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SUPPORTING INFORMATION
Delving into Delias Hübner (Lepidoptera: Pieridae): fine-scale biogeography,
phylogenetics and systematics of the world’s largest butterfly genus
Chris J. Müller, Pável F. Matos-Maraví and Luciano B. Beheregaray
Journal of Biogeography
Appendix S3 Delias systematics.
Taxonomic appraisal of Delias
The comprehensive taxon sampling in this study resolved 14 distinct clades. Both the
number of clades and their fine-scale composition differ from that of Talbot (1928–
1937), Yagishita et al. (1993) and Braby & Pierce (2007). Nonetheless, the broadscale phylogeny of the latter authors does not differ markedly from our own. Clade
relationships and composition within the new phylogenetic hypothesis was markedly
constant irrespective of the analysis [i.e. maximum parsimony (MP) or Bayesian
inference (BI)].
Our nysa clade corresponds to the singhapura clade of Braby & Pierce (2007),
which included the singhapura, nysa and georgina species groups of earlier authors
(Talbot, 1928–1937; Yagishita et al., 1993) and the brandti species group of Müller
(2001). Yagishita et al. (1993) separated the georgina group from the nysa group but
this is probably not warranted as D. nuydaorum Schröder and D. paoaensis Inomata
& Nakano (from the ‘georgina’ group of Yagishita et al., 1993) are both nested within
the ‘nysa’ complex in our analysis. The finer-scale relationships of the nysa clade
revealed in this study are in major disagreement to previous authors. The clade
comprises two main groups, with D. hempeli Dannatt, D. enniana Oberthür and D.
vidua Joicey & Talbot [previously assigned to the nysa as opposed to the singaphura
group by both Talbot (1928–1937) and Yagishita et al. (1993)] grouping with the
singaphura group. We agree with species level status for members of Yagishita’s
georgina group. Those that were known at the time were all included as subspecies
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of D. georgina Felder & Felder by Talbot (1928–1935) but since the publication of his
monograph sympatry between certain species has been recognized. We also concur
with Braby & Pierce (2007), who implied that D. nysa (Fabricius) and D. brandti
Müller are closely related. However, D. pulla and D. nysa were recovered as sister
species, with D. brandti sister to that pairing.
The isse clade also includes the dorimene clade and contains a mix of the
previously recognised species groups. Braby & Pierce (2007) made a similar inference
but included both with the hyparete species group in the isse clade, while in our
phylogeny the isse clade is sister to the nysa clade. Delias laknekei Miller, Simon &
Wills, a recently described species from New Ireland, Bismarck Archipelago, is sister
to D. lytaea (Godman & Salvin) from proximal New Britain. This relationship suggests
that the two species are allopatric and that D. lytaea is not likely to occur on New
Ireland, as was previously supposed (e.g. D’Abrera, 1971, 1977, 1990). Indeed, the
few records of D. lytaea from this island are dubious (Miller et al., 2007). Particularly
noteworthy in our phylogeny is the phylogenetic position of D. chrysomelaena in the
isse group, transferred from the ladas group (previously referred to as the
chrysomelaena group). We found that the composition of the morphologically
divergent ladas group, sampled entirely, is otherwise largely in agreement with that
proposed by Talbot.
For the most part, the pasithoe group forms an obvious grouping, as
suggested by previous authors (Talbot, 1928–1937; Yagishita et al., 1993), and is
corroborated in this study. Our phylogeny confirms that the distinctive D. benasu
Martin, transferred from the belladonna group by Morishita (1981), belongs in the
pasithoe group. Talbot (1928–1937) considered that D. benasu may be a link
between the pasithoe and belladonna groups but in our analyses D. benasu is sister
to D. woodi Talbot (also pasithoe group). Interestingly, D. blanca, transferred from
the nysa to the belladonna group by Yata (1985) is actually nested within the
pasithoe group. The belladonna group is sister to the hyparete clade and to the
pasithoe clade, in our MP and Bayesian analyses, respectively. Delias aganippe
Donovan is a particularly distinctive species that was provisionally placed, together
with the pasithoe group, within the belladonna group by Braby & Pierce (2007).
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However, it had weak support and grouped loosely with D. sagessa Fruhstorfer, so is
here given unique species group ranking.
