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Notes on the identity and typification of Zehneria thwaitesii (Schweinf.) C.Jeffrey and Zehneria tridactyla (Hook.f.) R. Fern. & A. Fern. (Cucurbitaceae)

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In Zehneria thwaitesii long standing uncertainties persisted with respect to taxonomic delimitation. This study is a first step in clarifying the case using both morphological and molecular phylogenetic analyses. We here provide a complete overview on the nomenclatural history of Z. thwaitesii. For the African plants hitherto included in the broadly circumscribed species the name Z. tridactyla is reinstated and a lectotype is designated. Diagnostic features distinguishing both taxa are summarised. Z. thwaitesii is now considered restricted to Asia.
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Phytotaxa 284 (2): 143–146
http://www.mapress.com/j/pt/
Copyright © 2016 Magnolia Press Correspondence PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Hugo de Boer: 26 Oct. 2016; published: 11 Nov. 2016
http://dx.doi.org/10.11646/phytotaxa.284.2.6
143
Notes on the identity and typification of Zehneria thwaitesii (Schweinf.) C.Jeffrey
and Zehneria tridactyla (Hook.f.) R. Fern. & A. Fern. (Cucurbitaceae)
C. BRÄUCHLER1$, MAYANK D. DWIVEDI2,3 & H. SCHAEFER2
1Restoration Ecology, Department of Ecology & Ecosystem Management, Technical University of Munich, Freising, Germany
2Plant Biodiversity Research, Department of Ecology & Ecosystem Management, Technical University of Munich, Freising, Germany
3Plant Systematic Laboratory, Department of Botany, University of Delhi, Delhi, India
$Author of correspondence: c.braeuchler@tum.de
In Zehneria thwaitesii long standing uncertainties persisted with respect to taxonomic delimitation. This study is a first step
in clarifying the case using both morphological and molecular phylogenetic analyses. We here provide a complete overview
on the nomenclatural history of Z. thwaitesii. For the African plants hitherto included in the broadly circumscribed species
the name Z. tridactyla is reinstated and a lectotype is designated. Diagnostic features distinguishing both taxa are sum-
marised. Z. thwaitesii is now considered restricted to Asia.
Keywords: Africa, Asia, Cucurbitales, Hooker, Kirk, nomenclature, typification, Welwitsch
Introduction
The cucurbit genera Melothria Linnaeus (1753: 35) and Zehneria Endlicher (1833: 69) have long been subject to discussion
with respect to both generic delimitation and circumscription of species included. The Old World species of the group
have either been accommodated in various genera (De Wilde & Duyfjes 2006, 2009) or united under one large genus
Zehneria Endlicher (1833: 69, Jeffrey 1962, Schaefer & Renner 2011). The latter concept has been supported by pollen
exine ornamentation (Van der Ham & Pruesapan 2006) and more recently by molecular phylogenetic analyses (Schaefer &
Renner 2011, De Boer et al. 2015). One of the species, Zehneria thwaitesii (Schweinfurth) Jeffrey (1962: 15), is subject to
molecular phylogenetic investigations at a larger geographic scale (Dwivedi et al., in prep.). In its current circumscription
this species is distributed from India to southern Tropical Africa with numerous names described from different parts of the
range (De Wilde & Duyfjes 2006, Renner & Pandey 2013). Our ongoing study, however, shows the Indian plants to form an
evolutionary lineage separate from the African taxa, a fact already recognised by Hooker (1871). He considered Melothria
tridactyla Hooker (1871: 562) (cited from the African countries Sudan, Mozambique, Angola, and Congo) as different
from the plants from Ceylon (Table 1). The typical element of Z. thwaitesii unambiguously originates from the latter island,
binding the name to that lineage. When included in a large Zehneria, the oldest available name for the African plants is
Z. tridactyla (Hooker) Fernandes & Fernandes (1962: 118) (≡ M. tridactyla Hooker (1871: 562), for which a lectotype is
designated below.
Nomenclatural treatment
Zehneria thwaitesii (Schweinfurth) Jeffrey (1962:15), non Z. deltoidea Miquel (1855 or 1857: 397)
Melothria thwaitesii Schweinfurth (1868: 44), nom. nov.
