Acanthus ilicifolius L. (Acanthaceae), a new larval host plant for Hypolycaena erylus himavantus Fruhstorfer, 1912 (Lepidoptera: Lycaenidae) from Thailand, with a review on larval host plants of H. erylus and its weaver ant association

Abstract

The mangrove, Acanthus ilicifolius L. (Acanthaceae) is recorded as a new larval host plant for the tropical butterfly Hypolycaena erylus himavantus Fruhstorfer, 1912 in Thailand. Hypolycaeana erylus (Godart, 1824), which is clearly polyphagous, is now known to utilize 25 hosts from 17 different plant families. The larvae of this species at the study site appeared to have an obligate myrmecophilous relationship with the weaver ant Oecophylla smaragdina, as larvae were heavily attended by the ant and no last instar larvae appeared to be unattended. This would appear to be (a) the first record of H. eryclus himavantus feeding as larvae on Acanthus ilicifolius, (b) the first record of a host plant in Thailand for this butterfly species and (c) the first butterfly larvae recorded feeding on foliage of Acanthus ilicifolius. A review of known larval host plants for this thecline butterfly is provided as well as a short discussion of weaver ant association with the larvae of this lycaenid.

Full text

Acanthus ilicifolius L. (Acanthaceae), a new larval host plant for Hypolycaena
erylus himavantus Fruhstorfer, 1912 (Lepidoptera: Lycaenidae) from Thailand,
with a review on larval host plants of H. erylus and its weaver ant association
T.J. Hawkeswood*, K.L. Dunn+ & B. Sommung#
*PO Box 842, Richmond, NSW, 2753, Australia (drtjhawkeswood@gmail.com)
+ kelvyn_dunn@yahoo.com
#Sisaket, Thailand (buppha19741@gmail.com)
Hawkeswood, T.J., Dunn, K.L. & Sommung, B. (2016). Acanthus ilicifolius L. (Acanthaceae), a new larval host plant
for Hypolycaena erylus himavantus Fruhstorfer, 1912 (Lepidoptera: Lycaenidae) from Thailand, with a review on larval
host plants of H. erylus and its weaver ant association. Calodema, 416: 1-8.
Abstract: The mangrove, Acanthus ilicifolius L. (Acanthaceae) is recorded as a new larval host plant for the tropical
butterfly Hypolycaena erylus himavantus Fruhstorfer, 1912 in Thailand. Hypolycaeana erylus (Godart, 1824), which is
clearly polyphagous, is now known to utilize 25 hosts from 17 different plant families. The larvae of this species at the
study site appeared to have an obligate myrmecophilous relationship with the weaver ant Oecophylla smaragdina, as
larvae were heavily attended by the ant and no last instar larvae appeared to be unattended. This would appear to be (a)
the first record of H. eryclus himavantus feeding as larvae on Acanthus ilicifolius, (b) the first record of a host plant in
Thailand for this butterfly species and (c) the first butterfly larvae recorded feeding on foliage of Acanthus ilicifolius. A
review of known larval host plants for this thecline butterfly is provided as well as a short discussion of weaver ant
association with the larvae of this lycaenid.
Introduction
The general biology and host plants of butterflies and moths of Thailand are poorly known apart
from some agriculturally important species (e.g. Ek-Amnuay, 2010). This leaves considerable scope
for the casual naturalist to investigate the biology of various common species of Lepidoptera in this
region of south-east Asia, and to record new ecological information on those study animals selected.
