Biological Journal of the Linnean Socieb (1987), 30: 313-323.

With 3 figures

Altitude, habitat and tropical insect diversity

HENK WOLDA

Smithsonian Tropical Research Instihie, P.O. Box 2072, Balboa, Republic of Panama

Accepted for publication 17 November 1986

Sample size and species richness of a series of light-trap samples taken in a number of localities in
the Republic of Panama are given. Species richness as well as sample size decreased gradually with
increasing altitude over a 10G2200 m range, in contrast to data from the literature which
demonstrated a maximum at intermediate elevations. It is suggested that differences in technique,
especially the continuous nature of the sampling in the present case are responsible for this
discrepancy. Moderate human disturbance, which leaves a relatively high tree species diversity in
the area has a surprisingly little, if any, effect on insect diversity. A large perturbation of the
environment does, however, decrease species richness of the insect fauna.

KEY WORDS:-insects - tropics - diversity - altitude - habitat - human disturbance.

CONTENTS
Introduction . . . . . . . . . . . . . . . . . . . 313
Procedures . . . . . . . . . . . . . . . . . . . 314
Localities . . . . . . . . . . . . . . . . . . . 314
Insect groups . . . . . . . . . . . . . . . . . . 315
Diversity . . . . . . . . . . . . . . . . . . . 316
Results . . . . . . . . . . . . . . . . . . . . 316
Discussion . . . . . . . . . . . . . . . . . . . . 319
Altitude effect. . . . . . . . . . . . . . . . . . 319
Human disturbance effect . . . . . . . . . . . . . . . 321
Acknowledgements . . . . . . . . . . . . . . . . . 322
References. . . . . . . . . . . . . . . . . . . . 322

INTRODUCTION

Studies of insect diversity (cf. Mound & Waloff, 1978) become especially
fascinating when including the rich tropical insect faunas (Janzen, 1976;
Prance, 1982). It has become clear that insect groups tend to have many more
species at lower than at higher latitudes, with the exception of bees (Michener,
1979) and some parasitic Hymenoptera (Gauld, 1987). Such a statement,
however, is vague because ecologically there is not just one ‘tropics’. Depending
upon the local rainfall regime an area within the lowland tropics may be a
desert, a savannah, a dry forest or a humid rainforest and one cannot profitably
discuss ‘the tropics’ without specifying the habitat. Altitudinal gradients are yet
another source of between habitat variation and so are disturbances caused by
313
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314 H.WOLDA
human activities. Altitude and disturbance will be the focus of attention in the
present paper.
In birds, a gradual decrease in species richness has been observed along a n
altitudinal gradient in Costa Rica from 0 to 3000 m (Stiles, 1983) and in Peru
from 400 to 3600 m (Terborgh, 1977). In insects, however, the available studies
suggest that maximum species richness occurs at intermediate altitudes (Janzen,
1973a, b; Janzen et al., 1976; Gagne, 1979; Gauld, 1985). Other studies include
only a small altitudinal range, but the results are consistent with such an
intermediate level maximum (Turner & Broadhead, 1974; Moron & Terrbn,
1984). Hebert (1980) finds a gradual decrease in species richness of moths
between 2200 and 2800 m in Papua New Guinea, and refers to other evidence
for a similar decrease between 2000 and 3400 m. He also states that
“preliminary studies indicate that the diversity of moth communities in Papua
New Guinea shows little further increase at lower elevations”. The decrease of
species richness with altitude there may only start at higher altitudes. Claridge
& Singhrao (1978) found little, if any, change in species richness of grasshoppers
over the entire range between 300 and 1800 m on the Mont Ventoux in France.
Giinthart (1984) found a decrease of species richness with altitude of Homoptera
between 2200 and 3060 m in Switzerland, but no comparable data from lower
elevations are available.
I n most instances no complete information is available about the insect fauna
at particular sites. All one has to rely on are samples taken from that fauna in
some, usually biased, manner. The present paper is no exception, and it will
discuss species richness of a number of insect groups as measured by light-trap
samples taken in different sites, at different altitudes, in the Republic of
Panama.

