ArticlePDF Available
Journal of the Lepidopterists’ Society
74(2), 2020, 127–131
Additional key words: behavior, Indonesia, island, nutritional ecology, volcanic sand
Puddling is a common and widespread behavior of
Lepidoptera and other arthropods in which fluid is
imbibed from mud puddles or other sources to obtain
sodium and perhaps other micronutrients (Downes
1973; Molleman 2010). While there have been sporadic
observations of puddling on seawater, the phenomenon
seems to be uncommon or under-reported (Pola &
García-París 2005; John & Tennent 2012). We
document frequent marine puddling by a diverse
community of butterflies on volcanic sands in Sulawesi,
Indonesia, and speculate on the prevalence of the
behavior at this locale.
Puddling is common in several insect orders but is
most conspicuous in Lepidoptera, particularly
butterflies (Papilionoidea). Puddling butterflies extend
their proboscis into a moist substrate and use their
cibarial pump to ingest aqueous solution. Desired
micronutrients are absorbed in the hindgut (Smedley &
Eisner 1995), and an anal jet of filtered water is ejected
every few seconds—sometimes shooting around half a
meter or more (John & Tennent 2012; all authors,
personal observation). Puddling butterflies are nearly
always male and sometimes form large aggregations
comprising hundreds or thousands of individuals
(Downes 1973; Beck et al. 1999; Burger & Gochfeld
2001; Molleman et al. 2005). While presumably coined
in reference to mud puddles, “puddling” is also used to
describe supplementary feeding (i.e., non-nectar
feeding) from stream banks, feces, urine, perspiration
and tears from living mammals, carrion, and—rarely—
seashores (Downes 1973; John & Tennent 2012; Plotkin
& Goddard 2013).
Most evidence suggests that puddling Lepidoptera
seek and sequester sodium (Arms et al. 1974; Adler &
Pearson 1982; Pivnick & McNeil 1987; Smedley &
Eisner 1995, 1996), though there is evidence that
nitrogenous compounds are preferred by some taxa
(Beck et al. 1999; Boggs & Dau 2004; Molleman 2010).
More than 99% of Lepidoptera species consume only
living plant tissue during their larval stages (Pierce
1995), and most plant tissues—particularly the leaves of
terrestrial plants—are a poor source of sodium (Slansky
& Rodriguez 1987). Both sodium and nitrogen are a
scarce resource for most herbivorous insects, including
nectivorous adult butterflies (Seastedt & Crossley 1981;
Liu et al. 2003). In many Lepidoptera species, sodium
accumulated by males through puddling is transferred
to females with their spermatophore during mating
(Pivnick & McNeil 1987; Smedley & Eisner 1996). In
various butterfly species where it has been studied,
sodium increases female longevity and fecundity,
decreases females’ need to forage, and increases male
mating success (Pivnick & McNeil 1987; Molleman et
al. 2004). However, not all butterfly species experience
fitness benefits from supplemental sodium (Molleman
et al. 2004). While it has been suggested that sodium
obtained through puddling enhances neuromuscular
function, as neurons and flight muscles require sodium
ions (Arms et al. 1974), there is little direct evidence for
this hypothesis (Molleman et al. 2005; Molleman 2010).
Interestingly, male Battus philenor butterflies that
consumed sodium courted females differently and more
vigorously than males deprived of the nutrient,
potentially proving an honest signal for females to assess
a male’s sodium titer (Mitra et al. 2016).
Despite the abundance of sodium in seawater, there
are few reported observations of puddling in coastal
marine environments. Pola and García-París (2005)
reported ca. 20 individuals of Papilio polytes drinking
from exposed reef shelves in Guam, providing
unequivocal evidence of direct marine puddling in
butterflies. John and Tennent (2012) reviewed the
occurrence of marine puddling in butterflies, including
published reports and previously unpublished personal
communications documenting this phenomenon. They
note observations of marine puddling near the water’s
edge, but also describe several observations of
butterflies gliding over the ocean and imbibing seawater
with their proboscis while in flight. They concluded that
observations of marine puddling remain rare or under-
reported. The phenomenon has been observed in
Papilionidae, Hesperiidae, Pieridae, Nymphalidae, and
Lycaenidae (Pola & García-París 2005; John & Tennent
2012), though the prevalence of this behavior within the
respective families and its geographical extent remain
poorly known.
