Estacionalidad de Insectos PDF

Seasonality of Tropical Insects
Author(s): Henk Wolda

Source: Journal of Animal Ecology, Vol. 49, No. 1 (Feb., 1980), pp. 277-290
Published by: British Ecological Society
Stable URL: http://www.jstor.org/stable/4289
Accessed: 10-05-2016 23:29 UTC
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Journal of Animal Ecology (1980) 49, 277-290
SEASONALITY OF TROPICAL INSECTS

I. LEAFHOPPERS (HOMOPTERA) IN LAS CUMBRES, PANAMA
BY HENK WOLDA
Smithsonian Tropical Research Institute, P.O. Box 2072, Balboa, Panama
SUMMARY
(1) The seasonal distribution of tropical insects was studied using a light trap in Las
Cumbres, Panama, over a period of 3- years.
(2) The phenology-measures Seasonal Range (SR) and Seasonal Maximum (SM) are
used here for the first time.
(3) Some 370 % of the species occur throughout the year. But almost all species, with
few exceptions, do have clearcut seasonal peaks in abundance.
(4) The results are compared with data on Homoptera from Finland, England and the
USA. Seasons as short as those commonly found as far north as Northern Finland also
occur in the wet tropics of Las Cumbres, Panama.
(5) Most species have their seasonal peak in abundance some time during the rainy
season. The concentration of peaks in the early dry season (January-February) is caused
by increased activity of species leaving the grass that is drying out, in search of alternative
food.
(6) Some of the seasonal peaks occurring at the very beginning and at the end of the
rainy season refer to dispersing insects, including some species which do not normally live
in the vicinity of the trap.
INTRODUCTION
The occurrence of seasonal fluctuations in the number of individuals in tropical insect
species is a well established fact (for references see Wolda 1978a, 1979a). However, it is
far from clear how such fluctuations compare with those found in other parts of the
world with different climatic regimes. On the basis of climatic data one might expect
tropical species to have longer seasons than their counterparts at higher latitudes. Many
species may be expected to occur around the year, or at least throughout the favourable
season, which usually is the rainy season. One might expect many of the species that do
occur around the year to have little, if any, seasonal variation in abundance, as shown for
instance for some butterflies (Ehrlich & Gilbert 1973; Owen & Chanter 1972). However,
is this true? The published data on seasonality in the tropics are insufficient to allow for a
general test of these, or any other, hypotheses concerning the seasonality of tropical
insects.
For such a test, the seasonal distribution of a large number of tropical species would
have to be documented and these data would have to be analysed quantitatively so that
direct comparisons can be made between different areas in the world. The present paper
will present such an analysis for Homoptera from Las Cumbres, Panama.
0021-8790/80/0200-0277$02.00 (?1980 Blackwell Scientific Publications
277
278 Seasonality of Homoptera in Las Cumbres, Panama

The insects are collected with a light trap. I believe that for many species these data
reflect reasonably well the real fluctuations in abundance in nature, that is of the active
adults of those species. In some instances I do have general observations supporting this
contention, such as for cicadas, where monitoring the songs gives an impression of
seasonal presence which coincides nicely with the light trap data. For a further discussion
see Wolda 1978a, b.
Attempts have been made in the literature to capture some aspect of the seasonal
distribution of a species in a single parameter. Slobodchikoff & Parrott (1977) used a
descriptor called 'Seasonal Niche Breadth' (Levins 1968). Unfortunately, this measure is
useful only when the information on phenology for each species is based on a large
number of individuals (Wolda 1979b). For smaller sample sizes the measure is practically
useless. I used 'Season Length' (Wolda 1977), which is much better than the Seasonal
Niche Breadth, but still dependent on sample size (Wolda 1979b). I proposed a measure
called 'Seasonal Range' (SR), which is an estimate of the length of the season in which the
phase of the insect under consideration, in this case adults, are present in a given year,
corrected for sample size (Wolda 1979b). This measure SR will be used together with a
new one called 'Seasonal Maximum' (SM) to analyse the light trap data on Homoptera
from Las Cumbres, Panama and to compare them with data from Kansas, USA (Blocker,
Harvey & Launchbaugh 1972 and personal communication), from England (Waloff
1973) and from Northern Finland (Kontkanen 1950).