PROCEDURES

Localities
The light-traps were modified “Pennsylvania” traps (Smythe, 1982), used in
the following localities in the Republic of Panama:
( I ) Miramar, Bocas del Tor0 Province, 9’N,82’15’W, sea level. A very
disturbed habitat, with pastures, the village of Miramar, and nearby abandoned
cacao plantations. Estimated annual rainfall 300 cm. No dry season, but rainfall
both in March and in September is less than in other months of the year. The
trap functioned from November 1978 to November 1979, every night from 6 to
10 p.m. (Wolda & Flowers, 1985).
(2) Corriente Grande, Bocas del Tor0 province, 9”17’30”N, 82’32’41”W,
100 m elevation. Except for some recent logging a relatively undisturbed forest
along the Changuinola river. Climate similar to that of Miramar, but estimated
annual rainfall 250 cm. The trap functioned only from January to May 1980,
every night from 6 to 10 p.m.
(3) Barro Colorado Island (BCI), Panama Canal Area,
9’9’1 9”N, 79’50‘1 5”W. Relatively undisturbed tropical forest. Annual rainfall
250 cm with a 4-month dry season (Leigh, Rand & Windsor 1982). Two traps,
one at 2-3 m above the forest floor and one in the canopy at 26-28 m. The traps
are being operated every night, all night, from late 1973 onwards.
 ALTITUDE AND INSECT DIVERSITY 315

(4) Las Cumbres, Province of Panama, 9”5’36”N, 79”31’54”W, 150 m above

sea level, a disturbed residential area with secondary forest nearby. Annual
rainfall 150 cm with a strong 4 months long dry season (Wolda, 1980). A light-
trap operated in 1974-1976 and again 1982-1984, all night, every night.
(5) Fortuna, province of Chiriqui, 8”44’N, 82”16’W, 1010 m altitude, a
relatively undisturbed forest. Annual rainfall near 500 cm with no dry season. A
trap operated here from September 1976 to July 1977 in the canopy and from
then on at a low level until July 1979, from 6 to 10 p.m. every night (Adames,
1977; McElravy, Wolda & Resh, 1982).
(6) Boquete, township of Alto Lino, province of Chiriqui, 8”48’N, 82”26‘W,
1350 m elevation, a small forest remnant surrounded by coffee plantations.
Annual rainfall 200 cm with a 4 months long dry season. A trap operated from
July 1975 to July 1978, every night, all night.
(7) Guadalupe Arriba, province of Chiriqui, 8”52’26”N, 82”33’13”, 2200 m
altitude. Wet cloud forest with the larger oak trees removed. Annual rainfall
unknown, but high, with a mild dry season of 3-4 months. A trap operated from
December 1983 to December 1985, every night from 6 to 10 p.m.

Insect groups
A number of insect groups are included in the present analysis, sorted to
species by Miguel A. Estribi (STRI, Panama), unless otherwise indicated:
Homoptera Auchenorrhyncha, Blattaria (analysed by Frank W. Fisk, Ohio
State University and Miguel Estribi), ‘Orthoptera’ (Tettigoniidae, Mantidae
and some Gryllidae) (analysed by Saturnino Martinez, STRI, Panama and
Miguel Estribi) , several Coleoptera familes, such as Pselaphidae and Anthicidae
(analysed by Donald S. Chandler, University of New Hampshire),
Curculionidae (including Anthribidae, Brentidae, Apionidae) (analysed by
Charles W. O’Brien, Florida A & M University), Cerambycidae, Scarabaeidae,
Elateridae, Eucnemidae and Chrysomelidae, the Lepidoptera family Sphingidae
(analysed by Saturnino Martinez and Miguel Estribi), ‘Pentatomidae’
(including Scutelleridae and Cydnidae) (analysed by Dodge Engelman,
Panama), and Dermaptera (analysed by A. Brindle, Manchester Museum).
There are a multitude of taxonomic riddles and in many cases the ‘species’
possibly should be called ‘morpho-species’, although with the help of specialists
the number of such cases is kept at a minimum.
Not all groups were sorted from all light-trap samples, as is documented-
in Table 1. Only the data on the Homoptera are complete. A unit’sample is
a full year of light-trap samples from a given site. Some sites have, for a number
of groups, several such year samples, and only in a few cases do the available
data cover less than one year (Corriente Grande, 4 months; the first year in
Fortuna, 9 months; Curculionidae from Guadalupe Arriba, 6 months).
In the discussion of diversity in Homoptera a few samples have been added
which were obtained by different methods, such as canopy fogging in the forest
at Pipeline Road (Wolda, 1979), sweepnetting in the forest understory on and
near BCI (Wolda & Wong, unpublished) and in young tree plantations near
lake Alajuela (Arauz, Jules & Quintero, 1980), light-trapping with mosquito
traps in the forest on the Majt island, and trapping with a different kind of
light-trap in the forest at Punta Escocts (Sutton, 1983).
316 H. WOLDA