Bryonia deltoidea Thwaites ex Arnott (1836: 19), nom. illeg., non Schumacher & Thonning (1829: 429)
Aechmandra deltoidea Arnott (1841: 274)
Melothria deltoidea (Arnott) Thwaites (1859: 124), nom. illeg., non (Schumacher & Thonning) Bentham (1849: 368)
Melothria zeylanica Clarke (1879: 626), nom. superfl. [M. zeylanica Koenig ex Wight & Arnott (1834: 129) is not relevant here, for it
was published in synonymy only to Vitis pedata (Lamarck 1783: 31) Wallich ex Wight (1833: 26)]
Neoachmandra deltoidea (Arnott) de Wilde & Duyfjes (2006: 18)
Type:—[SRI LANKA] Ceylon, Walker 273 (Lectotype K000742779!, designated by Jeffrey (1962: 371))
= Bryonia deltoidea Thwaites, nom. nud. [in sched. Zeyl. 1610, 2581, 3128]
Notes:—Arnott (1836: 2) cites specimens in the herbaria of Graham (then at Edinburgh) and Hooker (then at Glasgow)
as basis for his descriptions. The relevant specimens for Z. thwaitesii were in Hooker’s possession and are kept at E (via
BRÄUCHLER ET AL.
144 Phytotaxa 284 (2) © 2016 Magnolia Press
GL; E00301187!) and K (via Herbarium Hookerianum, K000742779!) today. The specimen at K is a Walker collection
(“Ceylon, Col. Walker” on sheet and “No. 273 Bryonia” on a separate label) and has been annotated in Arnott’s hand
(“Bryonia certe, does not agree with any of my Peninsular species, B. deltoidea Arn.”). The specimen at E is labelled
as holotype (http://data.rbge.org.uk/herb/E00301187) and has been annotated in Hooker’s hand only (with an error in the
number copied from the original label: “275” [which is the number of a Phyllanthaceae] instead of “273”). Both specimens
together most likely originally formed the holotype, as indicated by the existence of one original label only. The gathering
as a whole was in Hooker´s possession then. Since it is now split in two and the respective duplicates are kept in different
collections, a lectotypification was necessary and the E specimen is best to be regarded as isolectotype. The K specimen has
been effectively and validly designated as such (Art. 7.9 & 7.10, ICN, McNeill et al. 2012) by Jeffrey (1962). Schweinfurth
(1868) included Indian and African plants in one broadly circumscribed taxon when giving it a new name under Melothria
(M. thwaitesii). In fact, the female parts illustrated by him stem from a collection from Ceylon. From a nomenclatural point
of view, the citation of African and other material by Schweinfurth is not relevant, because his account formally represents
publication of a new name (Art. 6.11, ICN, McNeill et al. 2012) typified by the type of the illegitimate Bryonia deltoidea.
Zehneria tridactyla (Hooker) Fernandes & Fernandes (1962: 118)
Melothria tridactyla Hooker (1871: 562); Cogniaux (1881: 596); Hiern (1898: 402); Durand & Durand (1909: 232); Cogniaux (1916:
275)
Type (lectotype, designated here):—[Africa, Mozambique] Shupanga, climbing on trees and among grass, April 1862, J. Kirk, s.n.
(K000313374!).
Further original material:—MOZAMBIQUE. Shupanga, June[?] 1862, J. Kirk s.n. (Syntype K000313373!); Congo,
September 1863, R. Burton s.n., (Syntype K000313436!); Angola, shady places, Golungo Alto, fl. and fr. middle of December
1855, F.M.J. Welwitsch [826] (Syntype BM, G00458275!, K, and probably numerous other herbaria, but see note below);
[Sudan] Reliq. Kotschy. t. 29, excl. pl. femina [illustration of male plants collected in 1861 at the White Nile “im Gebiete der
Tschier, unter 7° n. Br” by Harnier, W.v.]
Notes:—Hooker (1871: 562) provides a detailed description and cites several syntypes. Although citation of
Schweinfurth´s illustration is a reference to a previously published description (Art. 38.1(a); illustration with analysis) that
illustration thus represents part of the original material but cannot be considered a type. According to Jeffrey´s annotation
on the two Kirk specimens from Shupanga (K000313373! & K000313374!) he considered Kirk´s collections as “holotype”,
but a formal lectotypification for M. tridactyla was not published so far. It may well be both Kirk specimens represent
parts of one and the same gathering. K000313373 however has only the standard printed label “Livingstone´s Zambesi
Expedition. J. Kirk June 1862” and the determination added in Hooker’s hand. K000313374 in addition to that printed one
also has Kirk´s handwritten label indicating it as having been collected in April 1862. According to Livingstone´s itinerary
(Livingstone & Livingstone 1865: 414-423), the expedition was at Shupanga both in April and June 1862. In spite of this
remaining uncertainty on the precise date of collecting, we here designate K000313374 as a lectotype, for it also is the most
complete specimen. The Welwitsch syntype was cited from BM by Jeffrey (1962) and Fernandes & Fernandes (1962), where
the collections of Welwitsch were kept as one in 1871 (Hiern 1896). Duplicates today found in other herbaria, have been
distributed at a later point of time (e.g. G00458275 has been received in 1881). According to Albuquerque et al. (2009) the
study/top set of Welwitsch’s collections has been transferred to COI/LISU so, in theory, the original syntype should be kept
there. According to Hiern (1898) and Fernandes and Fernandes (1962), the syntype of M. tridactyla has been distributed
under the number 826, which does not represent a true collector’s number and the corresponding plants may actually stem
from different collections (Albuquerque et al. 2009). Also, plants distributed under that number sometimes are mislabelled
individuals of M. triangularis that normally have been distributed as Welwitsch 827 (Hiern 1898, Fernandes & Fernandes
1962, Dolezal 1974). Due to these ambiguities we refrain from using the Welwitsch syntype as lectotype, though in contrast
to the Kirk syntype, duplicates may be found in numerous collections.