PROCEDURES
Locality and trapping
The data to be discussed were collected using a light trap of the Pennsylvania type,
modified for the tropics by my colleague Nicholas Smythe. In this trap the insects are
attracted by a Sylvania Fl 5T8 BL fluorescent light, fall through a funnel in a cloth bag
where they are killed by carbon tetrachloride. The trap was in operation all night, every
night, from October 1973 to March 1977. The insects were either sorted immediately or
stored in a freezer for later sorting. Several groups of insects have been sorted out and
many of these have been analysed by specialists to the species level. The present paper only
deals with the Homoptera, sorted into species by my assistant Mr Miguel Estribi and
myself. Other groups will be dealt with elsewhere.
For the vast majority of the homopteran species the classification should be correct.
We received considerable help from various specialists when taxonomic problems needed
to be resolved. It is possible that in some cases we may have to split, or lump, species as
the taxonomy of the groups is re-examined. For instance, the small green Empoasca
(Typhlocybinae) are all counted as just one taxon although there probably are several
species represented.
The trap was located in Las Cumbres, a town some 15 km North of Panama City,
Republic of Panama, at 9?5'36"N and 79?3 '54" W. The trap was on top of a slope. South
and west of the trap there are some bungalows, surrounded by trees, shrubs and lawns.
To the east and north the trap overlooks a valley with many wild, ornamental and fruit
trees. Fifty metres in that direction is the edge of a young second growth forest. The
elevation is 150 m above sea level.
Mean annual rainfall is 1500 mm, 84% of which falls from May to November and 900%
from April to November. The dry season usually starts some time in December and lasts
H.
WOLDA
279
30- (a)
20
I0
(b)
400
300
200
100
-11;2
1974
llL:1
1975
II
IF
1976
I,
FIG. 1. (a) The number of rainy days per month in Las Cumbres, Panama, and (b) rainfall
in mm during the study period.
until April or May. Towards the second half of January the effect of the dry season begins
to show in that the lawns and roadside grasses begin to dry and turn brown. The rainfall
per month for the study period, together with the number of days with rain per month, is
plotted in Fig. 1 (data from Caballero 1974, 1975, 1976, 1978). No data are available yet
from that source for 1977, but only little rain fell in January 1977 and February and
March of that year were completely dry.
The mean temperature per month is 27 ?C, with no apparent seasonal fluctuation. The
mean maximum temperature is about 31 ?C with the maxima in March and April, the
last months of the dry season, generally somewhat higher than during the rest of the year.
The mean minimum temperature is 230 C and tends to be somewhat lower in the dry
season than during the rainy season.
Measures of seasonality
Rather than trying to capture many aspects of the seasonal distribution in one single
parameter, as the Seasonal Niche Breadth pretends to do, I selected two aspects of the
seasonal distribution and tried to represent each by a parameter. This procedure seems
less confusing. The aspects selected are the length of the time, per year, that a species is
present and the relative height of the seasonal peak. These two aspects are not completely
independent, but sufficiently so to warrant two parameters instead of just one. For an
area where all species have short seasons, one parameter would be sufficient, but for
species which occur around the year it can be very important to be able to distinguish
between species which have a very sharp seasonal peak and those which show no seasonal
variation in abundance at all.
The Seasonal Range (SR) is determined by measuring the length of the season (SL)
in weeks. With the help of a simulation model based on a seasonally normal distribution,
the SL is transformed to SR, thus eliminating, at least to some extent, the effect of sample
size. For details on how the transformation from SL to SR is carried out, see Wolda
(1979b).
The Seasonal Maximum (SM) is determined as follows. In a year there are 13 periods of
4 weeks. The mean number per 4 weeks is determined simply by dividing the annual
total by 13. The running 4-week sum of the data is taken and the 4-week period with the
highest number of individuals is called the maximum. Seasonal Maximum is this maximum
number divided by the mean. Seasonal Maximum varies from 13, when all individuals
are found within one period of 4 weeks, to one, when the number of individuals is absolutely constant throughout the year. For actual data, however, because of statistical
variations both in the actual populations and in the sampling process, even in species
which do not have any biologically significant seasonal variation in abundance, SM will
always be larger than unity.
As one might expect, SM is also dependent on sample size. This can be demonstrated
by taking computer-drawn samples of various sizes from seasonally normal distributions,

with overlapping tails, with various seasonal standard deviations (SSD). An SSD of 1
means a very sharp seasonal peak, an SSD of 20 describes a distribution which is effectively
rectangular, i.e. non-seasonal. For further details of these distributions see Wolda 1979b.
