Further taxonomic notes on the brinjal eggplant complex: a response to

Meyer et al. (2013)

John Samuelsa,*, Alan Child

Abstract
The response to Samuels (2013a) offered by Meyer et al. (2013) has necessitated further discussion.
The designation “Asian S. incanum” used by Meyer et al. actually refers to S. insanum sensu Lester
and Hasan. Its use has confused the issue of the taxonomic relationship between Middle Eastern
Solanum incanum L. sensu stricto and Indian S. insanum, although distinction is, in fact, supported by
the molecular evidence of Meyer et al. (2012). The group of brinjal relatives that Meyer et al. call S.
undatum Lam. is distinct from S. cumingii Dunal (=S. melongena L. subsp. cumingii [Dunal]
Samuels). The validity of the name S. insanum L. has yet to be assessed, but it is an example of a
nomen ambiguum, and should be considered for rejection. We re-iterate the likelihood of a sole
domestication event in India, with transmission eastwards from the centre of origin to south-eastern
China. We also recommend a collaborative, multidisciplinary approach for future studies, and stand
by the necessity for consistency in the identification and nomenclature of experimental material.

Keywords: brinjal eggplant, domestication, nomen ambiguum, Solanum incanum, Solanum insanum,
Solanum melongena.

1. Introduction
The phylogeographic study of Meyer et al. (2012) presented viewpoints on nomenclature and
domestication that were commented on by Samuels (2013). One of the comments made in response to
their paper involves the taxonomic uncertainties relating to the species in question (Samuels, 2013a).
Unfortunately, by referring to “Solanum section Melongena Bitter pro parte” in their preamble,
Meyer et al. (2013) raise further uncertainty. It is thus necessary to specify the precise group of plants
studied by Meyer et al. Solanum melongena L., the brinjal eggplant (or brinjal), and its closest wild
and weedy relatives can be classified under Solanum section Melongena (Miller) Dunal, series
Incaniformia Bitter (Child and Lester, 2001; Lester et al., 2011). Apart from S. marginatum L. and
some lesser-known taxa, they may also be referred to as the “brinjal eggplant complex” (Daunay et
al., 1991; Samuels, 2013b).

2. Unbiased Approach
Meyer et al. (2013) state that the purpose of their 2012 study was to examine eggplant domestication
“...with as little influence as possible from any previously published nomenclatural scheme...” Such
an approach helps to alleviate any anthropogenic bias that might be introduced due to an a priori
classification based on morphological characters (Duminil and di Michele, 2009). Yet, Meyer et al.
make judgements on issues relating to species names, authorities and types-thus introducing their own
bias. Hence, current discussion of their work has had to focus on several pivotal taxonomic issues.

a
Novel Solanaceae Crops Project, Penzance, TR20 8XD, United Kingdom
*
Corresponding author; e-mail address: john.samuels@virgin.net

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 3. Possibility of misidentification
The possibility of misidentification of collections was suggested by Samuels (2013a) in an attempt to
understand some of the patterns in genomic data (contrary to what Meyer et al. [2013] suggest).
Confusion of brinjal with its wild and cultivated relatives has been noted (Lester and Hasan, 1991)
and the possibility of misidentification should not be underestimated. It has been reported concerning
herbarium specimens (e.g. Daunay and Hazra, 2012; Samuels, 2012a, 2013b; Vorontsova and Knapp,
2012), and germplasm collections (e.g. Daunay et al, 1999; Daunay and Hazra, 2012; Furini and
Wunder, 2004; Karihaloo et al., 2009; Knapp et al., 2013).

4. Recent system of Samuels (2013b)

Meyer et al. (2013) suggest that the taxonomic issues raised by Samuels (2013a) are based on
previous work (i.e. Samuels, 2012a, 2012b). This is incorrect as, of all the species at issue, original
work in these papers only relates to S. incanum L. sensu stricto, and reference to multiple sources of
information is made. Meyer et al (2013) mention that: “Samuels’ recent system could not be
considered by them because it was published after their paper.” The system they refer to (Samuels,
2013b) is based on the detailed examination of approximately 2500 herbarium specimens from around
20 herbaria (collected in Africa and Asia), as well as an ecogeographic survey of around 800
herbarium specimens (from Africa and the Middle East). This, alongside studies of other living
collections of brinjal and relatives (Samuels, unpubl.), has provided an “in-depth and geographically
broad evaluation of specimens,” as recommended by Meyer et al. (2013).

5. S. incanum sensu Meyer et al.

Most of the accessions studied by Meyer et al. (2012) are from various parts of Asia, including SW
Asia, where some S. incanum s. str. accessions were originally collected. Meyer et al. (2013)
comment that they only meant for “Asian accessions of S. incanum s. str.” to be included under the
taxon they called S. undatum Lam. This is confusing because, instead of “S. incanum s. str.,” they
probably mean S. incanum sensu Meyer et al. (2013: 307). It now seems they were originally referring
to South Asian (i.e. Indian) “S. incanum.” This corresponds to “S. insanum,” as defined by Lester and
Hasan (1991). Meyer et al. (2012: Figures 2 and 3) provide graphical data for accessions of both S.
incanum s. str. and S. incanum sensu Meyer et al. in the same figures; with the foregoing in mind, this
makes interpretation extremely difficult.

6. Nomenclatural difficulties
The comment (Meyer et al., 2013) that Samuels (2013a) argues that “Asian S. incanum, S. insanum
and S. undatum...should be maintained as distinct species...” is inaccurate. The latter author, in fact,
puts forward that S. incanum s. str., S. insanum sensu Lester and Hasan, and S. cumingii Dunal are
distinct taxa.

