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2016 Tikocaetal Pacific Science
2016 Tikocaetal Pacific Science
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Temporal Variation in Macro-Moth Abundance and Species Richness
in a Lowland Fijian Forest1
Siteri Tikoca,2,6 Simon Hodge,3,4 Sarah Pene,2 John Clayton,5 Marika Tuiwawa,2
and Gilianne Brodie 3
Seasonal patterns in the presence, abun plant abundance, and accessibility of nectar
dance, and activity of Lepidoptera have been ( Intachat et al. 2001, Pozo et al. 2008). In
explained as a function of both climatic and temperate regions, the timing and duration of
biotic factors, such as temperature, precipi flight periods of adult moths often display
tation, seasonality of vegetation, larval host well-defined seasonal sequences due to the ar
rival of different species by emergence and /or
migration, differences in life history traits,
1
This work was funded with grants from The Uni overwintering, voltinism, adult life span, and
versity of the South Pacific ( Institute of Applied Sciences growth rates of immature stages ( Wolda
and School of Biological and Chemical Sciences). Manu 1988, Highland et al. 2013, Jonason et al.
script accepted 1 June 2016.
2
Institute of Applied Sciences, The University of the
2014). However, in the tropics the effects of
South Pacific, Suva, Fiji Islands. climate and plant phenology on lepidopteran
3
Faculty of Science, Technology, and Environment, seasonality may not be as prominent because
The University of the South Pacific, Suva, Fiji Islands. many regions have relatively stable weather
4
Faculty of Agriculture and Life Sciences, Lincoln conditions (temperature and rainfall) and
University, Canterbury, New Zealand.
5
15 Whinny Brae, Broughty Ferry, Dundee DD5 many plant species are present all year round
2HU, Scotland. ( Wolda 1988, Barlow and Woiwood 1989).
6
Corresponding author (e-mail: stikoca@gmail.com). A number of previous studies have focused
on seasonal occurrences, fluctuations in abun
dance, and species composition of tropical
Pacific Science (2016), vol. 70, no. 4:447 – 461
doi:10.2984/70.4.5 moth assemblages (e.g., Barlow and Woiwood
© 2016 by University of Hawai‘i Press 1989, Kitching et al. 2000, Intachat et al.
All rights reserved 2001, Duarte and Schlindwein 2005, Beck
447
448 PACIFIC SCIENCE · October 2016
and Linsenmair 2006). Assemblages of noc cated in the province of Naitasiri, 7 km from
turnal moths have long been considered valu Suva, Fiji Islands (18.058° S, 178.467° E).
able biological indicators for monitoring the The vegetation at Colo-i-Suva contains a
effects of habitat degradation or conservation mixture of both exotic timber species and
success (e.g., Kitching et al. 2000, Beck et al. native species at various growth stages in the
2002, Hawes et al. 2009), and many projec understory. In a recent survey, Tuiwawa and
tions of climate change effects on ecological Keppel (2013) reported that Meliaceae and
communities predict substantial local extinc Myrtaceae were the most diverse families at
tions of moth species, changes in geographic the site, and noticeable species included
distribution, or modification of seasonality mahogany (Swietenia macrophylla, Meliaceae)
(e.g., Conrad et al. 2006, Altermatt 2010, that has naturally regenerated, Gironniera
Maclean and Wilson 2011). Therefore, to
celtidifolia (Cannabaceae), Dillenia biflora
monitor long-term changes in lepidopteran (Dilleniaceae), Barringtonia edulis (Lecy
assemblages in tropical forests, and thus eval thidaceae), and Endospermum macrophyllum
uate any potential effects of climate change, (Euphorbiaceae).
detailed baseline data are required, which Hand collecting specimens attracted to a
must account for the variation in the insect mercury-vapor (MV ) lamp has proved to be a
communities that occurs seasonally as well as successful method for obtaining nocturnal
spatially (Basset et al. 2015). moths in Fijian forests (Tikoca, Hodge, et al.