Diversity
Species richness is the number of species in a sample, but as this statistically
depends on the number of individuals in the sample, it is sometimes advisable
also to use diversity indices. All diversity indices have their problems and
limitations, but the alpha of the log series (Fisher, Corbet & Williams, 1943)
seems to serve better than others (Taylor, Kempton & Woiwood, 1976; Wolda,
1983; 1984).

RESULTS

The number of species in all samples is plotted against the number of

individuals for Homoptera (A) and Orthoptera (B) in Fig. 1. The expected
relationship between species and individuals, at a given value for the diversity

In I I I I I I I

I I I
0 1000 2000 3000
No. of individuals
Figure I . The relation between the number of species and the number of individuals of Homoptera
Auchenorrhyncha (A) and ‘Orthoptera’ (Tettigoniidae, Mantidae and some Gryllidae) (B) in a series
of samples taken in Panama. Samples taken in different years in the same site are surrounded by a
line. Light-trap samples, each usually covering a full year of daily samples: 0 , from forested; 0 ,
from disturbed areas. Circles with a vertical dash are sweep-net samples from Alajucla (AL) and
Barro Colorado Island (BCI). A circle with a horizontal line refers to a sample obtained by canopy
fogging at the Pipeline Road (PK). LC = Las Cumbres, M I R = Miramar, MA = Majt,
CG = Corriente Grande, PE = Punta Esrocks. F O R T = Fortuna, BOQ= Boquete and
GA = Guadalupe Arriba. Lines within the Las Cumbres Homoptera and the BCI Orthoptcra areas
are the expected relations between sample size and number of species based on a constant diversity
index alpha.
 ALTITUDE AND INSECT DIVERSITY 317

index alpha, is indicated for two localities with an extra large between-sample
variation in sample size. For Homoptera in Las Cumbres, the mean value for
alpha is 65.3 and the line for expected relation between sample size and number
of species is drawn within the Las Cumbres plot, showing that all the samples
conform reasonably well to the log series model in this respect. For the
Orthoptera from BCI, the first 3 years (1974-1976) have many fewer species
than all later years (the lower left-hand points in the BCI plot). Mean alpha for
all other points, 1977 to 1984, is 17.5. Figure 1B shows that the 1979-1984
points are very close to the line based on this value of alpha, while the richest
two years, 1977 and 1978, the two points on the right, have more species than
expected. A linear regression through all points is significantly steeper than that
through just the 1979-1984 points (P<O.Ol). This suggests that, over all 11
years, species richness increases more with sample size than the hypothesis of a
constant alpha would predict.
There were two traps operating simultaneously on BCI and only one at all
other sites, which may have affected the number of species caught. Figure 2
gives the information for each of the two BCI traps separately for Homoptera
over 5 years. The traps are not truly independent as each of the traps might

