Engler H.G.A. (ed.) Das Planzenreich IV 61 (Heft 19). Wilhelm
Engelmann, Berlin.
Winkler H.J.P. 1914. Neue Revision der Gattung Carpinus. Botanische Jahrbücher für Systematik,
Planzengeschichte und Planzengeographie 50 (Suppl.): 488‒508.
51
European Journal of Taxonomy 375: 1–52 (2017)
Yamamoto Y. 1932. Supplementa Iconum Plantarum Formosarum. Vol. 5. Department of Forestry,
Taihoku.
Yang C.C. 1997. Distribution of the Woody Plants in Sichuan [四川树木分布]. Guizhou Science and
Technology Publishing House, Guiyang.
Yi T.P. 1992. A new species of genus Carpinus L. from Sichuan. Bulletin of Botanical Research, Harbin
12 (4): 335‒337.
Ying S.S. 1988. Coloured Illustrated Flora of Taiwan. Published by the author, Taipei.
Yoo K.-O. & Wen J. 2002. Phylogeny and biogeography of Carpinus and subfamily Coryloideae
(Betulaceae). International Journal of Plant Sciences 163 (4): 641–650. https://dx.doi.org/10.1086/340446
Yoo K.-O. & Wen J. 2007. Phylogeny of Carpinus and subfamily Coryloideae (Betulaceae) based
on chloroplast and nuclear ribosomal sequence data. Plant Systematics and Evolution 267: 25–35.
https://doi.org/10.1007/s00606-007-0533-2
Zapałowicz H. 1908. Conspectus Florae Galiciae Criticus. Vol. 2. Academiae litterarum Cracoviensis,
Kraków.
Manuscript received: 20 May 2016
Manuscript accepted: 19 September 2016
Published on: 7 December 2017
Topic editor: Koen Martens
Desk editor: Alejandro Quintanar
Printed versions of all papers are also deposited in the libraries of the institutes that are members of the
EJT consortium: Muséum national d’Histoire naturelle, Paris, France; Botanic Garden Meise, Belgium;
Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London, United
Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History Museum
of Denmark, Copenhagen, Denmark; Naturalis Biodiversity Center, Leiden, the Netherlands; Museo
Nacional de Ciencias Naturales-CSIC, Madrid, Spain; Real Jardín Botánico de Madrid CSIC, Spain.
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