The Fiji Islands undergo a distinct wet 2016). Therefore sampling was performed
season (
November – April) and dry season using a manual light-trap system consisting of
(May – October) that is mainly regulated by a 125W MV lamp powered by a portable gen
the north and south movement of the South erator and a 2 by 2 m white sheet, which was
Pacific Convergence Zone (SPCZ). Through spread out and secured onto nearby trees or
out the year, the predominant winds are east branches and positioned in front of the light
to southeasterly trade winds that blow slightly source. All macro-moths that flew toward the
stronger during the cool dry season compared light and onto the white sheet were collected
with the warmer wet season. On the main is and placed into glass jars charged with ethyl
land of Viti Levu, the large central mountain acetate as a killing agent. Each sample of
ous ranges cause wet climatic zones on the moths consisted of the individuals collected
windward side and dry climatic zones on the for 4 hr from dusk.
leeward side (Mueller-Dombois and Fosberg Sampling was carried out for 12 months
1998). from December 2011 to November 2012 on
Although Chandra et al. (2013) provided 2 or 3 nights during the last week of each cal
some observations on the seasonality of the endar month when weather was forecast to
Fijian swallowtail butterfly, Papilio schmeltzi, be fine (i.e., low wind, no rain). Moths were
in Fijian forests, we could find no previous sampled on 30 nights in total (see Appendix
reports of studies specifically examining the for actual sampling dates). The catches
seasonality or phenology of a Fijian moth were averaged over the samples taken in each
assemblage. The current investigation was
calendar month and then divided into four
carried out over a 12-month period to exam seasonal periods: four “wet months”
ine how the abundance, species richness, and (December
–
March), four “dry months”
composition of adult macro-moths in a ( June – September), and the two bi-monthly
mixed-lowland forest varied over time. At periods on either side of the dry season identi
tempts were made to relate temporal variation fied as “transitional” periods (i.e., April – May,
in moth abundance and richness to climatic trans wet – dry; October – November, trans
factors such as rainfall and temperature. dry – wet). The families of macro-moths in
cluded in the survey are those covered by
materials and methods Robinson (1975) in his treatise of Fijian
moths. Robinson included the Limacodidae
Moths were sampled along the ridge of a low (which are not often thought of as macro-
land mixed forest, Colo-i-Suva, which is lo moths these days) but excluded all the Pyra
Temporal Patterns in Fijian Forest Moths · Tikoca et al.449
Chrysodeixis eriosoma ( Noctuidae, 57, 11 tightly grouped than those taken in the dry
months), Bulonga philipsi (Geometridae, 51, season (Figure 4), suggesting that the compo
10 months), Urapteroides anerces ( Uranidae, sition of the moth fauna was more consistent
46, 11 months), Calliteara fidjiensis (Erebidae, in the wet months than in the dry months.
41, 9 months), Oxyodes scrobiculata (Erebidae, The abundance of no individual species ex
39, 8 mo.), Polydesma boarmoides (Erebidae, 32, hibited very strong (rs > |0.75|) rank correla
11 mo.), and Ericaea leichardtii (Erebidae, 31, tions with the NMDS axis scores, although
9 months). In general, consistency of occur species showing some association (rs > |0.5|)
rence (number of months observed) and over with the wet-season samples were Spodoptera
all abundance were highly related (rs = 0.936, litura ( Noctuidae), Maceda mansueta ( Noli
P < .001, N = 116 species), and all of these dae), and Gymnoscelis sara (Geometridae).
common species were recorded in at least Species associated with the dry-season sam
eight of the 12 surveyed months (although ples were Simplicia cornicalis (Erebidae), Tira
only the combined Cleora spp. group was re cola plagiata ( Noctuidae), and Hydrillodes
corded in all 12). Conversely, some species surata (Erebidae).
were found over most of the year, but in rela
tively low numbers. For example, Agathia Climatic Factors
pisina (Geometridae) was recorded in 11
months, but only 22 specimens were recorded The average maximum daily temperature
in total. Similarly, Thyas miniacea (Erebidae) ranged from 26.2°C in July to 30.6°C in
and Stictoptera stygia ( Noctuidae) were re December, and the average minimum daily
corded in nine months, with 22 and 17 speci temperature ranged from 19.9°C in June to
mens, respectively. 23.4°C in January. Average daily rainfall
Although the 12 most common species peaked at 15.5 mm in March and was lowest
(>30 specimens recorded) were present over in August with 3.3 mm. Relative humidity was
most of the year, there appeared to be distinct always high, ranging from 79.7% in October
peaks in abundance for many of these species. to 87.0% in March.