For each SSD and each sample size ten replicate samples were obtained, which gives a
mean and a standard deviation for SM. For a number of SSD values the mean value of
SM is plotted against sample size in Fig. 2. The lines given are actually lines drawn by
eye through the calculated points. The peculiar bump in the line for SSD = 2 could be a
sampling error. For anyone who wishes to use these same lines the data describing them
are available on request. For smaller samples the effect of sample size is considerable.
Actual data would be represented by points scattered over the area of Fig. 2 and the lines
classify the points in groups, such as the ones above the line for SSD = 1, the ones
between the lines for SSD = 1 and SSD = 2, etc. The distribution of the points over
these groups describes the data set in terms of the sharpness of the seasonal peak,
eliminating, at least to a certain extent, the effect of sample size. The groups associated
with the lines for low values of SSD indicate sharp seasonal peaks, the ones with high
values for SSD refer to less pronounced seasonal variation.

Admittedly, the lines in Fig. 2, which are used for all data sets, are based upon a
seasonally normal distribution, while an actual distribution may deviate from normality,
sometimes even considerably so. However, I doubt that such deviations from normality
affect the shapes and position of the lines in Fig. 2 in any serious way.
r w _ _ I Z 155~~~~~~~5D
10
E
I0
_2
.E_
C:
U)
4
7
20
10
100
1000
10000
Sample size
FIG. 2. The relations between the Seasonal Maximum (SM) and sample size in computerdrawn samples from seasonally normal distributions with standard deviations (SSD)
varying from 1 to 20.
H.
WOLDA
281
RESULTS
Species richness
A total of 86 039 individual Homoptera were caught, representing about 540 species.
The distribution of the Homoptera over the various families is given in Table 1, with the
Cicadellidae given per subfamily. Some 750 of the individuals and about 5000 of the
species are Cicadellidae, as seems to be the case in more open environments in Panama.
In forests Fulgoroidea are usually more abundant. About 2000 of the species can be
classified as common, i.e. they were caught with a hundred individuals or more, while
about the same percentage (107 species) is represented by one single individual only. This
shows that after 31 years of intensive trapping many species present have not been
caught at all. The number of 540 species represents only a minimum figure for the total
number of species within reach of the trap.
TABLE 1. Homoptera collected with a light trap in Las Cumbres, Panama, over
a period of three and a half years
Common
species
Individuals Species N 2 100
Cicadidae
1834
10
Membracidae 231 30
Cercopidae 83 11
Psyllidae 93 15
Cicadellinae 19118 35 10
Agalliinae 6066 10 4
Xestocephalinae 9595 11 6
Hecalinae 1 1
Nirvaniinae 4 2
Ledrinae
2
1
Deltocephalinae 8834
Typhlocybinae 13375
Idiocerinae 1168
Gyponinae 4903
Neocoelidiinae 66 5
Coelidiinae 13 4
Achilidae
4598
70 19
89 17
13 2
33 5
21
10
Cixiidae 2218 32
Acanaloniidae 18 2
Achilixiidae 110
Fulgoridae 60 2
Nogodinidae 21 1
Kinnaridae 15 3
Flatidae 3294 23
Derbidae 3350 56 5
Delphacidae 6649 47 14
Tropiduchidae 6 2
Dictyopharidae
Total
86039
314
540
10
110
Seasonal maximum and seasonal range
Some examples of seasonal distributions are given in Fig. 3. These range from a very
short season in Amblyscarta resolubris (Fowler) to a virtually constant distribution in
Polana scinna DeLong and Freytag. All these examples have their maximum, if there is one,
in the wet season. In Fig. 4 some aberrant species are given. Pacarina signifera Walker, a
common cicada, is typically a dry season species. Muirolonia metallica Fowler has a
bimodal distribution while Pintalia sp. 62B has three peaks per year.

10_(a)
0S
25 (b)
7-~~~~~~
LLA
100 _ (d)
100 (e)
1974
1975
1976
77
FIG. 3. The seasonal distribution, in numbers per 4 weeks, for some species of Homoptera
in Las Cumbres, Panama. (a) Amblyscarta resolubris (Fowler) (Cicadellinae). (b) Myconus
uniformis (Metcalf) (Achilidae). (c) Muirilixius banksi Metcalf (Achilixiidae). (d)
Xestocephalus tessellatus Van Duzee (Xestocephalinae). (e) Polana scina DeLong and
Freytag (Gyponinae).
