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METAMORPHOSIS
LEPIDOPTERISTS’ SOCIETY OF AFRICA
Volume 25: 82–89
ISSN 1018–6490 (PRINT)
ISSN 2307–5031 (ONLINE)
Applying lepidopteran oviposition science to establish new butterfly larval
host plants from Mpumalanga and Limpopo provinces, South Africa.
Published online: 26 August 2014
Herbert H.H. Otto
P.O. Box 1511, Barberton, 1300.
E-mail: herbertotto@hotmail.com; 24765627@nwu.ac.za
Copyright © Lepidopterists’ Society of Africa
Abstract: This paper reviews lepidopteran oviposition science and descibes the many specific plant and
environmental cues that female butterflies and moths seek prior to oviposition. Single oviposition
observations are looked upon as an inconclusive method of determining larval host plants (LHP)s. Repeat
oviposition and larval feeding observations are therefore presented that provide conclusive evidence for 28
new LHPs for butterfly species from Mpumalanga and Limpopo Provinces, South Africa.
Key words: Oviposition only, oviposition stimuli, oviposition cues, repeat oviposits, LHP recognition and
identification.
Citation: Otto, H.H.H. (2014). Applying lepidopteran oviposition science to establish new butterfly larval host
plants from Mpumalanga and Limpopo provinces, South Africa. Metamorphosis 25: 82–89.
INTRODUCTION
Lepidopteran oviposition is a much researched science
(Nishida, 2005) and females of the butterfly families
Nymphalidae, Pieridae and Papilionidae have been
found to follow definite sets of host finding cues
(Honda & Nishida, 1999). They rely on several senses
including vision (Nishida, 2005), specifically colour
reliance (Bernard & Remington, 1991) and leaf shape
(Edge, 2008) or polarization (Schoonhoven et al.,
2005). The olfactory senses (Nishida, 2005) can detect
volatiles that assist with larval host-plant (LHP)
recognition (Visser, 1986; Honda et al., 1997; Honda et
al., 2001; Schoonhoven et al., 2005) and orientation
(Honda, 1995), whereas the gustatory senses also play
a part (Ozaki et al., 2011; Schoonhoven et al., 2005;
Nishida, 2005; Ryuda et al., 2013) in detecting less or
non-volatile secondary metabolites (Honda, 1995).
Tactile senses are also used (Feeny et al., 1983;
Nishida, 1995; Roessingh et al., 1991; Ryuda et al.,
2013). Environmental cues include plant microclimate
(Schoonhoven et al., 2005), a preference for younger
leaves (Murugan et al., 1997) or plant size & secondary
chemistry (Reudler Talsma et al., 2008; Bowers, 1984).
In some situations an oviposition stimulant binding
protein (OSBP) may be necessary (Tsuchihara et al.,
2009) for confirmation of the correct LHP. Other
influencing factors include the absence of predators, an
‘enemy free space’ (Schoonhoven et al., 2005),
temperature and season (Schoonhoven et al., 2005) and
deterrents (Murawski, 1993; Paré & Tumlinson, 1999)
or which plants surround the host plant (Metspalu et al.,
2003) before the female will be placated and satisfied
to oviposit. Some butterflies are induced to oviposit by
one host-specific compound, while other butterflies
need multiple compounds (Honda, 1995; Nishida,
1995; Honda & Nishida, 1999; Ryuda et al., 2013) or
stimulants (Ryuda et al., 2013; Nishida, 1995). Certain
compounds may also serve as oviposition deterrents
(Ono et al., 2004). Papilio polyxenes detects stimulants
and deterrents that lead to LHP recognition through
tarsal chemosensilla (Roessingh et al, 1991; Ryuda et
al., 2013). The aforementioned sensory and
interspecific chemical communications are also utilised
by Lycaenidae but myrmecophilous species may also
need the correct host-ants to be present (Heath, 1998,
2014; Pierce et al., 2002); Heath et al., 2008. Lycaenid
larvae also interact intraspecifically and with the host
ants using chemical communication (Henning, 1983).
Butterflies may even ‘medicate’ their young by
choosing a specific LHP to rid their young of a virus
(Lefèvre et al., 2010; Schoonhoven et al., 2005: 17) or
larvae may self-medicate by taking in Pyrrolizidine
Alkaloids (PA) to rid them of Tachinid fly parasites
(Singer et al., 2009) while some Danaid females use
PAs to find their LHPs (Honda et al., 1997). During
winter or times of drought some butterflies may lay
their eggs on plants or objects near the valid LHP
(Mensah & Kudom, 2012) but mostly they will lay their
eggs on a true LHP, when it is available.
MATERIALS & METHODS
Ovipositing females were observed and recorded at
various locations as set out in Tables 2 & 3. Gravid
female butterflies which display a fluttering flight
pattern around a potential LHP provide an indication
that the female is about to oviposit, and she may often
rest or nectar at the same plant. Ova and larvae were
usually collected and attempts made to breed them
through to adult imagos in plastic containers,
Received: 13 July 2014
Accepted: 26 August 2014
Copyright: This work is licensed under the Creative Commons
Attribution-NonCommercial-NoDerivs 3.0 Unported License.
To view a copy of this license, send a letter to Creative
Commons, Second Street, Suite 300, San Francisco, California,
94105, USA, or visit: http://creative commons.org/licenses/by-
nc-nd/3.0/
Otto/ Metamorphosis 25: 82–89 83
replenishing fresh foliage daily. A Canon 650D with
60mm Macrolens was used for photographs. LHP
identification was verified by Mr Doug McMurtry
(Hortus Siccus Mcmurtrianus = HSMC) and for the
genus Indigofera by Dr. Brian Schrire (Kew).
RESULTS
Table 1 presents observations made on two abundant
polyphagous butterfly species which utilise several
LHP families and genera. Table 2 presents 15 observed
repeat oviposits for nine butterfly species, which serve
as confirmation of a LHP. In two cases these oviposits
were repeated on three different occasions on different
dates. The repeat oviposits of C. forestan forestan on
Indigofera arrecta Hochst. ex A. Rich. were further
confirmed by subsequent rearing of two larvae on this
plant to successful eclosion of the adult butterflies.
Table 3 presents 28 new LHP records, recorded for 12
different butterfly taxa, all of which butterflies are
considered to be polyphagous. Plate 1 (Figs 1–10)
contains a selection of oviposition, larval feeding and
pupal photographs.
Indigofera arrecta is a LHP for four butterflies in the
Barberton area alone – Anthene definita definita (new
plant genus and species record). Coeliades forestan,
Chilades trochylus and Leptotes sp. (of the pirithous
group) are all new plant species records.
The search for new LHPs has also benefited the
botanical community, since the oviposit of Lampides
boeticus on Crotalaria schlechterii Baker f. served to
identify four individual plants of this rare plant species
in the unlikely vicinity of a mine tailings site near
Barberton.
DISCUSSION
The study of oviposition is an empirical science which
enables us to understand how gravid female butterflies
rely on sensory and phytochemical cues to guide them
to the correct host plant. Plant buds produce fewer
volatiles than plant corollas (Irwin & Dorsett, 2002)
therefore butterfly females that oviposit on buds and
their subsequent larvae feeding on buds would be
attracted to these specific volatiles.
Studies done on the Gypsy Moth, Lymantria dispar
indicate that these moths detect and steer around virus-
contaminated leaves (Parker et al., 2010) as confirmed
by field observations of similar butterfly behaviour.