On December 9, 2019, the first three authors (YKT,
JSW, and CWG) observed marine puddling along the
black sand beaches of Tangkoko Batuangus Nature
Reserve, North Sulawesi, Indonesia (1° 3433.2N,
Downloaded From:'-Society on 08 Jun 2020
Terms of Use:Access provided by University of Sydney
125° 0936.1E; Fig. 1). The butterflies were initially
seen descending from the canopy before flying up and
down the shoreline patrolling the beach. Occasionally,
individuals flew above the water close to the surface,
where they appeared to drink mid-flight. Similar reports
of aerial seawater drinking have been documented for
Papilio ulysses in Honiara, Solomon Islands (Tennent
1997). Mostly, however, butterflies were observed
puddling on the sand 0–10 m from the waterline, often
flying away only when disturbed by waves (Fig. 1).
Subsequent observations over the span of the next three
days revealed that this was a common and frequent
We observed multiple male individuals of 27
Papilionidae, Hesperiidae, Pieridae, Nymphalidae, and
Lycaenidae species engaging in marine puddling
between 09:00 – 12:00 h and up to 14:00 h at this one
marine site (Fig. 2; Table 1). While we observed no
puddling Riodinidae, we note that there are only four
riodinids out of 557 butterfly species known from
Sulawesi (Vane-Wright & de Jong 2003). Based on our
reading of the available literature, we observed more
butterfly species and individuals puddling on seawater
than have ever been documented before at a single
location (John & Tennent 2012). We also observed that
most butterflies puddled alone, with a maximum group
size of ca. five individuals. So-called “puddle clubs”
(Adler, 2008) in the Indo-Australian Archipelago
including Wallacea can include dozens to hundreds of
individuals in a single group (all authors, personal
observation). Small group size might be explained by
the distance between the ocean and forest cover, ca.
10–15 m. On the one hand, brightly colored butterflies
foraging on dark sand are easily spotted by predators,
and this perhaps limits the number of males willing to
stray so far from the protection of forest cover. On the
other hand, butterflies are not frequent beachgoers, and
puddling males are usually copy-cats. A butterfly is far
more likely to land at a salt source if conspecifics have
already alighted. Experimental studies exploit this
behavior by placing freshly killed, pinned specimens
next to puddles to act as decoys and attract additional
puddling males (Arms et al., 1974). Perhaps the distance
between the ocean and forest cover limits the number
of males that see other puddling males, thus preventing
the formation of large groups.
Why this phenomenon of marine puddling is locally
common at this site in Indonesia is unclear. One
possibility is that terrestrial sodium may be an especially
rare resource in this region of Sulawesi. Although most
afternoons were met with rain, we did not encounter
rivers, streams, or flooded banks—favored habitats for
puddling activity. More curiously, we were unable to
attract many butterflies with urine and fermented
shrimp paste: two baits that that are effective elsewhere,
particularly for Papilionidae and Nymphalidae. It is
possible that paucity of sodium in the terrestrial
environment has resulted in local adaptation for
obtaining the mineral from the sea. Indeed, the habitat
of Tangkoko Batuangus Nature Reserve is not dissimilar
FIG. 1. Marine puddling butterflies on the black sand beaches of Tangkoko Batuangus Nature Reserve. Butterflies were observed
puddling at the edge of the waterline, often taking flight only when disturbed by the incoming surf. Photographs by Jonathan Soong
Wei and Yi-Kai Tea.
Downloaded From:'-Society on 08 Jun 2020
Terms of Use:Access provided by University of Sydney
to Honiara and Guam (Tennent 1997; Pola & García-
París 2005); both are tropical, coastal forests bordered
by the sea. Although seawater intake in northeast
Sulawesi is unlikely to be the only possibility for
butterflies to obtain minerals (we observed lone
individuals of Graphium, Papilio, Cepora, and Vindula
puddling in drains and swamps), it appears to be the
preferred option.