20 000

-
u1
0
a
u
.-
?
u
I

10 000

500

250

I00

75
t+ +-
$ /+\+

50
+---+
1980 1981 1982 1983 1984
Years

Figure 2. The number of individuals, the number olspecics and the diversity indrx alpha in 5 years
of light-trap wmples of Hornoptera on BCl at a low level (m), in the canopy (A)and both traps
combined (01. 'l'hr standard errors of the talues for alpha are given by vertical lines.
318 H. WOLDA

have caught more had the other not been there, but the data serve as a first
approximation. In all years except 1983, the canopy trap caught many more
individuals than the lower trap, but in all 5 years the lower trap caught more
species. The two traps combined collected more species than either one
separately, but in relation to the number of individuals caught, as expressed by
alpha, the lower trap sample was as diverse as both traps combined. The canopy
trap was less diverse.
Figures 3 and 4 show the correlation of sample size and species richness with
altitude for two groups, cockroaches and scarab beetles, showing a large
variation between years within sites, but an undeniable gradual decrease in
species richness and sample size from low to higher elevations. Figure 1 shows a
similar decrease for Homoptera and Orthoptera from BCI and Las Cumbres via
Fortuna and Boquete to Guadalupe Arriba. For all groups similar plots were
made and regression lines were calculated. Invariably those lines, for
individuals, species and alpha, showed a decrease, usually a significant one. Also
invariably, samples taken at Guadalupe Arriba were small and species-poor.
One extreme case is given by the Dermaptera where the three data points for
individuals, species and alpha in Fortuna (1010 m) are not lower than the two
points for BCI (125 m). At Boquete (1350 m), however, samples are much
smaller and poorer, and at Guadalupe Arriba (2200 m) the sample had only a
few individuals. Fewer individuals and fewer species of chrysomelid beetles were
obtained at 1350 and 101Om than on BCI,-but the value of alpha did not

0 0
8
-
I
P

I I
90 -

lo

Altitude (rn)

Figure 3. The relationship between altitude and sample size (the number of individuals), the
number of species and the diversity index alpha of cockroaches (Blattaria).
 ALTITUDE AND INSECT DIVERSITY 319

20 000 0
-
O 0

-$ loo00

0
I 1 I
0
I50

s! 100
ii
cn
50

0 I I I I 0
30
0

0
Jz
20
-
n
a
10'

I I 1 l 8
0 500 1000 1500 2000
Altitude (m)

Figure 4. The relationship between altitude and sample size (the number of individuals), the
number of species and the diversity index alpha for scarab beetles (Scarabaeidae).

change over that range. At the other extreme are the anthicid beetles which
were not found at all in any of the highland samples.
The data for all groups are summarized in Table 1. In order to save space, in
each locality the individual, yearly samples are ignored and only the mean
number of individuals, the mean number of species and mean alpha is given.
Some families of beetles were combined for the same reason (Anthicidae and
Pselaphidae in one group, and Cerambycidae, Chrysomelidae, Scarabaeidae,
Elateridae and Eucnemidae in another). It should be kept in mind that the
sample from Corriente Grande (100 m) covers only 4 months so that both the
number of individuals and the number of species are expected to be lower than
they would be for a full year. The sites with more extensive data, covering the
range from 125-2200 m, are most indicative of any changes with altitude.

DISCUSSION

Altitude e$ect
In all insect groups tested, the present data show a decrease in species richness
with increasing altitude. With the exception of the earwigs and, possibly, the
chrysomelid beetles, where the decrease only started after 1000 m, the decrease
was gradual from the lowlands to 2200 m. There is no sign of an intermediate
level maximum such as is indicated in the literature. The collecting technique as
well as the habitats sampled may have something to do with this. The seasonal
320 H. WOLDA

Table 1. For each site (altitude in metres given in parenthesis): mean number of
individuals (N)and species (S), mean diversity index alpha ( a ) per year, and
number of years; -, No data

Corriente Barro Las

Miramar Grande Colorado Cumbres Fortuna Boquete Guadalupe
(0) (100) (125) (150) (1010) ( 1350) (2200)