Avoidance of plants which have been heavily browsed
by other insects has also been observed. Butterflies may
also oviposit on various plants which are in close
proximity to the true LHP, perhaps due to a lack of a
suitable LHP during times of drought, and the larvae
move to the true LHP on emergence (Mensah &
Kudom, 2012). A similar spring observation was made
of a gravid Papilio demodocus demodocus at Crocodile
Gorge laying eggs on other non-LHPs 30cm away from
the correct diminutive LHP, Sneezewood, Ptaeroxylon
obliquum. During the July 2013, Low’s Creek Colotis
evagore antigone lay eggs on dry grass stems close to a
Maerua species. Perhaps butterflies that lay eggs on
winter-stressed LHPs and create further duress for the
plant limit the survival of their progeny and have an
evolutionary disadvantage.
According to J.H. Fabre ovipositing females seem to
have a ‘botanical instinct’ (Schoonhoven et al., 2005:
15). For example, the genetic analysis and insect
association of two Populus species had a 98%
correlation (Schoonhoven et al., 2005: 16). Perhaps
oligophagous or polyphagous butterfly species like
Pieris brassicae select host plants with phytochemicals
in common such as glucosinulates (Schoonhoven et al.,
2005: 7). Although a specific phytophagous species is
known to be polyphagous, the insect may still have a
higher preference for certain species within its edible
range (Schoonhoven et al., 2005: 9), as can be seen in
Table 1. Leptotes in the pirithous-group prefer LHPs in
the family Fabaceae, while Hypolycaena philippus
philippus prefers Rubiaceae. Also larvae performed
best on the particular LHP the adult female selected
(Schoonhoven et al., 2005: 10).
It is hypothesised that polyphagous butterflies
(generalists) have the ability to sort though many
signals while monophagous butterflies (specialists) are
better adapted to distinguish between subtle differences
in specific signals (Berenbaum & Feeny, 2008). The
same authors also observed that poor quality leaves did
not impair the larval growth of generalists. Ovipositing
female African Monarchs, Danaus chrysippus
preferred younger leaves of Calotropis gigantea, which
increased the larvae’s growth rates, quickened the
larval development, improved fecundity and
maximised the longevity whereas the older leaves
containing less water and nutrients had an increased
cardiac glycoside concentration (Murugan et al., 1997).
It has been noted that phytophagous species with a
limited host plant spectrum prefer young leaves while
the caterpillars of polyphagous species choose mature
leaves due to the higher level of toxins in younger
leaves (Schoonhoven et al., 2005: 8). The larvae of
some moths can distinguish between leaflets from the
base, central or terminal areas and avoid base leaflets
(Schoonhoven et al., 2005: 11).
At close range the female Zebra Swallowtail, Eurytides
marcellus is very attracted to host plant volatiles and
having found the host plant she relies on vision to
determine the young/mature leaf ratio and whether leaf
buds are present (Damman & Feeny, 1988). Regarding
vision, Papilio species are reported to be able to
distinguish between UV, purple, blue, green, red and
broadband (Arikawa, 2003).
Oviposition stimuli can be non-volatile taste stimulants
(Ryuda et al., 2013), so that the butterfly must settle on
the leaf of the prospective LHP to establish whether it
is a LHP or not. On settling she drums her tarsi and the
chemosensilla to determine if the plant has the
necessary phytochemicals. Within the Papilionidae,
Papilio xuthus needs a concoction with a specific ratio
of 10 phytochemicals; P. polytes requires 5 different
stimulants and P. protenor needs 6 plant chemicals
(Ryuda et al., 2013). P. xuthus has three different
sensilla found in the tarsomeres, each with a different
sensory function (Ryuda et al., 2013).
Certain plants attract Asian Danaid butterfly females,
specifically Ideopsis similis, which needs a definite
Otto/ Metamorphosis 25: 82–89 84
combination and exact quantities of the 12 studied
phenanthroindolizidine alkaloids emitted by its LHP,
Tylophora tanakae, to lay her eggs on. When this
perfect alkaloid ratio was separated into its individual
alkaloids, the female laid fewer eggs. Five of the 12
individual alkaloids examined stimulated oviposition,
while 3 of the 5 specific alkaloids served to assist with
host plant recognition (Honda et al., 2001). Another
interesting fact is that the butterfly Ideopsis similis is
cytotoxic due to the presence of these compounds
assimilated from the leaves of Tylophora tanakae
(Koyama et al., 1999). In South Africa Tylophora also
serves as a LHP for Amauris and it may thus be inferred
that certain alkaloids will attract the female butterflies
and others will stimulate her to oviposit. Some
Passifloraceae plants from Central and South America
repel oviposition by developing 2 yellow knob-like
growths at the base of their leaves, resembling
Heliconid eggs and deterring any other butterfly
oviposits. Other Passion Vines develop sugary leaf
secretions to attract wasps and ants – natural butterfly
egg and larvae enemies (Murawski, 1993). Cabbage
Whites, Pieris brassicae ovipositing on Arabidopsis
plants show that the plant’s whole-genome microarray
gene expression changes within three days to rid the
plant of the eggs with the cells beneath the egg actually
dying and forming calluses (Little et al., 2007). The
plant also emits volatiles from insect-damaged leaves
to attract larval enemies like parasitoid wasps (Paré &
Tumlinson, 1999).
American Monarch (Danaus plexippus) butterfly
larvae feeding on Tropical Milkweed had reduced
parasitoid growth, infection and disease while
parasitoid infected females preferred the more toxic
Tropical Milkweed to the less toxic Swamp Milkweed,
indicating that the females ‘doctored’ their young by
choosing a LHP that is more medicinal (Lefèvre et al.,
2010). Some arctiids feed on plants with decreased
nutritional value yet these plants supply the insects with
chemicals that ensure the survival from parasitisation
(Schoonhoven et al., 2005: 17).
It is hypothesised that common butterflies lay their eggs
in clusters and this certainly holds true for some
nymphalids, pierids and acraeids while rarer butterflies
lay their eggs singly (Stamp, 1980). Acraea encedon
encedon may lay several eggs in clusters to increase
copulation success by having the few remaining males
in close proximity to the females because males are
lethally parasitized by Wolburgia bacteria (Jiggins et
al., 1998). Lycaeides melissa samuelis from North
America on the other hand has two different strategies;
the first brood before summer lays eggs singly, while
the second brood that overwinters, are laid in batches.
The study has two hypotheses; due to the
environmental conditions (summer drought) first brood
eggs laid singly are distributed over a greater area to
support survival, while second brood egg clusters of 1–
6 allows at least one larva to survive (Pickens & Root,
2008). Thus if eggs are laid in clusters during times of
scarcity, the survival of at least one larva is assured,
which could be as a result of cannibalism (Edge & van
Hamburg 2009). Many ovipositing examples
mentioned in this paper do not include Lycaenidae, yet
it may be extrapolated that species of the Lycaenidae
would also utilise several of the ovipositing cues (e.g.
Heath & Gardiner, 2009) used by butterflies of the
Papilionidae, Pieridae and Nymphalidae while their
sensory perception may even be heightened if
myrmecophagous Lyceanids are considered.
Certain butterflies also rely on LHP stimuli like
stachydrine, a secondary compound found in Citrus for
the Asian Swallowtail Papilio xuthus which is essential
for the larvae to feed (Murata et al., 2011; Ryuda et al.,
2013). Pyrrolizidine alkaloids (PAs) allow female
Danaid butterflies to recognise their LHPs (Honda et
al., 1997; Hartmann et al., 2005), serve as precursors
for sexual pheromones (Schulz et al., 2004; Trigo,
2011) and PAs sequestered from plants are utilized as
defensive substances (Orr et al., 1996; Nishida, 2002).