The thecline butterfly, Hypolycaena erylus (Godart, 1824), is a myrmecophilous species based on
observations of its early stages at various locations in Asia (e.g. Jacobsen, 1912; Tokeshi et al.,
2007). Because of this ecological adaptation it would be expected to be polyphagous (see Valentine
& Johnson, 1989) and on account of broader host potentiality it would be anticipated to occur in a
variety of habitats throughout its distribution in order to optimize its use of available resources
where the ants are in occupancy. Indeed, earlier writers have reported that it is a common species
where present and is widespread across its known range in the orient. It occurs from Sikkim through
to Indo-China, and the Philippines, extending through the (largely) Indonesian archipelago to the
island of New Guinea, including Papua New Guinea at its eastern-most edge (Corbet & Pendlebury,
1992; Parsons, 1998). In Malaysia, Corbet & Pendlebury (1992: 320) reported a broad spectrum of
habitat usage, remarking that it can be found in “all types of vegetation from coastal mangrove to
the forested hills” but is rare in the montane areas above 2,000 feet (ca. 660 m). Others have
reported a similar variety of habitats including lowland rainforest and temperate hill forest, at
elevations between sea level and about 1500 metres in India, Papua New Guinea, Thailand and
other countries of SE Asia (Kirton, 2014; Hoskins, 2016).
Taxonomically, the species fits poorly in Hypolycaena C. & R. Felder (Corbet & Pendlebury, 1992),
a genus of some 30 species in the Afrotropical and Indo-Australian regions, with high diversity in
tropical Africa (Parsons, 1998), and so provides a heterogeneous mixture of species that require
reassessment. Corbet & Pendlebury (1992) stated that characteristics of the genitalia of erylus,
along with a sex-brand on the male forewing, align it with the Australian H. phorbas (Fabricius
1793) group which includes a sibling species, H. litoralis Lambkin et al. 2005 from Torres Strait
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(Braby, 2016) and other allied species in Papua New Guinea (Parsons, 1998). Illustrations of set
material of H. erylus are available in various sources (eg. Corbet & Pendlebury, 1992; Parsons,
1998), and those of its close relatives can be sourced in Parsons (1998) and Braby (2016). The
adults photographed (Figs. 1, 2) clearly match H. erylus based on the underwing patterns, and that
species is the only member of the genus in northern Thailand (e.g. Kunte, 2015).
Fig. 1. Adult of Hypolycaena eryclus himavantus resting on foliage of Acanthus ilicifolius L. (Acanthaceae) at
Bangkok, Thailand, July 2016. Note the prickly leaves of A. ilicifolius, which are characteristic of that species. (Photo:
T.J. Hawkeswood & B. Sommung).
Hypolycaena erylus includes about 15 subspecies scattered across its geographic range (Parsons,
1998). There are two subspecies in Thailand; subspecies teatus Fruhstorfer, 1912 is restricted to
southern Thailand where it extends southward through the Malay Peninsula (see Corbet &
Pendlebury, 1992) and the other subspecies, himavantus Fruhstorfer, 1912, occurs broadly through
central and northern Thailand (Inayoshi, 2001). Whether the characteristics of these two subspecies
are well defined or not, or whether transitional populations occur in intervening areas, beyond the
accepted limits of each, is not clear. It is sufficient to suppose that they may exist, and if so, that
assignment of some of these populations to subspecies may be arguable. That said the population in
this report from Bangkok, which is at or near the southern limits of the range of himavantus, has
been associated with that subspecies based on geographic proximity. Without close examination of
the wing patterns, not all of which are visible in the images obtained, a taxonomic placement to this
lower level is not possible other than on circumstantial evidence.
The species is likely to use particular strata in its habitat for surveillance and roosting but fine
details are lacking; for H. phorbas in Papua New Guinea, Parsons (1998) reported that males
occupy perches from 1-3 m above ground and similar behavior may be expected in H. erylus. It has
been reported that both sexes of H. erylus, when not feeding, often perch high up on bushes or on
the lower branches of trees, but they also settle on low herbage and bask on occasions with their
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wings fully outspread (Hoskins, 2016). Typical of many butterflies, one would expect that a wide
basking stance (as reported behavior for the species) would present during cooler conditions, and
when temperatures are higher adults will likely close their wings (see Fig. 1 this paper) to limit
absorption of radiation (Braby, 2016). The species is a frequent visitor to moist spots on forest roads
(Corbet & Pendlebury, 1992) – an accessory feeding habit to flower foraging – where they
presumably seek mineralized nutrients or enrichments of other organic matter, including amino
acids [e.g. see Norris (1936) & Downes (1973) for discussions]; Corbet & Pendlebury (1992) did
not specify the sexes they had noted, but this puddling behavior is usually associated with males
(Downes, 1973).