Homoptera
"hf 7546 2547 22046 17734 9828 2859 5445
S 376 401 458 355 239 152 66
a 83.5 133.8 82.1 65.3 44.6 34.9 10.4
Years 1 1/3 11 6 3 3 2
Blattaria
"hf 182 370 2202 350 622 62 9
S 30 44 53 34 28 13 2
a 10.2 13.0 10.0 9.2 6.3 5.5 0.7
Years 1 1 /3 10 3 3 3 2
Orthoptera
N 99 32 1566 219 77 52 1
S 33 19 73 43 16 15 1
a 17.3 19.6 15.9 16.0 6.4 7.9 0.5
Years I 1/3 11 1 3 3 2
Pselaph. Anth.
N 8258 980 6452 5460 1773 I292 365
S 89 103 114 69 62 27 -
a 13.9 29.0 19.7 11.1 12.5 4.8 -
Years I 1/3 3 3 3 I 2
Curculionidae
N 3903 2867 2852 1 3656 2124 1122 274
S 151 236 703 360 218 173 25
a 31.2 61.0 131.0 99.0 65. I 57.7 10.8
Years 1 I 13 3 1 3 2 112
Var. Coleopt.
N 26676 2968 34705 - 8518 15086 1897
S 215 272 597 - 283 24 1 51
a 32.0 72.9 102.5 - 56.8 41.0 9.5
Years 1 I /3 3 0 3 2 2
Sphingidae
N 149 I05 700 262 389 - 10
S 32 18 44 36 23 - -
a 12.5 6.3 11.3 11.3 5.9 - -
Years I 1/3 6 1 3 0 2
Pentatom. etc
N 363 286 934 349 517 187 -
S 50 19 46 41 16 18 ~

a 15.7 4.6 10.5 12.9 3.4 5.2 -

Years I 1/3 5 1 3 3 0
Dermaptera
x 171 438 683 ~

971 45 12
S I1 11 16 - 14 5 -
a 2.6 2.0 3.0 - 2.5 1.5 -
Years 1 I /3 2 0 3 2 2

changes observed depend on the technique used, sweep-netting and light-

trapping producing different results (Wolda & Wong, in prep.). The data from
Costa Rica (Janzen, 1973a,b) are confounded by sweep samples having been
taken in different seasons. Morbn & Terr6n (1984) found only one extra species
of scarab beetle at 1120 m than at 650 m. For the light-trapped moths in Papua
New Guinea (Hebert, 1980), no data from low altitudes are presented, so it is
 ALTITUDE AND INSECT DIVERSITY 32 1

not clear at what altitude the decrease of species abundance started. Turner &
Broadhead (1974) find a steady increase in species of Psocoptera on mango trees
in Jamaica between 152 and 1220 m, but the authors explain this in relation to
increased human disturbance at lower altitudes. The results obtained by Gagne
(1979) in Hawaii may also be related to changes in human disturbance with
a1titude.
Some light-traps operated all night while others were functioning only
between 6 and 10p.m. The latter include the highland traps in Fortuna and
Guadalupe, but not Boquete, and it also includes the lowland trap of Corriente
Grande. This factor does not seem to play a major role (Fig. 1, Table 1).
The study by Janzen et al. (1976) clearly shows an intermediate altitude
maximum in species abundance. Whether the difference with the present results
is due to them sweepnetting in secondary habitats (“old fields”) in Venezuela,
while my studies were done with light-traps in undisturbed forests or in a
disturbed habitat with a high tree species diversity (Las Cumbres, Boquete), is
not known, but effects of technique and habitat seem likely. The most important
cause of the differences between the present data and many of those in the
literature probably is the continous operation of the light-traps over a long time
in contrast with the only short periods covered by most other studies.