The black sands of these shores might explain why
puddling is more common on this beach than
elsewhere. Black sands are typically rich in minerals
(Mallik et al. 1987) and formed by recent volcanic
activity. Black sands are frequently composed of basalt,
which can be rich in sodium (Hughes & Brown 1972).
Thus, the sand itself may provide additional sodium or
other attractive micronutrients, and the rarity of black
sand beaches might explain the dearth of puddling
observations on beaches. Alternatively, minerals in the
sand may interact with seawater to make it more
appealing to butterflies. Carol Boggs (in John &
Tennent 2012) suggests that seawater may be
underutilized as a salt resource because it contains
unwanted or unsuitable compounds. These substances
might be buffered or chemically modified by black
sands to increase the palatability of seawater.
Apart from the species detailed herein (Fig, 2; Table
1), no other Lepidoptera species were observed
definitively engaging in marine puddling. The coastline
of Tangkoko Batuangus Nature Reserve is bordered by
dense coastal vegetation, a favored habitat for
Apocynaceae plants. Parsonsia and other lactiferous
vines dominated, and so did species of Danaini, which
specialize on these plants. Several adult males in the
genera Danaus, Euploea, and Parantica were observed
TABLE 1. Species of butterflies observed engaging in marine puddling in Tangkoko Batuangus Nature Reserve, North Sulawesi.
Frequency of observations are denoted by (+) symbols, corresponding to the total number of days (out of four) that each species
was observed.
Species Family Subfamily Frequency
Papilio demoleus demoleus Papilionidae Papilioninae ++++
Papilio fuscus minor Papilionidae Papilioninae +
Papilio gigon gigon Papilionidae Papilioninae ++
Papilio sataspes sataspes Papilionidae Papilioninae +
Graphium agamemnon comodus Papilionidae Papilioninae ++++
Graphium codrus celebensis Papilionidae Papilioninae +++
Graphium eurypylus pamphylus Papilionidae Papilioninae ++++
Graphium meyeri meyeri Papilionidae Papilioninae ++
Graphium milon milon Papilionidae Papilioninae ++++
Graphium rhesus rhesus Papilionidae Papilioninae ++++
Badamia exclamationis Hesperiidae Coeliadinae ++
Appias hombroni hombroni Pieridae Pierinae +
Appias zarinda zarinda Pieridae Pierinae +++
Cepora celebensis celebensis Pieridae Pierinae ++
Cepora timnatha timnatha Pieridae Pierinae +
Hebomoia glaucippe celebensis Pieridae Pierinae ++
Catopsilia pomona flava Pieridae Coliadinae ++++
Charaxes nitebis nitebis Nymphalidae Charaxinae +
Cyrestis thyonneus celebensis Nymphalidae Cyrestinae +++
Danaus genutia leucoglene Nymphalidae Danainae +
Euploea eupator eupator Nymphalidae Danainae +
Vindula dejone celebensis Nymphalidae Helioconiinae ++
Doleschallia polibete celebensis Nymphalidae Nymphalinae +
Hypolimnas bolina bolina Nymphalidae Nymphalinae +
Anthene lycaenina Lycaenidae Polyommatinae ++
Catopyrops ancyra subfestivus Lycaenidae Polyommatinae +
Hypolycaena erylus gamatius Lycaenidae Theclinae +
Downloaded From:'-Society on 08 Jun 2020
Terms of Use:Access provided by University of Sydney
congregating in large numbers at the base of these
vines, often directly on seawater-tainted sand. While
most were likely imbibing pyrrolizidine alkaloids
(Boppré 1986), it is unclear if seawater was secondarily
or unintentionally being imbibed. For this reason, we do
not include these species as engaging in marine
puddling (except for Euploea eupator eupator [Fig. 2R]
and Danaus genutia leucoglene [not pictured], which
were seen actively drinking from seawater tainted sand).