For example, Spodoptera mauritia peaked in There were no significant relationships be
abundance in February, followed by Urapteroi tween total abundance and species richness
des amerces, which peaked in March (Figure 2). with any of the climatic factors measured
Bulonga philipsi, Chrysodeixis eriosoma, Oxyodes (Table 1, Figure 5). When considering the
scrobiculata, and Condica illecta subsequently abundance of the twelve common species in
peaked in abundance around April –
May dividually, only two exhibited relationships
(Figure 2). Cleora sp., Oeonistis delia, Polydesma with any of the climatic variables measured
boarmoides, and Ericaea leichardtii appeared to (Table 1). Hydrillodes surata exhibited signifi
have no distinct periods of high and low abun cant negative relationships with minimum
dance, whereas Hydrillodes surata and Calli and maximum daily temperature and with
teara fidjiensis were most abundant toward mean daily rainfall, whereas Bulonga philipsi
the end of the year (Figure 3). exhibited positive relationships with mini
Although there were no obvious trends in mum and maximum daily temperature (Table
species richness over the course of the year, 1, Figure 6).
the composition of the faunas found within
each of the seasonal periods showed some
discussion
distinctiveness, as illustrated by the results of
the NMDS (Figure 4). The monthly samples The macro-moth assemblage of Colo-i-Suva
from the wet and dry seasons formed distinct did not show major fluctuations in abun
groups on the basis of their positions on the dance and species richness over the course
first NMDS axis, with the wet – dry and of 12 months, and we found no evidence that
dry – wet transition periods taking up interme total abundance and diversity were related
diate positions (Figure 4). The four monthly to climatic factors. The lack of strong tem
samples taken in the wet season were more poral trends in these features of the adult
Temporal Patterns in Fijian Forest Moths · Tikoca et al.451
Figure 2. Fluctuations in abundance (mean moths per sample night) for common macro-moths captured in Colo-i-
Suva forest, Fiji, exhibiting peaks in abundance in the wet season and wet – dry transition period.
moth assemblage may initially appear unusual similarly trapped moths in the United King
because some form of synchrony with the dom, the seasonal patterns in the tropics were
presence of larval host plants, emergence of much less pronounced than those in the tem
young foliage, and the presence of flowers perate location. It is possible that the cli
to provide nectar might be expected ( Wolda mate within the forest is stable enough, and
1978, van Schaik et al. 1993). Barlow and the required plant species present in suffi
Woiwood (1989) reported similar findings cient quantities and in a suitable develop
for moths in tropical Malaysia, suggesting mental stage, to allow the moth assemblage
that although diversity and biomass were to remain relatively stable over the whole
much higher in the tropical forest than in 12-month cycle.
452 PACIFIC SCIENCE · October 2016
Figure 3. Fluctuations in abundance (mean moths per sample night) for common macro-moths captured in Colo-i-
Suva forest, Fiji, exhibiting no obvious periods of high or low abundance or peaking in abundance toward the end of
the year.
All of the most common species in our nset of heavy rainfall in tropical forests can
o
study occurred as adults over most of the year, trigger “bud-break” and flowering in many
although a few of these species did exhibit dis plants (e.g., DeVries and Walla 2001, Pinhei
tinct peaks in abundance, a finding similar to ro et al. 2002, Hilt et al. 2007). However, the
that described by Hilt et al. (2007) for moths correlation between moth abundance and
in a montane forest in southern Ecuador. climatic factors may not always be straight
Some previous studies conducted in tropical forward. For example, Intachat et al. (2001)
areas have indicated an increase in insect found that abundance and richness of geome
abundance in the wet season, because the trid moths in a Malaysian tropical forest were
Temporal Patterns in Fijian Forest Moths · Tikoca et al.453
Figure 4. Plot of axis 2 versus axis 1 scores obtained from nonmetric multidimensional scaling based on captures of all
macro-moth species (mean per month) obtained in Colo-i-Suva Forest over the 12-month period (stress = 13.7%).