TABLE 1. Morphological differences between African and Asian plants.
Characters Zehneria tridactyla (Africa) Zehneria thwaitesii (Asia)
Leaves Triangular, 3 lobed, lateral lobes at right angles with middle lobe Not lobed, deltoid or narrowly triangular
Fruit 1.9–2.5 cm long 2.5–3.5 cm long
Seeds Lenticular (biconvex), 2.5 mm long Flattened 3–4 mm long
NOTES ON Z. THWAITESII SCHWEINF. AND Z. TRIDACTYLA Phytotaxa 284 (2) © 2016 Magnolia Press 145
The differences provided in the table are based on the descriptions obtained from literature cited and type specimens of
the two species examined for this study.
Note:—Presence of ridges on the fruits cited for Zehneria thwaitesii by Philcox (1997) could not be confirmed because
the specimens consulted were lacking fully developed fruits.
Acknowledgements
The authors thank John McNeill, Stefan Dressler and two anonymous reviewers for valuable comments improving the
manuscript.
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The genus Zehneria Endl. is split into 5 genera, of which 4 are new: Indomelothria, Neoachmandra, Scopella, and Urceodiscus. Apart from several new combinations and taxa of subspecific rank, the following species are described as new: I. chlorocarpa, N. backeri, N. lancifolia, N. macrantha, N. nesophila, N. platysperma, U. arfakensis, U. carrii, U. hippocrepicus, U. parviflora, U. scabridula, U. viridis, Z. elbertii, Z. erythrobacca, Z. immarginata, Z. neocaledonica, Z. pedicellata, Z. pisifera, Z. tahitensis, and Z. trullifolia. The New World genus Melothria was introduced in Asia where it is represented by one species, M. pendula L.
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In order to provide additional data for the proposed subdivision of the palaeotropical genus Zehneria (Cucurbitaceae, tribe Benincaseae, subtribe Cucumerinae) by De Wilde and Duyfjes (2006)4. De Wilde , W. J. J. O. and Duyfjes , B. E. E. 2006a . Redefinition of Zehneria and four new related genera (Cucurbitaceae), with an enumeration of the Australasian and Pacific species. . Blumea , 51 : 1 – 88 . [Web of Science ®]View all references into five genera (Indomelothria, Neoachmandra, Scopellaria, Urceodiscus and Zehneria s.s.) pollen was examined using light and scanning electron microscopy for 18 Zehneria species including representatives of each of the proposed subdivisions, plus one species of the neotropical genus Melothria to which Zehneria once belonged. The pollen of Indomelothria, Scopellaria and Urceodiscus are described for the first time. Indomelothria, Neoachmandra and Zehneria s.s. have the same pollen type (3‐colporate, microreticulate to reticulate), which cannot be distinguished from that of Melothria. Scopellaria deviates by its irregularly striate‐reticulate ornamentation and short colpi, and Urceodiscus by its striate‐reticulate ornamentation. This result supports separation of Scopellaria and Urceodiscus as separate genera, but neither supports or rejects the segregation of Indomelothria and Neoachmandra. The pollen type shared by Indomelothria, Neoachmandra and Zehneria s.s. fits very well in the Cucumerinae. However, the striate‐reticulate ornamentation of Scopellaria and Urceodiscus pollen is unknown within this subtribe, and leaves some doubt with respect to the (sub)tribal assignment of both Scopellaria and Urceodiscus. Based on pollen morphology alone, the African Neoachmandra(?) peneyana does not belong to the tribe Benincaseae. Within the Cucurbitaceae, the 6‐aperturate pollen type of Neoachmandra (?) peneyana occurs only in the exclusively neotropical tribe Sicyeae.