Fig. 2. A close view of Hypolycaena eryclus himavantus resting on flowers of Acanthus ilicifolius L. at Bangkok,
Thailand, July 2016. (Photo: T.J. Hawkeswood & B. Sommung).
Observations
During two recent visits to the Royal Gardens No. 4 in Bangkok city, Thailand on 5 and 14 July
2016, about 1030-1130 hrs Thailand Time, two of the authors (TJH & BS) observed several
lycaenid butterflies flying around the flowers and leaves at the tops of several plants of Acanthus
ilicifolius L., growing at the edge of a human constructed waterway (Figs. 1 and 2). On close
inspection, several last instar larvae of this butterfly were found at the ends of two leaves; they were
attended by numerous weaver ants, Oecophylla smaragdina (Fabr.) (Hymenoptera: Formicidae)
(Fig. 3). The adult butterflies were very active in full sunshine; they usually spent less than 10
seconds pausing to feed at flowers or perching on flowers and leaves of the host plant. The males
being vigilant and aggressively territorial were easily disturbed; they would rapidly fly out and
patrol the habitat briefly before returning to the host to perch again. On two occasions, it appeared
that females were laying eggs on the younger foliage but a close examination of the leaves failed to
uncover any.
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Fig. 3. Ants attending last instar larvae of Hypolycaena eryclus himavantus on foliage of Acanthus ilicifolius L. at
Bangkok, Thailand, July 2016. (Photo: T.J. Hawkeswood & B. Sommung).
Discussion
The larvae encountered were not able to be collected and reared to adult to confirm the species
involved which undermines the identification rigor to an extent. Notwithstanding this, the
photographs of the larvae (Fig. 3) were closely compared with photos of larvae of several Arhopala
species from Sri Lanka and Australia – a genus that has larvae of similar size to this species and
which can be associated with the same ant species in the Asia-Pacific region – and which
reasonably eliminated them as not likely to be that genus. Secondly, they were compared with
larvae of Hypolycaena phorbas from eastern Australia, and H. erylus from Singapore, Sulawesi and
other locations. The larvae of H. erylus is quite variable but the features of the mature larvae
illustrated (Fig. 3) broadly resemble the green form of the mature larva of H. phorbas from
Australia. They are most similar, though, to those of H. erylus from Singapore, as illustrated by Tan
(2016) and somewhat similar to those from Sulawesi. We could not find any illustrations of the
mature larva of this species from Thailand in order to more suitably compare the larvae with local
material. On the evidence available, one of us (KLD) determined that the larvae almost certainly
belong to the H. phorbas group (as might be anticipated), and hence would be reasonably accepted
as H. erylus by association. Given the variability in the larvae of H. erylus it is likely larvae of
subspecies himavantus would not match exactly with material of those other subspecies,
particularly from populations farther afield. Geographic variation may explain some of the
differences that were noted; the larvae illustrated seem less well marked than those from elsewhere.
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The larvae of H. erylus utilize a broad range of host plants (Tokeshi et al., 2007) and this
acanthaceous mangrove, Acanthus ilicifolius L. now adds to that tally. It provides the first larval
host record (that we know of) for subspecies himavantus in Thailand and, indeed, may be the first
report of larvae of any butterfly species feeding on the foliage of A. ilicifolius. The mangrove is a
prickly shrub (Fig. 1), which extends from India, throughout Malesia to the Solomon Islands, New
Hebrides (Vanuatu) and New Caledonia (Barker, 1986). It favors the river banks, tidal canal sides or
swampy areas in mangrove forests and their vicinity (Barker, 1986; Jafee Bignoli, 2013). Indeed, A.
ilicifolius is a rare plant in Thailand but some colonies are preserved in parks and gardens. In
Malaysia it is similarly rare and has recently been listed under the IUCN as Least Concern [LC -
version 3.1] (Juffe Bignoli, 2013). On account of this plant’s scarcity, and the abundance of the
butterfly, it is probably an infrequently utilized host compared with those other more common
species that may be available to it elsewhere in its range. The mangrove is widely known, however,
due to its medicinal properties, and chemical extracts have been used to treat complaints of
neuralgia and rheumatism, particularly in China (Peng & Long, 2006; Liu & Lin, 2008); the root
when boiled in mustard oil is used for paralysis of limbs; water extracted from the bark is also used
to treat colds and skin allergies.