Human disturbance efect

T o see whether there is a n effect of habitat discernible in the present data,
pairs of samples were compared. Las Cumbres and BCI are at similar altitudes
(150 and 125 m, respectively), at a distance of 35 km from each other. BCI is on
the Atlantic side of the Isthmus, while Las Cumbres in on the Pacific slope. BCI
is a relatively undisturbed forest while Las Cumbres is a disturbed residential
area. In all groups, a yearly sample from Las Cumbres had fewer individuals
and fewer species than those from BCI, and in most groups, though not all,
alpha was also lower (Table 1). Figure 1 illustrates this in more detail for
Homoptera. It is conceivable that the fact that two traps were operating on BCI
against only one in Las Cumbres contributes to this difference. I t was shown
that the canopy trap on BCI caught more individuals but fewer species than the
lower trap (Fig. 2). During the first period in Fortuna, when the trap was in the
canopy, fewer species were caught than in the subsequent two years with the
trap at a low level, supporting the difference between canopy and low level
found on BCI. The Las Cumbres trap is on top of a slope, overlooking a wooded
valley, and may be comparable to a canopy trap, so that the difference between
Las Cumbres and BCI may be largely explained by the number of traps.
Boquete (1350 m) is only 20 km from Fortuna (1010 m), and is much more
disturbed than Fortuna. In almost all groups the samples from Boquete have
fewer species and fewer individuals than those from Fortuna. This may partly be
an effect of altitude but an effect of human disturbance seems also likely, but, if
present, it is small. Miramar (sea level), 65 km from Corriente Grande
(100 m), is a very disturbed habitat with some trees along pastures, although
these is a (disturbed) forested slope nearby. Corriente Grande is good forest. I n
most groups the samples from Corriente Grande, which cover only 4 months,
have more species than those from a full year in Miramar.
These three comparisons show that a moderate disturbance, one which leaves
322 H. WOLDA

a relatively high plant (tree) species diversity, does have a surprisingly small
effect on species richness. Only if the disturbance is extreme, such as in
Miramar, is a clear effect noticeable. The kind of species present change
(Wolda, 1983, 1984), but not so much their total number.

ACKNOWLEDGEMENTS

I gratefully acknowledge the help of all those who participated in the

collecting and/or sorting of the insect samples, such as A. Brindle, Carlos
Castillo, Donchandler, Dodge Engelman, Cecilio Estribi, Miguel Estribi, Frank
W. Fisk, Bonifacio de L e h , Saturnino Martinez, Charles W. O’Brien and
Nessim Yishui. Some of the collecting was partly supported by the
Environmental Sciences Program and the Scholarly Studies Program of the
Smithsonian Institution and by the Instituto de Recursos Hidriulicos e
Electrificacih (IRHE).