With Nymphalidae the larvae of the North American
Buckeye butterfly needs the secondary compound
iridoid glycosides which supports the growth of the
larvae (Bowers, 1984). Some Papilionidae like the
Black Swallowtails, Papilio polyxenes from Asia have
tarsal brushes and these tactile trademarks have to be
satisfied by the LHP extracts before egg-laying is
stimulated (Roessingh et al., 1991; Ryuda et al., 2013).
In the female Pierids of the Sulphur Butterfly, Colias
erate, D-pinitol as a stimulant has to be present before
the female will oviposit (Honda et al., 2012).
CONCLUSIONS
Insects ‘sense’ the correct LHP by different methods
including; visual, olfactory (Nishida, 2005),
mechanical and gustatory (Nishida, 2005; Ryuda et al.,
2013) stimuli.
A butterfly will oviposit on an alternative plant or
object other than its true LHP under these
circumstances:
1) During dry seasons the gravid female butterfly will
lay near the LHP (Mensah & Kudom, 2012) for the
emerging larva to move to the true LHP.
2) If the LHP has been oviposited on but more
suitable LHPs are rare in the local vicinity the
female may not want to overburden the plant with
too many larvae, thus laying on plants nearby
relying on the principle of survival of the fittest.
3) When the LHPs present have been stripped by
gregarious herbivores like moth larvae, the
butterfly larvae laid nearby may emerge in time for
the new flush on the LHP after the adult moths
have emerged.
4) Some Satyrinid female members of the Tribe
dirinii scatter their eggs while in flight.
With such complexity present in butterfly behaviour, it
must be apparent that butterflies do not randomly lay
eggs.
ACKNOWLEDGEMENTS
Thank you to Mr. Doug McMurtry who imparted his
invaluable plant knowledge and willingness to always
assist with identifying plants while safekeeping
vouchers in his private herbarium using his
considerable inspirational ease of finding and concisely
expressing the precise text and phrases. Dr. Brian
Otto/ Metamorphosis 25: 82–89 85
Schrire graciously assisted with expedient
identification of Indigofera. Thank you to Ian and Lize
Otto for their accommodation, hospitality, keen interest
and support. I am indebted to Dillan Hesselman and
Juan Joubert for dedicated field trip accompaniment to
find new LHPs and larvae. Thanks also to Ian, Tertia
and Allison Sharp for sharing their remarkable work of
locating and breeding butterflies on new LHPs in the
arid area of Hoedspruit.
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responses to phytochemicals from host- and non-
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495.
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TANIMURA, T. & OZAKI, K. 2013. Gustatory
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Otto/ Metamorphosis 25: 82–89 87
Table 1 – Two abundant polyphagous butterfly species which utilise several LHP families and genera
BUTTERFLY FAMILY,
TAXON
PLANT FAMILY PLANT TAXON
LYCAENIDAE
Leptotes sp. in pirithous group BIGNONIACEAE Tecomaria
FABACEAE Abrus, Acacia, Burkea, Indigofera, Mundulea, Neonotonia,
Rhynchosia, Sesbania & Vigna
PLUMBAGINACEAE Plumbago
VERBENACEAE Lantana
Hypolycaena philippus philippus CELASTRACEAE Gymnosporia
CONVOLVULACEAE Cuscuta
LORANTHACEAE Erianthemum
OLACACEAE Ximenia
RUBIACEAE Fadogia, Kraussia, Pachystigma, Vangueria
SAPINDACEAE Deinbollia
VERBENACEAE Clerodendrum
Table 2 – Repeat oviposition observations, which verify LHP records, with butterfly species, LHPs, plant
family, location, observer(s) and dates.
FAMILY,
TAXON
LHP PLANT FAMILY LOCATION, OBSERVER
(H.H.H. Otto unless mentioned)
DATES
HESPERIIDAE
Coeliades forestan
forestan
Indigofera arrecta Hochst.
Ex A. Rich.
FABACEAE Barberton (Litchie Farm Rd.)
Barberton (Litchie Farm Rd.)
15.ii.2014
23.iii.2014
LYCAENIDAE
Azanus jesous
jesous
Dichrostachys cinerea (L.)
Wight & Arn.
FABACEAE Letsitele
Barberton (Litchie Farm Rd.)
20.xii.2013
15.ii.2014
Chilades trochylus Indigofera vicioides Jaub &
Spach.var. rogersii (R.E.
Fr.) J.B. Gillett
FABACEAE Low’s Creek (Three Sisters Farm)
Low’s Creek (Three Sisters Farm)
Skukuza
16.ii.2013
4.iv.2013
29.iii.2013
Leptotes sp. in
pirithous group
Abrus laevigata E. Mey. FABACEAE Crocodile Gorge
Barberton (Litchie Farm Rd.)
i.2008
15.ii.2014
Leptotes sp. in
pirithous group
Acacia nigrescens Oliv. FABACEAE Shingwedzi
Low’s Creek
xii. 2011
7.vii.2013
Leptotes spe. in
pirithous group
Desmodium tortuosum
(Sw.) DC.
FABACEAE Barberton (Moody’s Rd.)
Barberton (Moody’s Rd.)
1.v.14
1.v.14
Leptotes sp. in
pirithous group
Neonotonia wightii (AM.)
J.A. Lackey
FABACEAE Low’s Creek (Three Sisters Farm)
(plains)
Low’s Creek (Three Sisters Farm)
(valley)
i.2009
ii.2011
Zizeeria knysna Mollugo nudicaulis Lam. MOLLUGINACEAE Low’s Creek (Three Sisters Farm)
Barberton (Ameide Rd.)
12.xii.2012
12.xii.2013
Zizeeria knysna Indigofera spicata Forssk. FABACEAE Barberton (Town)
Barberton (Town)
5.ii.2014
16.v.2014
Zizula hylax Ruellia patula Jacq. ACANTHACEAE Letsitele
Hoedspruit (Sharp, I)
20.xii.2013
xii.2013
Zizula hylax Ruellia malacophylla C.B.
Clarke
ACANTHACEAE Berg-en-Dal
Blyde River, Hoedspruit
30.xi.2013
14.xii.2013
PIERIDAE
Eurema hecabe
solifera
Acacia ataxacantha DC FABACEAE Nelspruit (Karino)
Barberton (Litchie Farm Rd.)
6.iv.2014
27.iv.2014
Eurema hecabe
solifera
Sesbania bispinosa (Jacq.)
W.Wight var. bispinosa
FABACAEA Marloth Park
Barberton (Litchie Farm Rd.)
Barberton (Litchie Farm Rd.)
iii.2010
15.ii.2014
29.iii.2014
Belenois aurota
aurota
Maerua rosmarinoides Pax CAPPARIDACEAE Barberton
Hoedspruit (Sharp, I.)
Barberton
xi. 2010
xi.2012
i.2013
Colotis evenina Maerua parvifolia (Sond.)
Gilg & Gilg-Ben.
CAPPARIDACEAE Shibavantsengele (KNP)
Low’s Creek (Three Sisters Farm)
xii. 2011
x.2012
Otto/ Metamorphosis 25: 82–89 88
Table 3 – New butterfly LHP records, with butterfly species, LHPs, plant family, location, observer, dates and
types of observation.
FAMILY,
TAXON
LHP PLANT
FAMILY
LOCATION, OBSERVER
(H.H.H. Otto unless mentioned)
DATES TYPE
HESPERIIDAE
Coeliades
forestan forestan
Indigofera arrecta Hochst. Ex
A. Rich.
FABACEAE Barberton (Litchie Farm Rd.) 15.ii.2014
23.iii.2014
11-12.iv.2014
12.v.2014
Oviposition
Oviposition
Larva
Larva
LYCAENIDAE
Azanus jesous
jesous
Dichrostachys cinerea (L.)