The lycaenid butterfly, H. erylus is clearly polyphagous, given that no less than 24 plant species
belonging to 16 families have been reported as host plants throughout the Indo-Malesian tropics
(Tokeshi et al., 2007). It appears to have a close association with mangrove communities,
nonetheless. Rau & Murphy (1990) and Murphy (1990) found larvae of Hypolycaena erylus on the
mangrove Rhizophora apiculata Blanco (Rhizophoraceae) in Thailand and Singapore respectively,
and reported extensive damage to the younger leaves attributed to this lycaenid as well as other leaf
feeding insects. Murphy (1990) also listed the following plants as larval hosts for H. erylus:
Rhizophora mucronata Lamk. and Bruguiera cyclindrica (L.) Blanco (Rhizophoraceae), Lumnitzera
racemosa Willd. (Combretaceae) and Scyphiphora hydrophylacea C.F. Gaertn. (Scyphiphoraceae)
[therein cited as “Sypiphora sp.”] as hosts in Singapore. Tokeshi et al. (2007) noted that in Sulawesi
(Indonesia), H. erylus feeds on the leaves of the mangrove apple, Sonneratia alba (Smith)
(Sonneratiaceae). Parsons (1998) also noted that in Papua New Guinea the larvae favor the younger
growth of those hosts utilized on that island. Of the 24 plant species purportedly used (of which we
do not have access to a full list), Tokeshi et al. (2007) mentioned three hosts, all of which are
mangroves: namely, Brugiera cylindrica (L.) Blanco, Rhizophora apiculata Blanco and R.
mucronata Lamk. [Plant authorities inserted by us],; in doing so, they did not provide the literature
sources for those records they listed, nor did they give references for the other unlisted hosts that
made up the tally they reported. Jacobson (1912) listed Meyna spinosa Roxb. ex Link [cited therein
as Vangueria spinosa Hook. f.] (Rubiaceae) as a host in Java, Indonesia. Hoskins (2016) noted that
the eggs of H. erylus are laid on Meyna pubescens (Kurz) Robyns, Meyna spinosa Roxb. ex Link
(Rubiaceae) [therein cited as Vangueria spinosa- recorded first by Jacobsen (1912) - see above] and
Cinnamomum verum J. Presl (Lauraceae) [therein cited as Cinnamomum zeylanicum] (localities not
cited) [Plant authorities inserted by us]. [Oviposition is a good guide as to the plant species utilized,
those rare mistakes aside (Braby, 2015), and so these may prove to be host plants with further
investigation]. Additionally, H. erylus larvae have been recorded feeding on the following plant
species in Singapore: Clerodendrum inerme (L.) Gaertn. (Lamiaceae), Saraca thaipingensis Cantley
ex Prain (Caesalpiniaceae), Hibiscus tiliaceus L. (Malvaceae) [as Talipariti tiliaceum], Ixora
javanica (Blume) DC. (Rubiaceae) and Guioa pleuropteris (Blume) Radlkf. (Sapindaceae)(Tan,
2016) [Plant authorities inserted by us].
Mangrove communities which include adjacent mudflats and saltmarshes are subject to regular tidal
inundation, sometimes with much of the vegetative community being covered by water at high tide.
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For these reasons, mangrove habitats are not a diverse floristic community (Anon., 2016) and
usually contain just a few species of plants specifically adapted to that saline environment. For the
same reason too, nectar sources are often limited in these communities and so those thecline
butterflies (eg. some Hypochrysops species) that are closely linked to hosts in these communities in
Australia for example, are often well adapted to seasonality of foraging at the few resources they
provide; in particular, the adults often target flowers of their own larval hosts for their foraging
requirements as herb-layer flowers are normally unavailable (Hill, 1986). However, H. erylus is not
a specialist species limited to the mangrove community and probably utilises the several mangrove
hosts recorded as additional resources, where available, as part of opportunism and polyphagy
swayed by the ant associations rather than the plant species itself. Supportively, Nagelkerken et al.