REFERENCES

ADAMES, A. J., 1977. Evaluacion ambiental y efectos del proyecto hidroelectrico Fortuna. Loteria, 254, 2.55,
256: 1-538.
ARAfJZ, A., JULES, G. & QUINTERO, V., 1980. Estudio ecoldgico de la entomofauna de la capa baja en parceles
experimentales en el Lago Alajuela. Lic. thesis, University of Panama.
CLARIDGE, M. F. & SINGHRAO, J. S., 1978. Diversity and altitudinal distribution of grasshoppers
(Acridoidea) on a Mediterranean mountain. Journal of Biogeography, 5: 239-250.
FISHER, R. A., CORBET, A. S. & WILLIAMS, C. B., 1943. The relation between the number of species
and the number of individuals in a random sample of an animal population. Journal of Animal Ecology, 12:
42-58.
GAGNE, W. C., 1979. Canopy associated arthropods in Acacia koa and Metrosideros tree communities along an
altitudinal transect on Hawaii island. PaciJic Insects, 21: 56-82.
GAULD, I. D., 1985. A preliminary survey of the Ophioninae (Hymenoptera: Ichneumonidae) of Brunei.
Brunei Museum Journal, 6: 169-185.
GAULD, 1. D., 1987. The so-called “anomalous diversity” of tropical Ichneurnonidae re-examined. Biological
Journal of the Linnean Society, 24: 299-312.
CONTHART, H., 1984. Zikaden (Horn. Auchenorrhyncha) aus der alpinen Hohenstufe der Schweizer
Zentralalpen. Mitteilungen dcr Schweizerischen Entomologischen Gcsellschaft, 57: 129-1 30.
HERBERT, P. D. N., 1980. Moth communities in montane Papua New Guinea. Journal of Animal Ecology, 49:
593-602.
JANZEN, D. H., 1973a. Sweep samples of tropical foliage insects: Description of study sites, with data on
species abundances and size distributions. Ecology, 54: 659-686.
JANZEN, D. H., 1973b. Sweep samples of tropical foliage insects: Effects of seasons, vegetation types,
elevation, time of day, and insularity. Ecology, 54: 687-708.
JANZEN, D. H., 1976. Why are there so many species of insects? Proceedings XV Congress of Entomology,
Washington: 84-94.
JANZEN, D. H., ATAROFF, M., FARINAS, M., REYES, S., RINC6N, N., SOLER, A., SORIANO, P. &
VERA, M., 1976. Changes in the arthropod community along an elevational transect in the Venezuelan
Andes. Biotropica, 8: 193-203.
LEIGH, E. G., RAND, A. S. & WINDSOR, D. M. (Eds), 1982. ‘The Ecology of a Tropical Forest. Seasonal
Rhythm and Long-term Changes. Washington, D.C.: Smithsonian Institution Press.
McELRAVY, E. P., WOLDA, H. & RESH, V. H., 1982. Seasonality and annual variability of caddisfly
adults (Trichoptera) in a “non-seasonal” tropical environment. Archiv fur Hydrobiologie, 94: 302-3 17.
MICHENER, C. D., 1979. Biogeography of the bees. Annals Missouri Botanical Garden, 66: 277-347.
MORdN, M. A. & TERRON, R. A., 1984. Distribuci6n altitudinal y estacional de 10s insectos necrofilos en
la Sierra Norte de Hidalgo, Mexico. Acta oofdgica Mexicana, 3: 1-47.
MOUND, L. A. & WALOFF, N., 1978. Diversity of Insect Faunas. Oxford: Blackwell Scientific Publications.
PRANCE, G. T., 1982. Biological DiversiJication in the Tropics, New York: Columbia University Press.
SMYTHE, N., 1982. The seasonal abundance of night-flying insects in a neotropical forest. In E. C. Leigh,
A. S. Rand D. M. Windsor (Eds), The Ecology of a Tropical Forest. Seasonal Rhythm and Long-term Changes:
309-318. Washington, D.C.: Smithsonian Institution Press.
STILES, F. G., 1983. Birds. In D. H. Janzen (Ed.) Costa Rican flatural History: 502-530. Chicago: University
Chicago Press.
 ALTITUDE AND INSECT DIVERSITY 323

S U I T O N , S. L., 1983. The spatial distribution of flying insects in tropical rain forests. In S. L. Sutton, T. C.
Whitmore & A. C. Chadwirk (Eds), Tropical Rain Foresl: Ecology and Management: 77-91. O x h r d : Blackwell
Scientific Publications.
TAYLOR, L. R., KEMPI‘ON, R. A. & WOIWOD, I. P., 1976. Diversity statistics and the log series model.
Journal oJ Animal Ecology, 45: 255-272.
TERBORGH, J., 1977. Bird species diversity on an Andean elevational gradient. Ecology, 58: 1007-1019.
‘TURNER, B. D. & BROADHEAD, E., 1974. The diversity and distribution of psocid populations on
MangiJera indica L. in Jamaica and their relationship to altitude and micro-epiphytic diversity. journal of
Animal Ecology, 43: 173-190.
WOLDA, H., 1979. Abundance and diversity of Homoptera in the canopy of a tropical forest. Ecological
Enlomology, 4: 181-190.
WOLDA, H., 1980. Seasonality of tropical insects. I. Leafhoppers (Homoptera) in Las Cumbres, Panama.
Journal of Animal Ecology, 49: 277-290.
WOLDA, H., 1983. Diversity, diversity indices and tropical cockroaches. Oecologia (Berlin), 58: 290-298.
WOLDA, H., 1984. Diversidad de la entomofauna y c6mo medirla. Informe Final IX Congreso Latino American0
de <oologia, Peru: 181-186.
WOLDA, H. & FLOWERS, R. W., 1985. Seasonality and diversity of mayfly adults (Ephemeroptera) in a
“non-scasonal” tropical environment. Biolropica, 17: 330-335.