Wight & Arn.
FABACEAE Letsitele
Barberton (Litchie Farm Rd.)
18.xii.2013
15.ii.2014
Oviposition
Oviposition
Anthene definita
definita
Indigofera arrecta Hochst. Ex
A. Rich.
FABACEAE Barberton (Litchie Farm Rd.) 9-21.iii.2014
29.iii-5.iv. 2014
Bred from
final instar.
Larva died
Cacyreus lingeus Plectranthus fruticosus L’Her. LAMIACEAE Barberton (Upper Rimer’s Creek
Rd.)
10.v.2014 Oviposition
Cacyreus lingeus Thorncroftia lotterii Edwards
& McMurtry
LAMIACEAE Barberton (Mountainlands) 10.v.2014 Oviposition
Chilades
trochylus
Indigofera arrecta Hochst. Ex
A. Rich.
FABACEAE Barberton (Moody’s Rd.) 7.v.2013 Oviposition
Chilades
trochylus
Indigofera schimperi Jaub. &
Spach var. schimperi
FABACEAE Letsitele 16.xii.2013 Oviposition
Chilades
trochylus
Indigofera spicata Forssk. FABACEAE Barberton (Litchie Farm Rd.) 30.iii.2014 Oviposition
Cupidopsis
jobates jobates
Rhynchosia minima (L.) DC.
var. minima
FABACEAE Crocodile Gorge 1.i.2014 Oviposition
Lampides
boeticus
Crotalaria lanceolata E. Mey. FABACEAE Barberton (Litchie Farm Rd.) 27.iv.2014 Oviposition
Lampides
boeticus
Crotalaria schlechterii Baker f. FABACEAE Barberton (Litchie Farm Rd.) 23.iii.2014 Oviposition
Lampides
boeticus
Sesbania bispinosa (Jacq.)
W.Wight var. bispinosa
FABACEAE Barberton (Litchie Farm Rd.) 30.iii.2014 Oviposition
Leptotes sp. in
pirithous group
Indigofera arrecta Hochst. Ex
A. Rich.
FABACEAE Barberton (Litchie Farm Rd.) 30.iii.2014 Oviposition
Leptotes sp. in
pirithous group
Indigofera galpinii N.E. Br FABACEAE Barberton (Ameide Rd.) 12.xii.2013 Oviposition
Leptotes sp. in
pirithous group
Desmodium tortuosum (Sw.)
DC.
FABACEAE Barberton (Moody’s Rd.) 1.v.2014 Oviposition
Zizula hylax Ruellia malacophylla C.B.
Clarke
ACANTHA-
CEAE
Berg-en-Dal (KNP) Hoedspruit
(Moriah farm, Blyde River)
xii.2013
14.xii.2013
Oviposition
Oviposition
Zizula hylax Ruellia patula Jacq. ACANTHA-
CEAE
Hoedspruit (Indabushy)
Letsitele (Sharp, I. T. & A.)
xii.2013
20.xii.2013
Bred
Oviposition
NYMPHALIDAE
Danaus
chrysippus
orientis
Ceropegia carnosa E. Mey. APOCYNA-
CEAE
Whyte Thorne, Nelspruit 15.iii.2014
15.iii.2014
Oviposition
Feeding
larvae
Precis tugela
tugela
Plectranthus fruticosus L’Her. LAMIACEAE Barberton (Litchie Farm Rd.) 29.iii-15/17.iv.
2014
Bred
Precis tugela
tugela
Plectranthus ecklonii Benth. LAMIACEAE Barberton (Litchie Farm Rd.) 29.iii-22.iv. 2014 Bred
Precis tugela
tugela
Plectranthus hadiensis
(Forssk.) Schweinf. ex Spreng.
LAMIACEAE Nelspruit (Whyte Thorne) 5-30.iv. 2014 Bred
Precis tugela
tugela
Plectranthus laxiflorus Benth. LAMIACEAE Barberton (Litchie Farm Rd.) 29.iii-26.iv. 2014 Bred
Precis tugela
tugela
Plectranthus verticillatus (L.f.)
Druce
LAMIACEAE Barberton (Litchie Farm Rd.) 29.iii-25.iv. 2014 Bred
Precis tugela
tugela
Pycnostachys urticifolia Hook. LAMIACEAE Barberton (Litchie Farm Rd.) 29.iii-22.iv. 2014 Bred
PIERIDAE
Eurema hecabe
solifera
Acacia ataxacantha DC FABACEAE Nelspruit (Karino)
Barberton (Litchie Farm Rd.)
6.iv.2014
27.iv.2014
Oviposition
Oviposition
Eurema hecabe
solifera
Dichrostachys cinerea (L.)
Wight & Arn. subsp. nyassae
FABACEAE Barberton (Litchie Farm Rd.) 12.iv- 3.v.2014
(larva died)
Oviposition
Larva
Eurema hecabe
solifera
Sesbania sesban (L.) Merr.
ssp. sesban var. nubica
FABACEAE Crocodile Gorge ii.2011 Bred from
final instar
Eurema hecabe
solifera
Sesbania bispinosa (Jacq.)
W.Wight var. bispinosa
FABACEAE Marloth Park
Barberton (Litchie Farm Rd.)
iii.2010
15.ii.2014
29.iii.2014
Oviposition
Oviposition
Oviposition
Otto/ Metamorphosis 25: 82–89 89
PLATE 1
Figure 1 – Coeliades forestan egg on Indigofera Figure 2 – Coeliades f. forestan larva on
arrecta Hochst. Ex A. Rich., Barberton (iii.2014) Indigofera arrecta Hochst. Ex A. Rich.,
Barberton (iv.2014)
Figure 3 – Anthene d. definita larva on Indigofera Figure 4 – Anthene d. definita pre-pupa.