For each year, every species which is represented by at least six individuals is used in the
analysis. There are 3 complete years, so that each species may contribute up to three data
points. The 246 species used give a total of 637 data points.
The values for SR are plotted, per group, in Fig. 5. A total of 235 points, 36.9%, are
52, which means that the species occurred the year around. Many species have shorter
seasons, or even very short seasons, much shorter than one might expect from the
climate. It is just possible that in some cases this is because the species does not live near
the trap and is only caught on its annual dispersal flight. I cannot rule out this possibility.
Most known dispersers that do not live near the trap are caught twice per year, making
the season rather long instead of short, once around May and once towards the end of
the year. In many cases, however, the shortness of the season is undoubtedly real.
25
t 25 - (b)
0rw 50-(c) A
1974
1975
1976
77
FIG. 4. The seasonal distribution, in numbers per 4 weeks, of some species of Homoptera.
(a) Pacarina signifera (Walker) (Cicadidae). (b) Muirolonia metallica Fowler (Cixiidae).
(c) Pintalia sp. '62B' (Cixiidae).
H.
WOLDA
283
10 (a)
N=17
10 -(b)
5.9%
_N=
13
4
I I I rr1 .-. 30 8%
10 ( c) N= 3 (unshaded)
_ N =2 (shaded)
20 - (d)
F r N- 188 36.7%
(0~~~~~~~~~~~~~~~5
F N= 69 ilrl flYEfltfKFl.,.r
10 (e)2023
N= 637
20
o/5
Seasonal Range
FIG. 5. Distribution of the values for the Seasonal Range (SR) in some groups within the
Homoptera. (a) Cicadidae, (b) Membracidae, (c) Cercopidae: (unshaded) Psyllidae:
(shaded), (d) Cicadellidae, (e) Fulgoroidea-Delph.; (f) Delphacidae, (g) Homoptera total.
The frequency of species occurring throughout the year is given in numbers and percent-
ages rather than in histograms. Data from Las Cumbres, Panama.
The 637 values for SM are plotted against sample size in Fig. 6. The size of each point
is directly proportional to the number of data points it represents. The lines are explained
above. The distribution of these data points over the spaces between these lines is given,
in percentages, in Table 2. There is a wide range of values, ranging from very sharp peaks
to no peaks at all, as illustrated in Fig. 3. Most cases are seasonal distributions which
resemble normal distributions with a seasonal standard deviation between 4 and 11. There
are a few species which do not show any seasonal variation in abundance, but those are
in the minority, even among the species that are present as active adults around the year.
Several species have well-defined sharp seasonal peaks. Among these there are some
where this peak is an increase in activity, rather than a maximum in abundance. This is
the case for some species which live in the lawns and start dispersing in search of suitable
places to live in the early dry season. Good examples of this are Plesiommata mo/lice/la
(Fowl.) (Cicadellinae) and A gal/ia modesta Osb. and B. These, and some other similar
species make up most of the points in the upper right hand corner of Fig. 6. In many
cases, however, the sharp peaks undoubtedly represent such sharp peaks in abundance
in the real world.
There tend to be differences between families in the average seasonal distribution of

their species (Fig. 5; Table 2). Cicadas have a rather short season, which is confirmed by

_
'
155D
10 a
E
E* . a ... :- .-.. * 7
En
o5
=p
20
10
100
1000
10000
Sample size
FIG. 6. Plot of values for Seasonal Maximum (SM) v. sample size for Homoptera from
Las Cumbres, Panama. The size of each plot is proportional to the number of data points
it represents. The simulation lines of Fig. 2 are also included.