Points on the graph are based on composition of moth assemblage obtained in each of 12 months, assigned to dry
season, wet season, and wet – dry ( WD) or dry – wet (DW ) transition periods (see Materials and Methods section for
details). The named moth species were strongly correlated with the axis indicated.
actually negatively related to rainfall and rela are typical of these seasons. However, there
tive humidity, but were positively correlated was at least one case where some consistency
to the incidence of tree flowering the previous was seen across the different analytical
month. Our data set was of insufficient dura approaches. The erebid Hydrillodes surata
tion to properly examine for the occurrence showed a general increase in abundance in the
of time lags relating moth abundance to cli dry-season months (Figure 3), and this was
matic conditions that occurred some months also seen in the NMDS analysis, where H.
earlier; however this possibility cannot be surata was correlated with the axis 1 scores
ruled out. associated with dry-season samples (Figure 4).
Although the number of species and abun Hydrillodes surata abundance was negatively
dance of moths remained fairly constant over correlated with temperature and rainfall (Fig
the monitoring period, the multivariate anal ure 6), indicating that it was most numerous
ysis suggested that the compositions of the when temperature and rainfall were relatively
assemblages within the dry and wet seasons low, both characteristics of the dry-season cli
were actually quite distinct. This separation mate in this part of Fiji.
of the moth samples into distinct seasons us It is possible that more species do show
ing NMDS appears to contradict the lack some relationship with the climatic fac
of relationships with the climatic factors that tors that we investigated, but by using a
454 PACIFIC SCIENCE · October 2016
TABLE 1
Values of Spearman’s Rank Correlation Coefficient Illustrating Strengths of Relationships between Climatic Factors
with Total Moth Abundance and Species Richness, and with Abundance of the 12 Most Common Species
N = 12 months.
*P < .05; **P < .01).
Figure 6. Scatterplots illustrating relationships between abundances of Hydrillodes surata and Bulonga philipsi with
climatic factors for which statistically significant relationships occurred (Spearman’s rank correlation, P < .05; see
Table 1).
With data based on only a single year of reoccur in subsequent years ( Wolda 1988).
observations we cannot make conclusions Although seasonal patterns were indicated for
regarding whether the phenomena we have a few Fijian moth species by Robinson (1975),
observed are true seasonal patterns that will we believe that ours is the first study that has
Temporal Patterns in Fijian Forest Moths · Tikoca et al.457
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Temporal Patterns in Fijian Forest Moths · Tikoca et al.459
Appendix
Macro-Moths Captured at Colo-i-Suva Forest Reserve, Fiji, by Mercury-Vapor Light Trapping from December
2011 to November 2012 (*, captured; — , not captured; N, number of specimens collected in total)
Sampling dates (day-month-year): 29-12-11, 30-12-11, 25-01-12, 26-01-12, 27-01-12, 22-02-12, 23-02-12, 24-02-12,
23-03-12, 24-03-12, 28-04-12, 29-04-12, 30-04-12, 30-05-12, 31-05-12, 28-06-12, 29-06-12, 26-07-12, 27-07-12,
28-07-12, 27-08-12, 28-08-12, 29-08-12, 28-09-12, 30-09-12, 20-10-12, 30-10-12, 31-10-12, 27-11-12, 28-11-12.