(2008) has previously noted that the few mangrove-inhabiting butterflies that can breed in a range
of natural habitats (localities not cited) are those that are either oligophagous (feeding on various
members of single plant lineages) or polyphagous (feeding on many plant groups), such as
Hypolycaena erylus (Lycaenidae) and Polyura schreiber (Nymphalidae). Both of these species,
which have been recorded feeding on one and more mangrove species (Murphy, 1990; this paper)
suitably optimize their survival and dispersal by their use of non-mangrove host plants as well
(Corbet & Pendlebury, 1992). Nonetheless, the H. phorbas group in Australia appears to show a
strong association with mangrove communities (KLD per. obs. and inferentially in McCubbin
(1971) for the mainland; Lambkin et al. (2005) for Torres Strait), despite the many hosts from 18
genera listed by Braby (2016) for H. phorbas in Australia and the two genera listed for H. litoralis
(both mangroves). Notwithstanding this spectrum of usage, their frequent presence in mangroves
(and adjacent littoral scrublands) may be influenced more so by the spatial requirements of the ant
(see Lokkers, 1986) rather than the butterflies’ intentional selection of that habitat.
The weaver ant, Oecophylla smaragdina attends the larvae of most, if not all species in the H.
phorbas group (Braby, 2016; Parsons, 1998). It is almost certainly an obligatory association in all
species in that subgrouping (as has been implied and stated for at least H. phorbas in Australia
(Eastwood & Fraser, 1999 and references therein; Braby, 2000). An obligate relationship, in the
case in the Thailand species, suitably protects the juvenile stages from predators as has been found
to occur in the Australian thecline butterfly, Jalmenus evagoras (Donovan, 1805) (Pierce et al.,
1987; Pierce & Nash, 1999). The first record of an association of H. erylus with weaver ants dates
back to the report by Jacobsen (1912) for the nominate subspecies in north and central Java, in
Indonesia. In Singapore and Thailand, others such as Murphy (1990) and Rau & Murphy (1990)
have noted that H. erylus defoliated Rhizophora apiculata Blanco (Rhizophoraceae) when weaver
ants were present on the host trees; supportively, Parsons (1998) remarked concerning H. phorbas
that ovipositing females ignore potential hosts that are not ant-occupied. In addition, Tokeshi et al.
(2007) reported that younger instars of H. erylus in Sulawesi were generally found in the arboreal
nest of the Oecophylla weaver ants, which may suggest that the survival of young lycaenid larvae is
enhanced by the increased ant guard associated with the close proximity to the nest areas. They
noted too that older instars leave the ant nests and forage openly, and suggested that this might be
because the amount of suitable leaves inside an ant's nest is limited. Nonetheless these larvae are
still aggressively guarded and tended by the weaver ants, which will attack other insects or spiders
wandering on the foliage. Tokeshi et al. (2007) concluded for the Indonesian population they
studied, that whether the lycaenid larvae were inside or outside of an ant's nest, they were
continuously attended by the ants; their evidence suggests that at least for that particular population
of H. erylus, myrmecophily would seem an obligate association, as it is with H. phorbas in
Australia. The observations of heavy ant attendance in Sulawesi and Singapore as well as in
Thailand suggest similarly that myrmecophily is the norm in this butterfly and is likely to be
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obligatory, perhaps with this species of ant only, across the whole range of this butterfly on the
various hosts it uses.
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Date of publication: 10 September 2016
Copyright: T.J. Hawkeswood, K.L. Dunn & B. Sommung
Editor: Dr T.J. Hawkeswood (drtjhawkeswood@gmail.com)
PO Box 842, Richmond, New South Wales, Australia, 2753
(Published as hard paper copy edition as well as electronic pdf)
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