The eichhorni clade was the focus of a study by Morinaka et al. (2002), who
demonstrated its monophyly, although only a single gene, ND5, was used in their
analysis. We only sampled three species from this group (D. eichhorni Rothschild &
Jordan, D. germana Roepke and D. gilliardi Sanford & Bennett), the grouping of
which coincided with that of Morinaka et al. (2002). The sagessa clade, herein newly
recognized, is another distinctive group of just a few species, none of which were
sampled by Braby & Pierce (2007). Indeed, they retained the classification of earlier
authors, who considered D. sagessa (and allies) to be within the geraldina group,
although Talbot (1928–1937) suggested that D. sagessa was not well placed and
might belong to the kummeri [nigrina] species group. The unique status given here is
supported by the sagessa group possessing very distinctive genitalia, with long apical
processes of the valva, quite different to members of the geraldina clade. In the
present study, the geraldina clade is taken to include only a few members of the
large, heterogeneous group of species originally proposed by Talbot (1928–1937)
and adopted by Yagishita et al. (1993) but comprises all three members of the
previously recognized stresemanni group. Indeed, Talbot (1928–1937) grouped D.
stresemanni Rothschild and D. lecerfi Joicey & Talbot on genitalic and androconial
affinities. Braby & Pierce (2007) grouped the cunningputi clade with (geraldina +
stresemanni) but the geraldina clade recognized herein is supported as a clade
distinct from the aroae (cunningputi) groups. The revised phylogeny shows that both
this and the aroae group contain species that were assigned to the geraldina group
by previous authors.
The subgenus Piccarda Grote was restored for the hyparete clade by Talbot
(1928–1937), based on characteristic androconial discs and male genitalia, and is a
well supported group in the present study. The species D. eileenae Joicey & Talbot,
which resembles some members of the dorimene group and was hence included in
that group by Talbot (1928–1937), was rightfully transferred to the hyparete group
by Yagishita et al. (1993). The belisama clade is another well-supported group but its
relationships were poorly resolved in the analysis of Braby & Pierce (2007). In our
analysis it is sister to (albertisi + nigrina). The albertisi clade comprises the niepelti,
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iltis, clathrata, albertisi and bornemanni groups of previous authors. We found that
there is much overlap of these groups, so their separation is not warranted. Shared
morphological characters are also present between the species (e.g. those of D.
albertisi Oberthür and D. discus Honrath with D. elongatus Kenrick). The albertisi and
nigrina clades were broadly grouped by Braby & Pierce (2007). However, both are
distinct, well-supported groups. Within the nigrina clade, both D. harpalyce Donovan
and D. wollastoni Rothschild were recovered as outlying members, although the
sequence (only COI) for the latter is very short. Talbot (1928–1937) also suggested D.
wollastoni was an outlying member of the nigrina group, based on examination of
the then unique type. Delias mayrhoferi Bang-Haas (=schunichii Morita) was
considered of uncertain status (e.g. Gotts & Ginn, 2003) but in our analysis, it is sister
to D. eximia Rothschild.
Barcoding Delias
Delias are characterized by several species groups containing numbers of species
that are phenotypically very similar, i.e. cryptic species. This is particularly evident
within the albertisi and nigrina species groups, all of which are endemic to the
uplands of mainland New Guinea. We assessed three sets of sibling species, using
the COI gene using two to four exemplars of each species. In all cases, we attempted
to use specimens from two locations where the species occur in sympatry. The first
set of species from the nigrina clade (D. kummeri Ribbe, D. ligata Jordan and D.
isocharis Rothschild & Jordan), well known for their subtle but consistent differences
of pattern on the underside hindwing, are widespread throughout much of the
central cordillera of New Guinea. For both MP and Bayesian analyses (see Fig. S1a
and S1b, respectively, below), specimens of each of these species grouped with high
support and the genetic differences between them averaged 3.9%, 3.2% and 3.4%
for kummeri-ligata, kummeri-isocharis and ligata-isocharis, respectively. Delias ligata
was originally described as a form of D. kummeri but Talbot (1928–1937) noted
differences in the androconia and male valva and that there were no intermediates.
Delias weiskei Ribbe and D. leucias Jordan, also in the nigrina clade, are sister taxa
exhibiting few external differences and have extensively overlapping distributional
ranges in mainland New Guinea. Our analysis showed a genetic divergence averaging
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3.3% between the taxa. A similar scenario was recorded for the group of closely
related species in the albertisi clade, namely D. callista Jordan, D. iltis Ribbe, D.
hapalina Jordan and D. luctuosa Jordan. Despite the similar external facies, the
pairwise divergence was significant between these species, with averages of 6.0%
(callista-luctuosa), 4.9% (callista-iltis), 4.0% (luctuosa-iltis), 7.1% (callista-hapalina),
4.0% (luctuosa-hapalina) and 3.3% (iltis-hapalina). The utility of the COI marker for
barcoding cryptic species has been controversial. In the case of Delias, the COI gene
shows high definition and resolves the relationships between closely related species.
Müller et al. (2010) showed that it had variable application for delineation of closely
related species of Charaxes (Nymphalidae) in the region but generally resolved
relationships.
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Miller, L.D., Simon, M.J. & Wills, L. (2007) A new species of Delias (Lepidoptera:
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Figure S3 Phylogenetic trees for the COI dataset of selected members of the nigrina
and albertisi species groups. (a) Strict consensus tree of 53 equally parsimonious
trees. Numbers below branches are bootstrap values > 50% for the node to the right.
(b) Fifty percent majority rule consensus phylogram from the Bayesian analysis.
Numbers to the left of nodes are the posterior probabilities of those nodes.
(a)
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(b)