arrecta Hochst. Ex A. Rich. Barberton. (iii.2014) Barberton. (iii.2014)
Figure 5 – Anthene d. definita pupa about to Figure 6 – Anthene d. definita adult female,
eclose. Barberton. (iii.2014) freshly eclosed. Barberton. (iii.2014)
Figure 7 – Danaus chrysippus orientis larva on Figure 8 – Chilades trochylus egg on Indigofera
Ceropegia carnosa E.Mey. Whyte Thorne, spicata Forssk. (iii.2014)
Nelspruit. (iii.2014)
Figure 9 – Leptotes sp. ovipositing on Figure 10 – Leptotes sp. egg on Desmodium
Desmodium tortuosum (Sw.) DC. (iv.2014) tortuosum (Sw.) DC. (iv.2014)

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2014.08.26 Metamorphosis 25_82-89 Otto Final

  • 1. METAMORPHOSIS LEPIDOPTERISTS’ SOCIETY OF AFRICA Volume 25: 82–89 ISSN 1018–6490 (PRINT) ISSN 2307–5031 (ONLINE) Applying lepidopteran oviposition science to establish new butterfly larval host plants from Mpumalanga and Limpopo provinces, South Africa. Published online: 26 August 2014 Herbert H.H. Otto P.O. Box 1511, Barberton, 1300. E-mail: herbertotto@hotmail.com; 24765627@nwu.ac.za Copyright © Lepidopterists’ Society of Africa Abstract: This paper reviews lepidopteran oviposition science and descibes the many specific plant and environmental cues that female butterflies and moths seek prior to oviposition. Single oviposition observations are looked upon as an inconclusive method of determining larval host plants (LHP)s. Repeat oviposition and larval feeding observations are therefore presented that provide conclusive evidence for 28 new LHPs for butterfly species from Mpumalanga and Limpopo Provinces, South Africa. Key words: Oviposition only, oviposition stimuli, oviposition cues, repeat oviposits, LHP recognition and identification. Citation: Otto, H.H.H. (2014). Applying lepidopteran oviposition science to establish new butterfly larval host plants from Mpumalanga and Limpopo provinces, South Africa. Metamorphosis 25: 82–89. INTRODUCTION Lepidopteran oviposition is a much researched science (Nishida, 2005) and females of the butterfly families Nymphalidae, Pieridae and Papilionidae have been found to follow definite sets of host finding cues (Honda & Nishida, 1999). They rely on several senses including vision (Nishida, 2005), specifically colour reliance (Bernard & Remington, 1991) and leaf shape (Edge, 2008) or polarization (Schoonhoven et al., 2005). The olfactory senses (Nishida, 2005) can detect volatiles that assist with larval host-plant (LHP) recognition (Visser, 1986; Honda et al., 1997; Honda et al., 2001; Schoonhoven et al., 2005) and orientation (Honda, 1995), whereas the gustatory senses also play a part (Ozaki et al., 2011; Schoonhoven et al., 2005; Nishida, 2005; Ryuda et al., 2013) in detecting less or non-volatile secondary metabolites (Honda, 1995). Tactile senses are also used (Feeny et al., 1983; Nishida, 1995; Roessingh et al., 1991; Ryuda et al., 2013). Environmental cues include plant microclimate (Schoonhoven et al., 2005), a preference for younger leaves (Murugan et al., 1997) or plant size & secondary chemistry (Reudler Talsma et al., 2008; Bowers, 1984). In some situations an oviposition stimulant binding protein (OSBP) may be necessary (Tsuchihara et al., 2009) for confirmation of the correct LHP. Other influencing factors include the absence of predators, an ‘enemy free space’ (Schoonhoven et al., 2005), temperature and season (Schoonhoven et al., 2005) and deterrents (Murawski, 1993; Paré & Tumlinson, 1999) or which plants surround the host plant (Metspalu et al., 2003) before the female will be placated and satisfied to oviposit. Some butterflies are induced to oviposit by one host-specific compound, while other butterflies need multiple compounds (Honda, 1995; Nishida, 1995; Honda & Nishida, 1999; Ryuda et al., 2013) or stimulants (Ryuda et al., 2013; Nishida, 1995). Certain compounds may also serve as oviposition deterrents (Ono et al., 2004). Papilio polyxenes detects stimulants and deterrents that lead to LHP recognition through tarsal chemosensilla (Roessingh et al, 1991; Ryuda et al., 2013). The aforementioned sensory and interspecific chemical communications are also utilised by Lycaenidae but myrmecophilous species may also need the correct host-ants to be present (Heath, 1998, 2014; Pierce et al., 2002); Heath et al., 2008. Lycaenid larvae also interact intraspecifically and with the host ants using chemical communication (Henning, 1983). Butterflies may even ‘medicate’ their young by choosing a specific LHP to rid their young of a virus (Lefèvre et al., 2010; Schoonhoven et al., 2005: 17) or larvae may self-medicate by taking in Pyrrolizidine Alkaloids (PA) to rid them of Tachinid fly parasites (Singer et al., 2009) while some Danaid females use PAs to find their LHPs (Honda et al., 1997). During winter or times of drought some butterflies may lay their eggs on plants or objects near the valid LHP (Mensah & Kudom, 2012) but mostly they will lay their eggs on a true LHP, when it is available. MATERIALS & METHODS Ovipositing females were observed and recorded at various locations as set out in Tables 2 & 3. Gravid female butterflies which display a fluttering flight pattern around a potential LHP provide an indication that the female is about to oviposit, and she may often rest or nectar at the same plant. Ova and larvae were usually collected and attempts made to breed them through to adult imagos in plastic containers, Received: 13 July 2014 Accepted: 26 August 2014 Copyright: This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 Unported License. To view a copy of this license, send a letter to Creative Commons, Second Street, Suite 300, San Francisco, California, 94105, USA, or visit: http://creative commons.org/licenses/by- nc-nd/3.0/
  • 2. Otto/ Metamorphosis 25: 82–89 83 replenishing fresh foliage daily. A Canon 650D with 60mm Macrolens was used for photographs. LHP identification was verified by Mr Doug McMurtry (Hortus Siccus Mcmurtrianus = HSMC) and for the genus Indigofera by Dr. Brian Schrire (Kew). RESULTS Table 1 presents observations made on two abundant polyphagous butterfly species which utilise several LHP families and genera. Table 2 presents 15 observed repeat oviposits for nine butterfly species, which serve as confirmation of a LHP. In two cases these oviposits were repeated on three different occasions on different dates. The repeat oviposits of C. forestan forestan on Indigofera arrecta Hochst. ex A. Rich. were further confirmed by subsequent rearing of two larvae on this plant to successful eclosion of the adult butterflies. Table 3 presents 28 new LHP records, recorded for 12 different butterfly taxa, all of which butterflies are considered to be polyphagous. Plate 1 (Figs 1–10) contains a selection of oviposition, larval feeding and pupal photographs. Indigofera arrecta is a LHP for four butterflies in the Barberton area alone – Anthene definita definita (new plant genus and species record). Coeliades forestan, Chilades trochylus and Leptotes sp. (of the pirithous group) are all new plant species records. The search for new LHPs has also benefited the botanical community, since the oviposit of Lampides boeticus on Crotalaria schlechterii Baker f. served to identify four individual