TABLE 2. Seasonal Maximum (= sharpness of the seasonal peak in abundance)

for leafhoppers (Homoptera) from Las Cumbres, Panama as compared with
Homoptera from Kansas, England and Finland. The distribution of the species
over the various categories is given in percentages. The categories range from
a very sharp seasonal peak on the left to the virtual non-existence of any
obvious peak on the right. The number of data on which the percentages are
based are given (N). For further details see text
SSD-classes of SM < 1 1-2 2-4 4-7 7-11 11-20 > 20 N
Las Cumbres, Panama
Cicadidae 5-9 17.6 47 23.5 5.9 17

Membracidae - 7.7 30.8 23.1 15 4 23-1 13
Cercopidae
667
33-3
3
Psyllidae
100
2
Cicadellidae 0-6 3-8 11.6 36-2 29.6 15-4 2 9 345
Fulgoroidea-Delphac. 1 6 12.2 46-3 30 8 6.4 2-7 188
Delphacidae 4.3 31.9 42 11-6 5-8 4-3 69
Total Las Cumbres 0.5 4.1 14-7 39.1 27 11-3 3-3 637
Kansas, USA 3-8 25 40-4 23-1 5-8 1.9 - 52
England 9-1 32.7 43-6 12.7 1-8 55
Finland 23.2 58.1 16.3 2.3 43
data on their singing. The one species listed here as having an SR of 52 I tend to interpret
as an error in estimation. In 1975 the species Herrera marginella (Walker) was caught
with 6 individuals only over a period of 25 weeks. Using my standard procedure (Wolda
1979b) this gives an SR of 52. With samples this small some errors are bound to appear.
I think this is one instance in which the 25 is real, the 52 is not. The SR values are grouped
in classes 0-10, 10-20,. . ., 40-51.9, and 52 and differences between groups are tested by
x2. The same test is used to compare groups in the distribution of the SM values. The
Cercopidae and Psyllidae are left out of the comparisons as the number of data points is
too low. The differences between Membracidae. Cidadellidae and Fulgoroidea (Minus
the Delphacidae) are not significant, or only barely so at the 5%o level. The Delphacidae
H.
WOLDA
285
13 5
I0-
26
~~~~55
127
189
235
20
30
40
50
Seasonal Range
FIG. 7. The relation between SR and SM for the Homoptera from Las Cumbres, Panama.
Horizontal lines indicate the range of SR values used for calculating a given point, and the
mean value of SM for those data. The vertical lines are at the mean value of SR for that
subset of data and indicate one standard deviation in each direction. The numbers indicate
the number of data points represented.
have significantly shorter seasons and significantly sharper seasonal peaks, on the
average, and the Cicadas again have significantly shorter seasons, even than the Delphacidae, and sharper seasonal peaks.
As mentioned before, SR and SM, although far from identical, are correlated measures
of seasonality. For the 637 Homoptera data points the relation between SR and SM is
shown in Fig. 7. The SR values are grouped as indicated by the horizontal lines. These
lines are drawn at the level of the mean value of SM for these data (SM here is not
corrected for sample size). At the mean value for SR within each group, a vertical line
indicates one standard deviation of SM in each direction. The numbers indicate how
many data points there are in each group. There is a clear relation between SR and SM,
but the fairly large standard deviations in SM indicate that the knowledge of only one of
the two parameters has only limited predictive value for the value of the other.
An important problem in an analysis such as this is presented by those species that are
multimodal in distribution. Some of the more common species show two or more welldefined peaks in abundance per year (Fig. 4). In other cases there could be more than
one peak but the picture is not entirely clear. Some of the less common species seem to
have more than one peak, but the interpretation of such data is rather subjective. In
some cases the light trap data simply are not good enough to demonstrate the number of
peaks (generations) per year. A good example is Aeneolamia postica (Walker) (Cercopidae). It was caught in the present trap only occasionally, but at the lights of a house
nearby, on a wall facing a grassland where the species occurs in large numbers, the species
is very common. The distribution per day is given in Fig. 8, indicating some six or seven
peaks during the rainy season. Such a resolution can only be obtained for species that
are very common in the trap. The maxima in abundance shown in Fig. 8 are real and not
an artefact produced, for example, by phases of the moon.

300
200
I oo9
50
25
0- vZ
--7 a
II I1
~ ~MI
~ J~~?iL
l
:X: :x:1
I X[ I I 1 Ev
1972
1973
FIG. 8. The seasonal distribution, in number of individuals per day, of Aeneolamia

postica (Walker) (Cercopidae) from the wall lights of a house not far from the light trap.
The timing of the seasonal maxima
I have shown that many species do not occur throughout the year, at least not as active
adults, and of those that do, the vast majority has more or less well-defined seasonal peaks
in abundance. The question arises whether the seasonal peaks are in any way related to
the alternation of the wet and dry seasons. It is also of interest to see whether the seasonal
peaks of the different species all tend to occur during the same part of the year or whether
there is a tendency for the peaks to be spaced out, which would suggest that there are
strong interactions between some of the species.
In most cases it was not difficult to decide which week in the year was the peak week.