Family Species N D J F M A M J J A S O N
Cossidae Acritocera negligens 2 — — — — — — * — — — — —
Erebidae Achaea robinsoni 6 * — — * — * — * — — — —
Erebidae Adetoneura lentiginosa 3 — — — * * — — — — — — —
Erebidae Aedia sericea 2 — — — — — — — — — — * —
Erebidae Anomis samoana 1 — — — — — — — — — * — —
Erebidae Argina astraea 3 — — — — — — — — — — * —
Erebidae Asota woodfordi 2 — — — * — — — — — — * —
Erebidae Bocana manifestalis 11 * * * — — * — * — — — —
Erebidae Caliteara flavobrunnea 1 — — — — * — — — — — — —
Erebidae Calliteara fidjiensis 41 * * * * * — — * — * * *
Erebidae Cosmophila flava 2 — — — — * * — — — — — —
Erebidae Dysgonia duplicata 1 — — — — — — — — — — * —
Erebidae Dysgonia hicanora 1 — — — — — * — — — — — —
Erebidae Dysgonia prisca 12 * — — — * * — * * * * *
Erebidae Ericaea inangulata 11 — — * — * * — — — — * —
Erebidae Ericaea leichardtii 31 * * * — * * — * — * * *
Erebidae Eudocima fullonia 1 — — — — * — — — — — — —
Erebidae Gonitis involuta 1 — — — — — * — — — — — —
Erebidae Hydrillodes surata 61 * * * * — * * * * * * *
Erebidae Hypena cornicalis 2 — — — — — * — — — — — —
Erebidae Hypena rubrescens 12 — — — — — * — * * * — —
Erebidae Hypenagonnia barbara 3 — — * — — — * — — — — —
Erebidae Hypenagonnia diana 2 — — * * — — — — — — — —
Erebidae Hypenagonnia emma 3 * * * — — — — — — — — —
Erebidae Hypospila similis 9 — — — — * * — * * * — —
Erebidae Lophocoleus rubrescens 1 — — — — — — — — — * — —
Erebidae Mecodina variata 5 — — — — — — — * * * — —
Erebidae Mocis frugalis 22 * — * * * * * * — — — —
Erebidae Mocis trifasciata 1 — — — — — — — * — — — —
Erebidae Neogabara plagiola 4 — * — * — — — — — — * —
Erebidae Oeonistis delia 103 — * * * * * * * * * * *
Erebidae Ophiusa disjungens 1 — — — — * — — — — — — —
Erebidae Oruza cariosa 1 — — — — — — — * — — — —
Erebidae Oxyodes scrobiculata 39 * * — — * * * * * — * —
Erebidae Palaeocoleus sypnoides 14 * * * — — * — * — — * *
Erebidae Polydesma boarmoides 32 * * * * * * — * * * * *
Erebidae Rhesalides curvata 2 — — — — — — — — — — * —
Erebidae Rusicada vulpina 3 * — — — — — * — — — * —
Erebidae Serodes mediopallens 3 — — — — — * * — — — * —
Erebidae Serrodes campana 3 — — * — * — — — — — — —
Erebidae Simplicia cornicalis 12 — — — — — — — * * * * *
Erebidae Thyas miniacea 22 * — * * * * * * * — * —
Erebidae Trigonodes cephisa 3 — — — — — * — — — — — —
Geometridae Agathia pisina 22 * * * * * * * * * * — *
Geometridae Anisodes gloria 1 — — — — — — — — — — * —
Geometridae Anisodes monetara 27 — * — * — * * * * * * *
Geometridae Anisodes oblivaria 5 * * — — * * — — — — — —
Geometridae Anisodes prionodes 1 — — — — — — * — — — — —
Geometridae Anisodes samoana 2 — — — — — * — — — — — —
Geometridae Brabira apatopleura 1 — — — * — — — — — — — —
Geometridae Bulonga philipsi 51 * * * * * * — * — * * *
460 PACIFIC SCIENCE · October 2016
Appendix (continued)
Family Species N D J F M A M J J A S O N
Geometridae Chlorochaeta cheromata 4 * * * — — — — — — — — —
Geometridae Chloroclystis encteta 2 — — — — — — — — — — * —
Geometridae Chloroclystis nina 1 — — — — — — — — — * — —
Geometridae Chloroclystis rubicunda 2 — — — — — — — * — — — —
Geometridae Cleora spp. 117 * * * * * * * * * * * *
Geometridae Eoasthena gnophobathra 1 — — — — — — — * — — — —
Geometridae Episteira nigrilinearia 1 — — * — — — — — — — — —
Geometridae Gelasma stuhlmanii 8 * * * * * — — — — — — —