plants of this rare plant species in the unlikely vicinity of a mine tailings site near Barberton. DISCUSSION The study of oviposition is an empirical science which enables us to understand how gravid female butterflies rely on sensory and phytochemical cues to guide them to the correct host plant. Plant buds produce fewer volatiles than plant corollas (Irwin & Dorsett, 2002) therefore butterfly females that oviposit on buds and their subsequent larvae feeding on buds would be attracted to these specific volatiles. Studies done on the Gypsy Moth, Lymantria dispar indicate that these moths detect and steer around virus- contaminated leaves (Parker et al., 2010) as confirmed by field observations of similar butterfly behaviour. Avoidance of plants which have been heavily browsed by other insects has also been observed. Butterflies may also oviposit on various plants which are in close proximity to the true LHP, perhaps due to a lack of a suitable LHP during times of drought, and the larvae move to the true LHP on emergence (Mensah & Kudom, 2012). A similar spring observation was made of a gravid Papilio demodocus demodocus at Crocodile Gorge laying eggs on other non-LHPs 30cm away from the correct diminutive LHP, Sneezewood, Ptaeroxylon obliquum. During the July 2013, Low’s Creek Colotis evagore antigone lay eggs on dry grass stems close to a Maerua species. Perhaps butterflies that lay eggs on winter-stressed LHPs and create further duress for the plant limit the survival of their progeny and have an evolutionary disadvantage. According to J.H. Fabre ovipositing females seem to have a ‘botanical instinct’ (Schoonhoven et al., 2005: 15). For example, the genetic analysis and insect association of two Populus species had a 98% correlation (Schoonhoven et al., 2005: 16). Perhaps oligophagous or polyphagous butterfly species like Pieris brassicae select host plants with phytochemicals in common such as glucosinulates (Schoonhoven et al., 2005: 7). Although a specific phytophagous species is known to be polyphagous, the insect may still have a higher preference for certain species within its edible range (Schoonhoven et al., 2005: 9), as can be seen in Table 1. Leptotes in the pirithous-group prefer LHPs in the family Fabaceae, while Hypolycaena philippus philippus prefers Rubiaceae. Also larvae performed best on the particular LHP the adult female selected (Schoonhoven et al., 2005: 10). It is hypothesised that polyphagous butterflies (generalists) have the ability to sort though many signals while monophagous butterflies (specialists) are better adapted to distinguish between subtle differences in specific signals (Berenbaum & Feeny, 2008). The same authors also observed that poor quality leaves did not impair the larval growth of generalists. Ovipositing female African Monarchs, Danaus chrysippus preferred younger leaves of Calotropis gigantea, which increased the larvae’s growth rates, quickened the larval development, improved fecundity and maximised the longevity whereas the older leaves containing less water and nutrients had an increased cardiac glycoside concentration (Murugan et al., 1997). It has been noted that phytophagous species with a limited host plant spectrum prefer young leaves while the caterpillars of polyphagous species choose mature leaves due to the higher level of toxins in younger leaves (Schoonhoven et al., 2005: 8). The larvae of some moths can distinguish between leaflets from the base, central or terminal areas and avoid base leaflets (Schoonhoven et al., 2005: 11). At close range the female Zebra Swallowtail, Eurytides marcellus is very attracted to host plant volatiles and having found the host plant she relies on vision to determine the young/mature leaf ratio and whether leaf buds are present (Damman & Feeny, 1988). Regarding vision, Papilio species are reported to be able to distinguish between UV, purple, blue, green, red and broadband (Arikawa, 2003). Oviposition stimuli can be non-volatile taste stimulants (Ryuda et al., 2013), so that the butterfly must settle on the leaf of the prospective LHP to establish whether it is a LHP or not. On settling she drums her tarsi and the chemosensilla to determine if the plant has the necessary phytochemicals. Within the Papilionidae, Papilio xuthus needs a concoction with a specific ratio of 10 phytochemicals; P. polytes requires 5 different stimulants and P. protenor needs 6 plant chemicals (Ryuda et al., 2013). P. xuthus has three different sensilla found in the tarsomeres, each with a different sensory function (Ryuda et al., 2013). Certain plants attract Asian Danaid butterfly females, specifically Ideopsis similis, which needs a definite
  • 3. Otto/ Metamorphosis 25: 82–89 84 combination and exact quantities of the 12 studied phenanthroindolizidine alkaloids emitted by its LHP, Tylophora tanakae, to lay her eggs on. When this perfect alkaloid ratio was separated into its individual alkaloids, the female laid fewer eggs. Five of the 12 individual alkaloids examined stimulated oviposition, while 3 of the 5 specific alkaloids served to assist with host plant recognition (Honda et al., 2001). Another interesting fact is that the butterfly Ideopsis similis is cytotoxic due to the presence of these compounds assimilated from the leaves of Tylophora tanakae (Koyama et al., 1999). In South Africa Tylophora also serves as a LHP for Amauris and it may thus be inferred that certain alkaloids will attract the female butterflies and others will stimulate her to oviposit. Some Passifloraceae plants from Central and South America repel oviposition by developing 2 yellow knob-like growths at the base of their leaves, resembling Heliconid eggs and deterring any other butterfly oviposits. Other Passion Vines develop sugary leaf secretions to attract wasps and ants – natural butterfly egg and larvae enemies (Murawski, 1993). Cabbage Whites, Pieris brassicae ovipositing on Arabidopsis plants show that the plant’s whole-genome microarray gene expression changes within three days to rid the plant of the eggs with the cells beneath the egg actually dying and forming calluses (Little et al., 2007). The plant also emits volatiles from insect-damaged leaves to attract larval enemies like parasitoid wasps (Paré & Tumlinson, 1999). American Monarch (Danaus plexippus) butterfly larvae feeding on Tropical Milkweed had reduced parasitoid growth, infection and disease while parasitoid infected females preferred the more toxic Tropical Milkweed to the less toxic Swamp Milkweed, indicating that the females ‘doctored’ their young by choosing a LHP that is more medicinal (Lefèvre et al., 2010). Some arctiids feed on plants with decreased nutritional value yet these plants supply the insects with chemicals that ensure the survival from parasitisation (Schoonhoven et al., 2005: 17). It is hypothesised that common butterflies lay their eggs in clusters and this certainly holds true for some nymphalids, pierids and acraeids while rarer butterflies lay their eggs singly (Stamp, 1980). Acraea encedon encedon may lay several eggs in clusters to increase copulation success by having the few remaining males in close proximity to the females because males are lethally parasitized by