Though extensive, this report on marine puddling
activity is likely not exhaustive. The following species
were seen patrolling the beach, but were not observed
puddling, viz: Graphium androcles androcles, Papilio
peranthus adamantius, Pachliopta polyphontes
polyphontes, Troides helena hephaestus, and Troides
hypolitus cellularis. It is likely that the list of marine
FIG. 2. A selection of butterflies from different families engaging in marine puddling. Marine puddling was observed from mem-
bers of all butterfly families found on Sulawesi except Riodinidae (representatives of Hesperiidae not pictured). (A–J) Papilion-
idae, in order of inset letters: Papilio gigon gigon, P. demoleus demoleus, P. sataspes sataspes, P. fuscus minor, Graphium rhesus
rhesus, G. milon milon, G. eurypylus pamphylus, G. meyeri meyeri, G. codrus celebensis, and G. agamemnon comodus; (K–N)
Pieridae: Catopsilia pomona flava, Cepora celebensis celebensis, Hebomoia glaucippe celebensis, Appias zarinda zarinda; (O–R)
Nymphalidae: Vindula dejone celebensis, Cyrestis thyonneus celebensis, Doleschallia polibete celebensis, Euploea eupator eupator
(S–T) Lycaenidae: Anthene lycaenina, Catochrysops ancyra subfestivus. Photographs by Yi-Kai Tea, Jonathan Soong Wei, and
Cheong Weei Gan.
Downloaded From:'-Society on 08 Jun 2020
Terms of Use:Access provided by University of Sydney
puddlers will no doubt increase with additional
observation. Nonetheless, this study provides the most
taxonomically extensive documentation of this behavior
yet observed at a single location.
No specimens were collected during the course of this study.
We thank Rod Eastwood and an anonymous reviewer for their
insightful comments on a previous version of this manuscript.
DJL’s research is funded by grant WW-227R-17 from the Na-
tional Geographic Society Committee for Exploration and Re-
search and by NSF DEB-1541557.
ADLER, P.H. 2008. Puddling behavior by Lepidoptera. In Capinera, JL
(ed.) Encyclopedia of Entomology. Dordrecht: Springer Nether-
lands. 3072–3073 pp.
ADLER, P.H. & D.L. PEARSON. 1982. Why do male butterflies visit
mud puddles? Can. J. Zool. 60: 322–325.
ARMS, K., FEENY, P. & LEDERHOUSE, R.C. 1974. Sodium: Stimulus for
puddling behavior by tiger swallowtail butterflies, Papilio glaucus.
Science 185: 372–374.
BECK, J., MÜHLENBERG, E. & K. FIEDLER. 1999. Mud-puddling be-
havior in tropical butterflies: in search of proteins or minerals?
Oecologia 119: 140–148.
BOGGS, C.L. & B. DAU. 2004. Resource specialization in puddling
Lepidoptera. Environ. Entomol. 33: 1020–1024.
BOPPRÉ, M. 1986. Insects pharmacophagously utilizing defensive
plant chemicals (pyrrolizidine alkaloids). Naturwissenschaften 73:
BURGER, J. & M. GOCHFELD. 2001. Smooth-billed ani (Crotophaga
ani) predation on butterflies in Mato Grosso, Brazil: Risk
decreases with increased group size. Behav. Ecol. Sociobiol. 49:
DOWNES, J.A. 1973. Lepidoptera feeding at puddle-margins, dung,
and carrion. J. Lepid. Soc. 27: 89–99.
HUGHES, D.J. & G.C. BROWN. 1972. Basalts from Madeira: A petro-
chemical contribution to the genesis of oceanic alkali rock series.
Contrib. Mineral. Petr. 37: 91–109.
JOHN, E. & W.J. TENNENT. 2012. Marine (seawater) puddling by but-
terflies: Is the sea an underutilised sodium resource? Entomol.
Gaz. 63: 135–145.
LIU, W., J.E.D. FOX, & Z. XU. 2003. Nutrient budget of a montane
evergreen broad-leaved forest at Ailao Mountain National Nature
Reserve, Yunnan, southwest China. Hydrol. Process. 17:
The black sand placer deposits of Kerala beach, southwest India.