Geometridae Gymnoscelis sara 2 — — — — — * — — — — * —
Geometridae Horisme chlorodesma 19 — — * * — * — * — * * *
Geometridae Luxiaria sesquilinea 8 * * * * — — — * — — — —
Geometridae Mnesiloba eupitheciata 11 * — — — — — — * * — — —
Geometridae Nadagara irretracta 1 — — — — — — — — — — * —
Geometridae Petelia aesyla 4 — — * * — * — — — — — —
Geometridae Polyclysta gonycrota 3 — — — — — — — * — — — —
Geometridae Pseuderythrolopus 2 — — — — — — — — — — * *
bipunctatus
Geometridae Pyrrhorachis pyrrhogona 13 — * * * * * * * — — — —
Geometridae Scardamia eucampta 3 * — — — — — * * — — — —
Geometridae Scotocyma miscix 1 — — — — — * — — — — — —
Geometridae Thalasodes pilaria 6 — — — — * — — * — — — —
Geometridae Thalassodes chloropis 4 * * — — — — — — — — — —
Geometridae Thalassodes figurata 2 — — — * — — — * — — — —
Geometridae Thalassodes fiona 26 — * * * * * * * * * — *
Geometridae Thalassodes liquescens 2 * — — — — — — — — — — —
Limacodidae Beggina albafascia 3 — — — — * — * — — — * —
Limacodidae Beggina bicornis 2 — — — — — — — * — — * —
Limacodidae Beggina mediopunctata 3 — — — * * — * — — — — —
Limacodidae Beggina minima 16 * * * * * — — — — * * *
Limacodidae Beggina sp. 6 — — * * — — — * — — — *
Limacodidae Beggina unicornis 7 — — * — * * — * — — — —
Limacodidae Beggina zena 12 — — * — * * * — * — * —
Noctuidae Aegilia vitiscribens 5 * — — — — — — * — — * *
Noctuidae Agrotis ipsilon 1 — — — — — — — — — — * —
Noctuidae Callopistria maillardi 3 — — — — — — — — — * — —
Noctuidae Chasmania tibialis 1 — — — — — — — — — — * —
Noctuidae Chrysodeixis eriosoma 57 * * * * * * * * — * * *
Noctuidae Condica conducta 2 — — — — — — — — — * * —
Noctuidae Condica illecta 90 * * * * * * * * — * * *
Noctuidae Dactyloplusia impulsa 1 — — — — — — — — — — * —
Noctuidae Gyrtonia purpurea 1 — — — — — — — — — — * —
Noctuidae Leucania venalba 1 — — * — — — — — — — — —
Noctuidae Leucania yu 4 — — — — * — * — — * * —
Noctuidae Penicillaria jocosatrix 1 — — — — — — — — — — * —
Noctuidae Rusicada nigritasis 2 — — — — — — * — — — — —
Noctuidae Sasunaga oenistis 1 — — — — — — — — — * — —
Noctuidae Sasunaga tomaniiviensis 1 — — — — — — * — — — — —
Noctuidae Spodoptera litura 29 * * * * * * * * — — * —
Noctuidae Spodoptera mauritia 87 * * * * * * * * * — * *
Noctuidae Stictoptera stygia 17 * * * * * * — * * — * —
Noctuidae Stictoptera vitiensis 11 — — * — * — — * — — * —
Noctuidae Tiracola plagiata 27 — — — — * — * * * * * —
Nolidae Austrocarea albipicta 1 — — — — — — — — — * — —
Nolidae Barasa triangularis 2 — — — — — — — — — — * —
Nolidae Earias flavida 9 — — — * * — — — — — * —
Nolidae Maceda mansueta 22 * * * * * * — * — * — —
Nolidae Maceda savura 8 * — — — * — — — — — * —
Temporal Patterns in Fijian Forest Moths · Tikoca et al.461
Appendix (continued)
Family Species N D J F M A M J J A S O N
Sphingidae Daphnis placida 1 * — — — — — — — — — — —
Sphingidae Macroglossum godeffroyi 2 — — — — — — — * — — * —
Sphingidae Macroglossum hirundo 1 — — — * — — — — — — — —
Thyrididae Banisia anthina 27 * * — — * * * * — * — *
Thyrididae Banisia messoria 4 * — — — — — * * — * — —
Thyrididae Striglina navigatorum 13 * * — * * — * * — * * —
Uranidae Epiplema simmondsi 3 — — — * — — — — * — — —
Uranidae Urapteroides anerces 46 * * * * * — * * * * * *
Yponomeutiidae Atteva aleatrix 19 * — * — — — * * * * — —