Application of the name S. incanum has been incredibly confused (Knapp et al., 2013) and it is
therefore important that it is applied in an unambiguous way. Adopting it (if only in part) for another
difficult taxon such as S. insanum L. (Meyer et al., 2012) only adds to the taxonomic confusion. The
name S. undatum, which Meyer et al. consider synonymous with S. cumingii, was not adopted by
Samuels. Meyer et al. (2013) adopt S. undatum on the grounds of taxonomic priority over S. cumingii.
However, this is only in order if both names refer to the same taxon; this is not the opinion of the
present authors. Samuels (2013a) pinpointed S. cumingii as a more appropriate name for the brinjal
wild relatives known as S. melongena group F, and without reference to the characteristics of the S.
undatum type specimen-contrary to what Meyer et al. (2013: 307) comment. Further examination of
the relevant types and protologues indicates clear distinction between the two taxa.

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 7. One species or two?
Meyer et al. (2012: 692, 696, 697) state that S. incanum (presumably S. incanum sensu Meyer et al.
[=S. insanum]) and S. undatum may be a single species. They add that morphological evidence
supports this view, although they provide no quantitative data to substantiate this. The molecular
support they mention (p. 696) seems to relate to nrITS data-in particular, the details of clade “W,” Fig.
2A. This shows three accessions of S. incanum from Israel and one Indonesian S. undatum accession
grouped together. This is confusing, as this seems to suggest conspecificity of S. incanum s. str. with
S. undatum.

On p. 697, Meyer et al (2012) suggest that “there may actually be two different species” amongst
accessions of S. incanum from different floristic regions. They add that their AFLP and nrITS
analyses provide evidence of distinction between Middle Eastern S. incanum and Asian “S. incanum”
accessions. The present authors concur with the delimitation of the two taxa.

8. The fate of S. insanum-revisited

Meyer et al. (2013) mention that the work of Knapp et al. (2013) supports the proposal to unify their
Asian wild taxa, and that the name S. insanum is “now used” for this group (with S. undatum and S.
cumingii synonymous with S. insanum)! The study of Meyer et al. (2012) seems to have been
formative in the unification of S. insanum and S. undatum/S. cumingii; this has thereby led to a
“chicken-and-egg” situation.

No nomenclatural opinions on S. insanum are given by Samuels (2013a)-contrary to Meyer et al.

(2013: 307)-but, apparently, there has been disagreement amongst some of the latter authors over the
validity of the name. Other viewpoints are held on the position of S. insanum (e.g. Lester et al., 2011;
http://www.plosone.org/annotation/listThread.action?root=63081). S. insanum has been described as
an unreliable taxon and a name that continues to be used in inconsistent ways-an example of a nomen
ambiguum that should be considered for rejection (Samuels, 2013b). It seems that, inadvertently,
Meyer et al. (2012, 2013) contribute to the argument for its rejection. Samuels (2013b) suggests that
S. cumingii (as S. melongena subsp. cumingii [Dunal] Samuels, =S. undatum auct. non Lam.) is a
more appropriate name for the feral forms of brinjal, found in many parts of South and SE Asia. This
would explain the AFLP clustering pattern (Meyer et al., 2012: Figure 3B) in which Malesian S.
undatum accessions group with accessions in cluster MC1, and not with those in cluster MC2.

9. Domestication
Meyer et al. (2012: 686) consider “Asian S. incanum” (=S. insanum) and S. undatum to be the
progenitor taxa involved in the domestication of brinjal. This equates to one taxon, S. insanum sensu
Knapp et al. (2013), but this has yet to be ratified or refuted, pending more detailed study.

Rather than multiple events in India, China and Malaysia (as proposed by Meyer et al.) Samuels
(2013a) suggested that a sole domestication event took place in India. Other studies re-enforce this
theory, e.g. that of Isshiki et al. (1994) which showed that there was an higher variation in isozyme
genotypes in brinjal accessions from South Asia than in accessions from China. In addition, Hurtado
et al. (2012) provided SSR marker evidence of high genetic diversity in accessions from Sri Lanka,
compared with those from China. Also using SSRs, Cericola et al. (2013) showed that PCA based on
geographical data produced distinctly geographical clusters of brinjal accessions (India, Indochina,
Indonesia, China). In addition, there was infiltration of Indian genotypes into the Chinese cluster,
suggesting transmission of brinjal from an initial region of domestication in India, eastwards to China
(Samuels, 2013b). In fact, first records of S. melongena pinpointed by Wang et al. (2008) in ancient
literature corroborate this by showing a temporal progression eastwards across south-eastern China.

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 10. Closing comments
Genomic research sometimes operates in isolation from other methodologies, largely because of the
highly specialised nature of its methods. Other studies on Solanum have combined molecular analyses
with different techniques, such as those using phenotypic markers. For example, Sunseri et al. (2010)
analysed the genetic diversity of African eggplants using an effective combination of morphological,
agronomic and molecular markers. For future studies we recommend such multidisciplinary inputs
which allow the combination of different skill sets. This collaborative approach may bring us nearer
to a consensus perspective on difficult taxa such as S. insanum, and to finally determine the details of
the domestication of brinjal. We also stand by the requirement for consistency in identification and
nomenclature of experimental material because, although names are “only the tags with which we
name biological entities,” they should nevertheless form part of an unambiguous and workable
system.

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