The maximum 4-week period used to calculate SM was taken and the week with the
highest number within that 4-week period is the peak used here. For several of the smaller
samples this did not work so well and then other criteria were applied, e.g. when there
was no one clear maximum and the sample was too small to decide about bimodality, the
meanpoint of the season was used. For species which, in my subjective judgment, were
clearly multimodal, all peaks were used in the analysis. The distribution of all the peaks
so obtained is given in Fig. 9 both for each year separately and for the 3 years combined.
There are very few peaks in abundance in the dry season. The large number of peaks
indicated in Fig. 9 in the early dry season, i.e. weeks 1 to 5, is mostly not real. That is,
they refer largely to species which lived in the lawns during the rainy season and now find
themselves without food in the dried out grassy areas and start flying around in large
numbers looking for suitable places to live. These maxima are peaks in activity rather
than peaks in abundance. In a number of cases I cannot be sure yet whether the suspected
peak in activity is that or is a real peak in abundance, so I have not weeded out the species
which have a maximum in the trap which does not correspond to a maximum in abundance in the field. Part of the irregularities in the pictures for each year is due to the phases
of the moon. At full moon many species have a low abundance in the trap and thus fewer
species will have their maximum abundance during such weeks with full moon. The
general picture obtained from Fig. 9 is that the peaks tend to be spaced all over the rainy
season. Some of the peaks at the very beginning and some at the end of the rainy season
refer to seasonal dispersers, i.e. species that tend to have dispersal flights during those
times of the year and that do not necessarily live in the vicinity of the trap. The other
H.
WOLDA
287
20 (a
I 0~~~~~~~~~~~~~~~~~~
20 (b)
20o
ILi
r
a))
-Lz
50 (d)
40 30
20-
IP
I0
10
20
30
40
50
Week of maxima
FIG. 9. The distribution of the seasonal peaks over the years. See text for details. (a) 1974,
(b) 1975, (c) 1976, (d) 1974-1976.
species, then, may have a slight tendency to have their annual peaks during the middle of
the rainy season, but in general the peaks seem to be staggered rather than concentrated,
which suggest fairly strong interactions between the species involved, forcing them to
have their peaks spaced out. The dry season, apparently, is so inimical to most Homoptera
that they cannot have their seasonal peaks during that season, away from all the other
peaks of the rainy season. Only some species have been able to overcome this problem.
Geographical comparisons in seasonality

The measures SR and SM permit direct comparisons between different geographic
areas for which data are available. The ones discussed in the present paper from Las
Cumbres can be compared with data from Northern Finland (Kontkanen 1950), from
Kansas, USA (Blocker, Harvey & Launchbaugh 1972 and personal communication) and
from England (Waloff 1973).
The values for SR are plotted in Fig. 10. The one large value of SR = 50 for Kansas I
assume to be an error of estimation like the one discussed before, as the sample size for
that case was very small (n = 8). The differences between the four distributions are
highly significant as tested by x2 after grouping of the data. In Northern Finland the
length of the season for flying adult Homoptera is, on the average, very short, much
shorter than for most tropical species. In England and Kansas the seasons are of intermediate length. Very long seasons, or species which occur throughout the year, are found
in the tropics only. However, short seasons, as short as the ones found in the subarctic,
are also found in the tropics of Las Cumbres, although there they are relatively rare. Data
I0 ? (ab)
10I(b)n n u- A Iol|
n
In1
rr,
10c
2o0
(d)
235
____ ___ ____ ___ ___ ____ ___ ___ 36-9 %
10
20
30
40
50
52
Seasonal Range
FIG. 10. The distribution of SR values in four samples of Homoptera ranging from the
subarctic of Northern Finland to Las Cumbres in tropical Panama. (a) Finland, (b)
England, (c) Kansas, (d) Las Cumbres.
from the salt marshes of North Carolina (Davis & Gray 1966) suggest that the seasons
of the Homoptera there may be as long as they are in the tropics (Wolda 1977) but the
data do not permit calculation of the SR values as sample sizes are not given.
The distribution of the species over the SM classes is given in Table 2. England and
Kansas are here not significantly different from each other, but all other differences are
highly significant. Sharp seasonal peaks predominate in Finland, but are also found in the
other areas, including the tropics. Although, on the average, the peaks in Las Cumbres
are much less pronounced than those of Finland, with Kansas and England occupying
intermediate positions. Unfortunately, the data given by Davis & Gray (1966) do not
permit calculations of the SM values for the Homoptera of the 'subtropics' of North
Carolina.