Mar. Geol. 77: 129–150.
MITRA, C., E. REYNOSO, G. DAVIDOWITZ & D. PAPAJ. 2016. Effects of
sodium puddling on male mating success, courtship and flight in
a swallowtail butterfly. Anim. Behav. 114: 203–210.
MOLLEMAN, F. 2010. Puddling: From natural history to understanding
how it affects fitness. Entomol. Exp. Appl. 134: 107–113.
BRAKEFIELD. 2005. Is male puddling behaviour of tropical but-
terflies targeted at sodium for nuptial gifts or activity? Biol. J.
Linn. Soc. 86: 345–361.
MOLLEMAN, F., B.J. ZWAAN & P.M. BRAKEFIELD. 2004. The effect of
male sodium diet and mating history on female reproduction in
the puddling squinting bush brown Bicyclus anynana (Lepi-
doptera). Behav. Ecol. Sociobiol. 56: 404–411.
PIERCE, N.E. 1995. Predatory and parasitic Lepidoptera: Carnivores
living on plants. J. Lepid. Soc. 49: 412–453.
PIVNICK, K.A. & J.N. MCNEIL. 1987. Puddling in butterflies: Sodium
affects reproductive success in Thymelicus lineola. Physiol. Ento-
mol. 12: 461–472.
PLOTKIN, D. & J. GODDARD. 2013. Blood, sweat, and tears: A review
of the hematophagous, sudophagous, and lachryphagous Lepi-
doptera. J. Vector Ecol. 38: 289–294.
POLA, M. & M. GARCÍA-PARÍS. 2005. Marine puddling in Papilio poly-
tes (Lepidoptera: Papilionidae). Fla. Entomol. 88: 211–213, 213.
SEASTEDT, T.R. & D.A. CROSSLEY. 1981. Sodium dynamics in forest
ecosystems and the animal starvation hypothesis. Am. Nat. 117:
SLANSKY, F. & J.G. RODRIGUEZ. 1987. Nutritional Ecology of Insects,
Mites, Spiders, and Related Invertebrates. Wiley-Interscience,
New York.
SMEDLEY, S.R. & T. EISNER. 1995. Sodium uptake by puddling in a
moth. Science 270: 1816–1818.
SMEDLEY, S.R. & T. EISNER. 1996. Sodium: A male moth's gift to its
offspring. Proc. Natl. Acad. Sci. USA. 93: 809–813.
TENNENT, W.J. 1997. Unusual behaviour in Papilio ulysses L., 1785
(Lep.: Papilionidae). Entomol's. Rec. J. Var. 109: 156–157.
VANE-WRIGHT, R.I. & R. DE JONG. 2003. The butterflies of Sulawesi:
Annotated checklist for a critical island fauna. Zool. Verh. 343:
YI-KAI TEA*School of Life and Environmental
Sciences, University of Sydney, Sydney, Australia;
Department of Ichthyology, Australian Museum
Research Institute, Australian Museum, 1 William
Street, Sydney, NSW 2010 Australia. JONATHAN SOONG
WEI 892 Upper Bukit Timah Road, #04-21, Singapore
678187; CHEONG WEEI GAN Nature Society Singapore,
510 Geylang Road, #02-05, Singapore 389466. AND
DAVID J. LOHMAN Biology Department, City College of
New York, City University of New York, 160 Convent
Ave., New York, NY, 10031 USA; Ph.D. Program in
Biology, Graduate Center, City University of New York,
365 5th Ave., New York, NY, 10016 USA; and
Entomology Section, National Museum of Natural
History, T.F. Valencia Circle, Rizal Park, Ermita,
Manila, 1000 Philippines.
*Corresponding email:
Submitted for publication 18 March 2020; revised and
accepted 5 April 2020.