Wolburgia bacteria (Jiggins et al., 1998). Lycaeides melissa samuelis from North America on the other hand has two different strategies; the first brood before summer lays eggs singly, while the second brood that overwinters, are laid in batches. The study has two hypotheses; due to the environmental conditions (summer drought) first brood eggs laid singly are distributed over a greater area to support survival, while second brood egg clusters of 1– 6 allows at least one larva to survive (Pickens & Root, 2008). Thus if eggs are laid in clusters during times of scarcity, the survival of at least one larva is assured, which could be as a result of cannibalism (Edge & van Hamburg 2009). Many ovipositing examples mentioned in this paper do not include Lycaenidae, yet it may be extrapolated that species of the Lycaenidae would also utilise several of the ovipositing cues (e.g. Heath & Gardiner, 2009) used by butterflies of the Papilionidae, Pieridae and Nymphalidae while their sensory perception may even be heightened if myrmecophagous Lyceanids are considered. Certain butterflies also rely on LHP stimuli like stachydrine, a secondary compound found in Citrus for the Asian Swallowtail Papilio xuthus which is essential for the larvae to feed (Murata et al., 2011; Ryuda et al., 2013). Pyrrolizidine alkaloids (PAs) allow female Danaid butterflies to recognise their LHPs (Honda et al., 1997; Hartmann et al., 2005), serve as precursors for sexual pheromones (Schulz et al., 2004; Trigo, 2011) and PAs sequestered from plants are utilized as defensive substances (Orr et al., 1996; Nishida, 2002). With Nymphalidae the larvae of the North American Buckeye butterfly needs the secondary compound iridoid glycosides which supports the growth of the larvae (Bowers, 1984). Some Papilionidae like the Black Swallowtails, Papilio polyxenes from Asia have tarsal brushes and these tactile trademarks have to be satisfied by the LHP extracts before egg-laying is stimulated (Roessingh et al., 1991; Ryuda et al., 2013). In the female Pierids of the Sulphur Butterfly, Colias erate, D-pinitol as a stimulant has to be present before the female will oviposit (Honda et al., 2012). CONCLUSIONS Insects ‘sense’ the correct LHP by different methods including; visual, olfactory (Nishida, 2005), mechanical and gustatory (Nishida, 2005; Ryuda et al., 2013) stimuli. A butterfly will oviposit on an alternative plant or object other than its true LHP under these circumstances: 1) During dry seasons the gravid female butterfly will lay near the LHP (Mensah & Kudom, 2012) for the emerging larva to move to the true LHP. 2) If the LHP has been oviposited on but more suitable LHPs are rare in the local vicinity the female may not want to overburden the plant with too many larvae, thus laying on plants nearby relying on the principle of survival of the fittest. 3) When the LHPs present have been stripped by gregarious herbivores like moth larvae, the butterfly larvae laid nearby may emerge in time for the new flush on the LHP after the adult moths have emerged. 4) Some Satyrinid female members of the Tribe dirinii scatter their eggs while in flight. With such complexity present in butterfly behaviour, it must be apparent that butterflies do not randomly lay eggs. ACKNOWLEDGEMENTS Thank you to Mr. Doug McMurtry who imparted his invaluable plant knowledge and willingness to always assist with identifying plants while safekeeping vouchers in his private herbarium using his considerable inspirational ease of finding and concisely expressing the precise text and phrases. Dr. Brian
  • 4. Otto/ Metamorphosis 25: 82–89 85 Schrire graciously assisted with expedient identification of Indigofera. Thank you to Ian and Lize Otto for their accommodation, hospitality, keen interest and support. I am indebted to Dillan Hesselman and Juan Joubert for dedicated field trip accompaniment to find new LHPs and larvae. Thanks also to Ian, Tertia and Allison Sharp for sharing their remarkable work of locating and breeding butterflies on new LHPs in the arid area of Hoedspruit. LITERATURE CITED ARIKAWA, K. 2003. Spectral organization of the eye of a butterfly, Papilio. Journ. of Comparative Physiology A 189: 791–800. BERENBAUM, M.R. & FEENY, P.P. 2008. Chemical Mediation of Host-Plant Specialisation: The Papilionid Paradigm. In Tilmon, K. (ed.) Specialization, speciation and radiation: The evolutionary biology of herbivorous insects. BERNARD, G.D. & REMINGTON, C.L. 1991. Color vision in Lycaena butterflies: Spectral tuning of receptor arrays in relation to behavioral ecology. Proceedings of the National Academy of Science, USA 88: 2783–2787. BOWERS, M.D. 1984. Iridoid glycosides and host- plant specificity in larvae of the Buckeye Butterfly, Junonia coenia, (Nymphalidae). Journal of Chemical Ecology 10(11): 1567–1577. DAMMAN, H. & FEENY, P. 1988. Mechanisms and consequences of selective oviposition by the zebra swallowtail butterfly. Animal Behavior 36: 563– 573. EDGE, D.A. 2008. Adult behaviour of Orachrysops niobe (Trimen) (Lepidoptera: Lycaenidae). Metamorphosis 19(3): 116–126. EDGE, D.A. & VAN HAMBURG, H. 2009. Larval feeding behaviour and myrmecophily of the Brenton Blue butterfly Orachrysops niobe (Trimen). Journal of Research on the Lepidoptera 42: 21–33. FEENY, P., ROSENBERRY, L. & CARTER, M. 1983. Chemical aspects of oviposition behavior in butterflies. In Ahmad S. (ed.) Herbivorous insects: host-seeking behavior and mechanisms. Academic, New York. HARTMANN, T., THEURING, C., BEUERLE, T., KLEWER, N., SCHULZ, S., SINGER, M.S., BERNAYS, E.A., 2005. Specific recognition, detoxification and metabolism of pyrrolizidine alkaloids by the polyphagous arctiid E. tigmene acrea. Insect Biochemistry and Molecular Biology 35: 391–411. HEATH, A. 1998. Further aspects on the life history of the myrmecophilous species Chrysoritis dicksoni (Gabriel), Lepidoptera: Lycaenidae). Metamorphosis 9(4): 160–172. HEATH, A. 2014. Uncovering secrets of the ‘cuckoo’ butterfly species Chrysoritis dicksoni (Gabriel, 1947), a social parasite of Crematogaster ants: A summary to the end of the 20th century with current conclusions. Metamorphosis 25: 5–10. HEATH, A. & GARDINER, A. 2009. Addendum to Heath et al., 2008: Oviposition behaviour and foodplant observations. Metamorphosis 20(1): 8– 10. HEATH, A., MCLEOD, L., KALISZEWSKA, Z.A., FISHER, C.W.S. & CORNWALL, M. 2008. Field notes including a summary of trophic and ant- associations for the butterfly genera Chrysoritis Butler, Aloeides Hübner and Thestor Hübner (Lepidoptera: Lycaenidae) from South Africa. 