Annual Variability
Changes in abundance from year to year can best be discussed in terms of a parameter
called Annual Variability (AV, see Wolda 1978b). This is, briefly, the variance of the
distribution of log R, where R, indicates the change in abundance (Nt+1/Nt) for the ith
species. As a measure for abundance in a given year the total number of individuals
caught in the trap is used. It is usually advisable to include only those data where the
smallest of the two N-values is at least five (Wolda 1978b).
For the years 1974/75 AV for the Las Cumbres Homoptera is 0. 103 (N = 197) and for
1975/76 AV is 0. 115 (N = 201). These values are what one would expect for data from
humid areas, tropical or temperate (Wolda 1978b).
As I have discussed before (Wolda 1978b), one might expect species with a sharp
seasonal peak to be more variable in abundance from year to year, to give larger values of
AV, than species which do not have such sharp peaks, or do not have clear maxima at all.
Some data, including part of the data discussed here, supported that hypothesis. The Las
Cumbres data in their entirety are now re-analysed to test this hypothesis. Annual
H.
WOLDA
289
Variability is calculated for those species which have, in 2 successive years, values of
SM less than 4 and for those species which have, in 2 successive years, a value of SM
larger than 6.5, or, if the sample was small, larger than 7. Otherwise the effect of sample
size on SM is ignored here. The latter species have relatively sharp seasonal peaks, the
former species do not. The latter group has an AV of 0.176, the former has an AV of
0.075. The difference is highly significant, again supporting the hypothesis.
DISCUSSION
The seasonal distribution of insects, or of other organisms, is usually discussed with the
aid of long tables or long series of figures of various kinds. Several examples can be found
in Lieth (1974). Without some kind of a formal analysis of such data, however, the
interpretation of the results is rather difficult. To the best of my knowledge the present
paper is a first attempt at a quantitative comparison between insects from the subarctic,
the temperate zone and the tropics in terms of their seasonal distribution. Some of the
results are hardly surprising. One might have expected to see that many tropical insects
occur throughout the year as active adults, while subarctic insects have very short active
seasons. However, up to now that expectation was not much more than a hypothesis and
seasonality at different latitudes has not been properly documented. The present data,
to a large extent, conform to expectation. However, there are also some surprises. The
fact that seasons as short as those found in the subarctic also occur in the tropics is
unexpected. The dry season, inimical to most species, has a strong effect on the seasonal
distribution. However, species which are effectively non-seasonal, although uncommon,
do occur in Las Cumbres in spite of the existence of a strong dry season. Some species
manage to exploit the dry season conditions such that they have their annual maximum
during that season. A few even occur in the dry season exclusively and are absent during
the rainy months of the year.
An explanation of the various patterns of seasonality found in the tropics at this stage
would be premature. For the vast majority of the species dealt with in the present paper,
the life history is completely, or almost completely, unknown. In the absence of information on foodplants of nymphs and adults, on predators, competitors, etc. I prefer to
postpone speculations, however easy these would be when there are no data to set limits
to one's imagination. The distribution of the seasonal peaks as given in Fig. 9 suggests
that interspecific interactions of some kind are an important factor in the determination
of the seasonal distribution of Homoptera in the tropics.
ACKNOWLEDGMENTS
It is a pleasure to express my gratitude to Miguel Estribi and Saturnino Martinez who

helped to sort the insects and to my wife Tineke who took care of the trap whenever I
was not there. The help received from Michel Boulard (Paris, France), Lois O'Brien
(Tallahassee, Florida), David Young (Raleigh, North Carolina) and Dwight DeLong
(Columbus, Ohio) in the identification of species is gratefully acknowledged. I am
particularly grateful to Derrick Blocker (Manhattan, Kansas) for sending me the only
partly published data for use in my comparisons.
REFERENCES
Blocker, H. D., Harvey, T. L. & Launchbaugh, J. L. (1972). Grassland leafhoppers. I. Leafhopper
populations of upland seeded pastures in Kansas. Annals of the Entomological Society of America,
65, 166-172.

Caballero, J. M. (1974). Meterologia, afno 1973. Estadistica Panamenla, 34, Serie L.
Caballero, J. M. (1975). Meteorologia, afio 1974. Estadistica Panamefna, 35, Serie L.
Caballero, J. M. (1976). Situaci6n fisica, meteorologia, afno 1975. Estadistica Panamenta, seccion 121,
clima.