Downloaded From:'-Society on 08 Jun 2020
Terms of Use:Access provided by University of Sydney
... Members of the G. sarpedon complex (Papilionidae: Papilioninae: Leptocircini) are widely distributed throughout tropical and subtropical Asia, Australia, and Melanesia (Tsukada & Nishiyama 1980). Males are frequently encountered mud puddling (e.g., Tea et al. 2020), and both sexes can be seen nectaring at a variety of different flowers. However, research on the G. sarpedon complex is complicated by disagreement over species limits in this geographically widespread group (Page & Treadaway 2013). ...
Full-text available
After molecular and morphological analyses, the taxon septentrionicolus Page & Treadaway, 2013 is shown to be a distinct species, and Graphium adonarensis (Rothschild, 1896) is placed as conspecific with Graphium sarpedon (Linnaeus, 1758). Graphium huangshanensis Wu & Ma, 2016 syn. nov. is synonymised with G. septentrionicolus.
... Danaus species of the Americas, for example, are apparently not known to perform leaf-scratching (Ramos et al. 2020, Lawson et al. 2021. It is possible that the dearth of natural history observations on Sulawesi may have precluded observations elsewhere on the island (but see Tea et al. 2020). ...
Full-text available
Milkweed butterflies (Lepidoptera: Nymphalidae: Danainae) are chemically defended and aposematic (Ackery and Vane Wright 1984). The chemical ecology of several species have been studied extensively, including the iconic and highly migratory Monarch Butterfly, Danaus plexippus (Brower and Glazier 1975; Boppré 1993). Danaine larvae feed on chemically defended plants from which they also extract phytochemical protection.
Samia watsoni was reared in Yunnan from eggs received from Zhejiang in 2021. A mature larva was found in the field on Pterostyrax corymbosus (Styracaceae) in 2020, so this tree was used as the hostplant in captivity. Larvae accepted other Styracaceae in captivity. There are probably other hostplants in nature, but captive larvae refused Ailanthus and Liquidambar. The mature larva is light bluish green, white dorsally, with red dorsal scoli and others blue. In captivity the cocoons were wrapped in a leaf attached to the hostplants, but half were spun at the bottom of the cage with weak peduncles, suggesting that some cocoons may occur on the ground in nature. Adults were observed to drink water along the edges of streams, and a functional proboscis is reported for other Attacini.
Full-text available
Recent years have witnessed considerable range expansion of the migratory Indo-Australian papilionid Papilio demoleus Linnaeus, 1758 from the Gulf States. Following dispersal / migration into central Syria, a rapid and extensive colonisation of the eastern Mediterranean coastlines of Syria soon ensued, with penetration into neighbouring regions of Turkey and Lebanon. Further westward spread had been anticipated and here we report on the species' first appearance in Cyprus. We hypothesise that westward trans-Mediterranean migration brought small numbers of immigrants to the island, and from the pristine nature of the individuals, it is considered that those seen in August 2021 were the progeny of an earlier, unnoticed migration. We also record the Levant's first known example of marine puddling by P. demoleus.
Full-text available
In many Lepidoptera species usually only males puddle for sodium. Two explanations have been offered for this: (1) neuromuscular activity: males need increased sodium for flight because they are more active flyers than females; and (2) direct benefits: sodium is a type of direct benefit provided by males to females via ejaculate during mating. Surprisingly, there is little direct experimental evidence for either of these. In this study, we examined both explanations using the pipevine swallowtail butterfly, Battus philenor L. If sodium increases neuromuscular activity, males consuming sodium should be better fliers than males without sodium. If males collect sodium for nuptial gifts that benefit their mates, males consuming sodium may have greater mating success than males without sodium. In that case, females then need an honest cue/signal of the quality of male-provided direct benefits that they can assess before mating. If sodium affects male courtship flight by increasing neuromuscular activity, how a male courts could serve as such a premating cue/signal of male benefit quality. Therefore, sodium may benefit males in terms of obtaining mates by increasing their neuromuscular activity. In this study we found that males that consumed sodium courted more vigorously and had greater mating success than males that consumed water. In addition, the courtship displays of males consuming sodium were significantly different from those of males consuming water, providing a possible honest cue/signal of male benefit quality that females can assess. Interestingly, we did not find evidence that sodium consumption affects male flight outside of courtship. That only aspects of male flight related to mating were affected by sodium, while aspects of general flight were not, is consistent with the idea that sodium may benefit males in terms of obtaining mates via effects on neuromuscular activity.