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  • 6. Otto/ Metamorphosis 25: 82–89 87 Table 1 – Two abundant polyphagous butterfly species which utilise several LHP families and genera BUTTERFLY FAMILY, TAXON PLANT FAMILY PLANT TAXON LYCAENIDAE Leptotes sp. in pirithous group BIGNONIACEAE Tecomaria FABACEAE Abrus, Acacia, Burkea, Indigofera, Mundulea, Neonotonia, Rhynchosia, Sesbania & Vigna PLUMBAGINACEAE Plumbago VERBENACEAE Lantana Hypolycaena philippus philippus CELASTRACEAE Gymnosporia CONVOLVULACEAE Cuscuta LORANTHACEAE Erianthemum OLACACEAE Ximenia RUBIACEAE Fadogia, Kraussia, Pachystigma, Vangueria SAPINDACEAE Deinbollia VERBENACEAE Clerodendrum Table 2 – Repeat oviposition observations, which verify LHP records, with butterfly species, LHPs, plant family, location, observer(s) and dates. FAMILY, TAXON LHP PLANT FAMILY LOCATION, OBSERVER (H.H.H. Otto unless mentioned) DATES HESPERIIDAE Coeliades forestan forestan Indigofera arrecta Hochst. Ex A. Rich. FABACEAE Barberton (Litchie Farm Rd.) Barberton (Litchie Farm Rd.) 15.ii.2014 23.iii.2014 LYCAENIDAE Azanus jesous jesous Dichrostachys cinerea (L.) Wight & Arn. FABACEAE Letsitele Barberton (Litchie Farm Rd.) 20.xii.2013 15.ii.2014 Chilades trochylus Indigofera vicioides Jaub & Spach.var. rogersii (R.E. Fr.) J.B. Gillett FABACEAE Low’s Creek (Three Sisters Farm) Low’s Creek (Three Sisters Farm) Skukuza 16.ii.2013 4.iv.2013 29.iii.2013 Leptotes sp. in pirithous group Abrus laevigata E. Mey. FABACEAE Crocodile Gorge Barberton (Litchie Farm Rd.) i.2008 15.ii.2014 Leptotes sp. in pirithous group Acacia nigrescens Oliv. FABACEAE Shingwedzi Low’s Creek xii. 2011 7.vii.2013 Leptotes spe. in pirithous group Desmodium tortuosum (Sw.) DC. FABACEAE Barberton (Moody’s Rd.) Barberton (Moody’s Rd.) 1.v.14 1.v.14 Leptotes sp. in pirithous group Neonotonia wightii (AM.) J.A. Lackey FABACEAE Low’s Creek (Three Sisters Farm) (plains) Low’s Creek (Three Sisters Farm) (valley) i.2009 ii.2011 Zizeeria knysna Mollugo nudicaulis Lam. MOLLUGINACEAE Low’s Creek (Three Sisters Farm) Barberton (Ameide Rd.) 12.xii.2012 12.xii.2013 Zizeeria knysna Indigofera spicata Forssk. FABACEAE Barberton (Town) Barberton (Town) 5.ii.2014 16.v.2014 Zizula hylax Ruellia patula Jacq. ACANTHACEAE Letsitele Hoedspruit (Sharp, I) 20.xii.2013 xii.2013 Zizula hylax Ruellia malacophylla C.B. Clarke ACANTHACEAE Berg-en-Dal Blyde River, Hoedspruit 30.xi.2013 14.xii.2013 PIERIDAE Eurema hecabe solifera Acacia ataxacantha DC FABACEAE Nelspruit (Karino) Barberton (Litchie Farm Rd.) 6.iv.2014 27.iv.2014 Eurema hecabe solifera Sesbania bispinosa (Jacq.) W.Wight var. bispinosa FABACAEA Marloth Park Barberton (Litchie Farm Rd.) Barberton (Litchie Farm Rd.) iii.2010 15.ii.2014 29.iii.2014 Belenois aurota aurota Maerua rosmarinoides Pax CAPPARIDACEAE Barberton Hoedspruit (Sharp, I.) Barberton xi. 2010 xi.2012 i.2013 Colotis evenina Maerua parvifolia (Sond.) Gilg & Gilg-Ben. CAPPARIDACEAE Shibavantsengele (KNP) Low’s Creek (Three Sisters Farm) xii. 2011 x.2012
  • 7. Otto/ Metamorphosis 25: 82–89 88 Table 3 – New butterfly LHP records, with butterfly species, LHPs, plant family, location, observer, dates and types of observation. FAMILY, TAXON LHP PLANT FAMILY LOCATION, OBSERVER (H.H.H. Otto unless mentioned) DATES TYPE HESPERIIDAE Coeliades forestan forestan Indigofera arrecta Hochst. Ex A. Rich. FABACEAE Barberton (Litchie Farm Rd.) 15.ii.2014 23.iii.2014 11-12.iv.2014 12.v.2014 Oviposition Oviposition Larva Larva LYCAENIDAE Azanus jesous jesous Dichrostachys cinerea (L.) Wight & Arn. FABACEAE Letsitele Barberton (Litchie Farm Rd.) 18.xii.2013 15.ii.2014 Oviposition Oviposition Anthene definita definita Indigofera arrecta Hochst. Ex A. Rich. FABACEAE Barberton (Litchie Farm Rd.) 9-21.iii.2014 29.iii-5.iv. 2014 Bred from final instar. Larva died Cacyreus lingeus Plectranthus fruticosus L’Her. LAMIACEAE Barberton (Upper Rimer’s Creek Rd.) 10.v.2014 Oviposition Cacyreus lingeus Thorncroftia lotterii Edwards & McMurtry LAMIACEAE Barberton (Mountainlands) 10.v.2014 Oviposition Chilades trochylus Indigofera arrecta Hochst. Ex A. Rich. FABACEAE Barberton (Moody’s Rd.) 7.v.2013 Oviposition Chilades trochylus Indigofera schimperi Jaub. & Spach var. schimperi FABACEAE Letsitele 16.xii.2013 Oviposition Chilades trochylus Indigofera spicata Forssk. FABACEAE Barberton (Litchie Farm Rd.) 30.iii.2014 Oviposition Cupidopsis jobates jobates Rhynchosia minima (L.) DC. var. minima FABACEAE Crocodile Gorge 1.i.2014 Oviposition Lampides boeticus Crotalaria lanceolata E. Mey. FABACEAE Barberton (Litchie Farm Rd.) 27.iv.2014 Oviposition Lampides boeticus Crotalaria schlechterii Baker f. FABACEAE Barberton (Litchie Farm Rd.) 23.iii.2014 Oviposition Lampides boeticus Sesbania bispinosa (Jacq.) W.Wight var. bispinosa FABACEAE Barberton (Litchie Farm Rd.) 30.iii.2014 Oviposition Leptotes sp. in pirithous group Indigofera arrecta Hochst. Ex A. Rich. FABACEAE Barberton (Litchie Farm Rd.) 30.iii.2014 Oviposition Leptotes sp. in pirithous group Indigofera galpinii N.E. Br FABACEAE Barberton (Ameide Rd.) 12.xii.2013 Oviposition Leptotes sp. in pirithous group Desmodium tortuosum (Sw.) DC. FABACEAE Barberton (Moody’s Rd.) 1.v.2014 Oviposition Zizula hylax Ruellia malacophylla C.B. Clarke ACANTHA- CEAE Berg-en-Dal (KNP) Hoedspruit (Moriah farm, Blyde River) xii.2013 14.xii.2013 Oviposition Oviposition Zizula hylax Ruellia patula Jacq. ACANTHA- CEAE Hoedspruit (Indabushy) Letsitele (Sharp, I. T. & A.) xii.2013 20.xii.2013 Bred Oviposition NYMPHALIDAE Danaus chrysippus orientis Ceropegia carnosa E. Mey. APOCYNA- CEAE Whyte Thorne, Nelspruit 15.iii.2014 15.iii.2014 Oviposition Feeding larvae Precis tugela tugela Plectranthus fruticosus L’Her. LAMIACEAE Barberton (Litchie Farm Rd.) 29.iii-15/17.iv. 2014 Bred Precis tugela tugela Plectranthus ecklonii Benth. LAMIACEAE Barberton (Litchie Farm Rd.) 29.iii-22.iv. 2014 Bred Precis tugela tugela Plectranthus hadiensis (Forssk.) Schweinf. ex Spreng. LAMIACEAE Nelspruit (Whyte Thorne) 5-30.iv. 2014 Bred Precis tugela tugela Plectranthus laxiflorus Benth. LAMIACEAE Barberton (Litchie Farm Rd.) 29.iii-26.iv. 2014 Bred Precis tugela tugela Plectranthus verticillatus (L.f.) Druce LAMIACEAE Barberton (Litchie Farm Rd.) 29.iii-25.iv. 2014 Bred Precis tugela tugela Pycnostachys urticifolia Hook. LAMIACEAE Barberton (Litchie Farm Rd.) 29.iii-22.iv. 2014 Bred PIERIDAE Eurema hecabe solifera Acacia ataxacantha DC FABACEAE Nelspruit (Karino) Barberton (Litchie Farm Rd.) 6.iv.2014 27.iv.2014 Oviposition Oviposition Eurema hecabe solifera Dichrostachys cinerea (L.) Wight & Arn. subsp. nyassae FABACEAE Barberton (Litchie Farm Rd.) 12.iv- 3.v.2014 (larva died) Oviposition Larva Eurema hecabe solifera Sesbania sesban (L.) Merr. ssp. sesban var. nubica FABACEAE Crocodile Gorge ii.2011 Bred from final instar Eurema hecabe solifera Sesbania bispinosa (Jacq.) W.Wight var. bispinosa FABACEAE Marloth Park Barberton (Litchie Farm Rd.) iii.2010 15.ii.2014 29.iii.2014 Oviposition Oviposition Oviposition
  • 8. Otto/ Metamorphosis 25: 82–89 89 PLATE 1 Figure 1 – Coeliades forestan egg on Indigofera Figure 2 – Coeliades f. forestan larva on arrecta Hochst. Ex A. Rich., Barberton (iii.2014) Indigofera arrecta Hochst. Ex A. Rich., Barberton (iv.2014) Figure 3 – Anthene d. definita larva on Indigofera Figure 4 – Anthene d. definita pre-pupa. arrecta Hochst. Ex A. Rich. Barberton. (iii.2014) Barberton. (iii.2014) Figure 5 – Anthene d. definita pupa about to Figure 6 – Anthene d. definita adult female, eclose. Barberton. (iii.2014) freshly eclosed. Barberton. (iii.2014) Figure 7 – Danaus chrysippus orientis larva on Figure 8 – Chilades trochylus egg on Indigofera Ceropegia carnosa E.Mey. Whyte Thorne, spicata Forssk. (iii.2014) Nelspruit. (iii.2014) Figure 9 – Leptotes sp. ovipositing on Figure 10 – Leptotes sp. egg on Desmodium Desmodium tortuosum (Sw.) DC. (iv.2014) tortuosum (Sw.) DC. (iv.2014)