Caballero, J. M. (1978). Situaci6n fisica, meteorologia, afio 1976. Estadistica Panamefna, seccion 121,
clima.
Davis, L. V. & Gray, I. E. (1966). Zonal and seasonal distribution of insects in North Carolina salt
marshes. Ecological Monographs, 36, 275-295.
Ehrlich, P. R. & Gilbert, L. E. (1973). Population structure and dynamics of the tropical butterfly
Heliconius ethilla. Biotropica, 5, 69-82.
Kontkanen, P. (1950). Quantitative and seasonal studies on the leafhopper fauna of the field stratum
on open areas in North Karelia. Annales Zoologici Societatis Zoologicae Botanicae Fennicae
'Vanamo', 13, 1-91.
Levins, R. (1968). Evolution in Changing Environments. Princeton University Press, Princeton.

Lieth, H. (1974). Phenology and Seasonality Modelling. Springer, New York, Heidelberg, Berlin.
Owen, D. F. & Chanter, D. 0. (1972). Species diversity and seasonal abundance in Charaxes butterflies
(Nymphalidae). Journal of Entomology, Series A, 46, 135-143.
Slobodchikoff, C. N. & Parrott, J. E. (1977). Seasonal diversity in aquatic insect communities in an allyear stream system. Hydrobiologia, 52, 143-151.
Waloff, N. (1973). Dispersal by flight of leafhoppers (Auchenorrhyncha: Homoptera). Journal of

Applied Ecology, 10, 705-730.
Wolda, H. (1977). Fluctuations in abundance of some Homoptera in a neotropical forest. Geo-EcoTrop, 3, 229-257.
Wolda, H. (1978a). Seasonal fluctuations in rainfall, food and abundance of tropical insects. Journal of
Animal Ecology, 47, 369-381.
Wolda, H. (1978b). Fluctuations in abundance of tropical insects. The American Naturalist, 112,
1017-1045.
Wolda, H. (1979a). Abundance and diversity of Homoptera in the canopy of a tropical forest. Ecological
Entomology, 4, 181-190.
Wolda, H. (1979b). Seasonality parameters for insect populations. Researches on Population Ecology,
20, 247-256.
(Received 4 May 1979)

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Seasonality of Tropical Insects

Author(s): Henk Wolda

Journal of Animal Ecology (1980) 49, 277-290

SEASONALITY OF TROPICAL INSECTS

Smithsonian Tropical Research Institute, P.O. Box 2072, Balboa, Panama

The occurrence of seasonal fluctuations in the number of individuals in tropical insect

278 Seasonality of Homoptera in Las Cumbres, Panama

1973) and from Northern Finland (Kontkanen 1950).

Locality and trapping

280 Seasonality of Homoptera in Las Cumbres, Panama

by taking computer-drawn samples of various sizes from seasonally normal distributions,

values for SSD refer to less pronounced seasonal variation.

Seasonal maximum and seasonal range

282 Seasonality of Homoptera in Las Cumbres, Panama

ages rather than in histograms. Data from Las Cumbres, Panama.

There tend to be differences between families in the average seasonal distribution of

284 Seasonality of Homoptera in Las Cumbres, Panama

TABLE 2. Seasonal Maximum (= sharpness of the seasonal peak in abundance)

Cicadidae 5-9 17.6 47 23.5 5.9 17

286 Seasonality of Homoptera in Las Cumbres, Panama

FIG. 8. The seasonal distribution, in number of individuals per day, of Aeneolamia

The timing of the seasonal maxima

Geographical comparisons in seasonality

288 Seasonality of Homoptera in Las Cumbres, Panama

____ ___ ____ ___ ___ ____ ___ ___ 36-9 %

It is a pleasure to express my gratitude to Miguel Estribi and Saturnino Martinez who

Blocker, H. D., Harvey, T. L. & Launchbaugh, J. L. (1972). Grassland leafhoppers. I. Leafhopper

290 Seasonality of Homoptera in Las Cumbres, Panama

Levins, R. (1968). Evolution in Changing Environments. Princeton University Press, Princeton.

Waloff, N. (1973). Dispersal by flight of leafhoppers (Auchenorrhyncha: Homoptera). Journal of

(Received 4 May 1979)

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__ _ __ _ _ _ ___ 36-9 %