Full-text available
All species and subspecies of butterflies recorded from Sulawesi and neighbouring islands (the Sulawesi Region) are listed. Notes are added on their general distribution and hostplants. References are given to key works dealing with particular genera or higher taxa, and to descriptions and illustrations of early stages. As a first step to help with identification, coloured pictures are given of exemplar adults of almost all genera. General information is given on geological and ecological features of the area. Combined with the distributional information in the list and the little phylogenetic information available, endemicity, links with surrounding areas and the evolution of the butterfly fauna are discussed.
Full-text available
While there are numerous documented records of butterflies 'mud puddling' in order to obtain salts and minerals by imbibing liquid from a freshwater source, a lack of published evidence regarding similar exploitation of seawater suggests this is a rarely used sodium resource, or is a behavioural activity infrequently observed or recognised. As background to introducing new reports of seawater exploitation, we first briefly review papers on puddling in general, noting an almost total absence of references to seawater use. Following a trawl amongst entomological colleagues, further instances of butterflies imbibing seawater are presented, including additional observations by the authors, among them the first known report of marine puddling by the Palaearctic hesperiid Thymelicus acteon and a second report of direct seawater intake by the tropical papilionid Papilio ulysses. The possible significance of these observations is briefly discussed.
Full-text available
Lepidoptera feed at mud puddles, dung, and carrion in a behavior known as puddling. Sodium and sometimes protein are feeding cues, are actively collected, and play a potentially important role in lepidopteran nutritional and mating ecology. We showed that montane butterfly species have feeding preferences among mud, herbivore dung, and carnivore dung, and that these preferences differed among butterfly species. The puddling substrates varied in soluble sodium content, with mud having the lowest concentrations and carnivore dung having the highest. Within one species, Pieris napi L., visit frequencies to mud versus dung matched visit frequencies to sand trays filled with sodium solutions matching the concentrations seen in mud or dung. This suggests that the preference hierarchy of this species is driven by soluble sodium concentration. Overall, the results indicate that lepidopteran species specialize on different puddling substrates, likely obtaining different arrays of nutrients. This suggests that there are species- or family-specific roles for puddling nutrients in the overall nutrient budget of the insects.
Moths and butterflies whose larvae do not feed on plants represent a decided minority slice of lepidoteran diversity, yet offer insights into the ecology and evolution of feeding habits. This paper summarizes the life histories of the known predatory and parasitic lepidopteran taxa, focussing in detail on current research in the butterfly family Lycaenidae, a group disproportionately rich in aphytophagous feeders and myrmecophilous habits. -Author
Although adult Lepidoptera are not often considered medically relevant, some butterflies and moths are notorious for their consumption of mammalian body fluids. These Lepidoptera can be blood-feeding (hematophagous), tear-feeding (lachryphagous), or sweat-feeding (we use the term "sudophagous"). Blood-feeding Lepidoptera have been observed piercing the skin of their hosts during feeding, while tear-feeding Lepidoptera have been observed frequenting the eyes of hosts in order to directly obtain lachrymal fluid. These behaviors have negative human health implications and some potential for disease transmission. In this study, articles concerning feeding behavior of blood, sweat, and tear-feeding Lepidoptera were reviewed, with emphasis on correlations between morphological characters and feeding behaviors. Harmful effects and vector potential of these Lepidoptera are presented and discussed.
This study is the first to demonstrate that Na budgets of male and female Lepidoptera differ. At the time of emergence, male imported cabbage butterflies, Pieris rapae L., have significantly more total body Na than females. Older males collected from the field show a significantly lower level of body Na than freshly emerged males, whereas freshly emerged females and older, field-collected females show no difference. It is suggested that feeding from soil may help restore losses of Na in males. A single female, through oviposition, may lose nearly 75% of the total body Na with which it emerged. The lepidopteran mating system involving transfer of spermatophores is postulated as one causative factor for sexual differences in body Na levels.