Contributed Pqers
The Magnitude of Global Insect Species Richness
KEVIN J. GASTON
Biodiversity Division
Department of Entomology
The Natural History Museum
Cromwell Road
London SW7 SBD, LJ.K
Abstract: Recent suggestions that insects number tens of
millions of species have received much attention. Little con-
sideration, hoWelJW, has been given to how such estimates
compare with what else we know about insect species rich-
ness. Perhaps most significantly, the specialist knowledge of
the taxonomic community at large has generalIy been ig-
nored Collation of published and unpublished information
from this source provides little to encourage belief in truly
vast numbers of undescribed species of insects. For the insect
groups f o r which figures are available, estimates of global
total numbers of species are typically less than ten times the
numbers of described species. Although minimum global es-
timates are more readily constructed than upper estimates,
these figures uniformly fail to support assertions that there
are 30 million or more species of insects. Rathw, a,figure of
less than ten million seems more tenable, and one of around
five million feasible.
Additional, more circumstantial, evidence tends to sup-
port the idea that insect species numbers may not be as vast
as has been claimed First, the contribution of canopy spe-
cialists to global richness m a y be less than often suggested.
Second, a higher proportion of species than commonly en-
tertained may have moderate to large geographic ranges.
Third, the number of groups failing to increase in richness in
the tropics may have been underestimated. Finally, the pro-
portion of undescribed species encountered by many taxon-
omists seems insufficient to justcyy estimates of vast species
numbers.
Introduction
The question of how many insect species there are has
stimulated interest for more than a century (Westwood
Puper submitted Junualy 17, 1991; revised manuscript accepted
April 16, 1991.
Resumen: L a s recientes sugerencias de que las especies de
insectos llegan a [as decenas de millones, ban recibido
mucha atencidn. Sin embargo, se le ha dado poca consid-
eracion a como es que estas estimaciones se comparan con
todo lo que se conoce acerca de la riqueza en especies de
insectos. E s mus s i g n i f i c a t i v o el hecho d e q u e el
conocimiento del especialista de la comunidad taxondmica
ha sido en gran parte ignorado. El cotejo de la informacidn
taxonomica publicada y no publicada, ofrece poca eviden-
cia para estimular la creencia de que verdaderamente exista
un vast0 numero de especies de insectos por describir. Para
los grupos de insectos de 10s que se tiene informacih cuan-
tificada, el numero total de especies estimadas a nivel mun-
dial es tipicamente diez veces menor que el numero de es-
pecies y a descritas. Aunque las estimaciones minimas
globales son mas fucilmente caiculadas que las estima-
ciones m h i m a s , estos datos uncformemente fallan para
apoyar Ias aseveracioanes de que b u y mas de treinta mil-
lones de especies de insectos. M a s que esto, un dato de menos
de diez millones parece SCT mas sostenible y, uno de cerca de
cinco millones, enteramente posible.
Evidencia adicional, mas circunstancial tiende a apqyar
la ideu de 4ue el numero de especies de insectos puede no ser
tan vast0 como se ha denunciado. Primero, la contribucidn
de 10s especialistas del dose1 a la riqueza global ha sido
sugerida con poca frecuencia Segundo, una proporcidn m&
alta de especies de las que comunmente se ban tratadopuede
que tengan distribuciones geograficas de rangos moderados
y grandes. Tercero, el numero de grupos que estu fallando
para incrementar la riqueza en 10s trripicospuede haber sido
subestimado. Finalmente, la proporcion de especies no de-
scritas enconhadas por muchos ta.xdnomos son insuficientes
para justijicar estirnaciones de vastos numeros de especies.
1833; Metcalf 1940; Sabrosky 1952; May 1986, 1988;
Stork 1988;Adis 1990). However, widespread comment
and debate on the subject is a recent phenomenon, en-
gendered by both a growing need to understand the
magnitude of the extinction crisis and suggestions that
there may be vastly more insects than previously
thought. The latter were principally fueled by T. L. Er-
283
Conservation Biology
Volume 5 , No. 3, Srptemkr 1991
284 Global Insect Species Richness Gaston

win, who proposed, on the basis of samples of beetles Table 1. Estimates of the numbers of species described in the
from the canopies of nineteen specimens of one species major insect orders.
of Panamanian tree, that there may be 30 million species Coleoptera 370,000 Ilammond 1974
of arthropods (principally insects) in tropical regions 350,000 Southwood 1978
alone (Erwin 1982). Erwin (1988) has subsequently 340,500 Lawrence 1982
suggested that this figure may be too small. 290,000 Arnett 1967, 1985
Diptera 120,000 Southwood 1978
The calculations upon which Erwin’s estimates are
98,500 Arnett 1985
based are undoubtedly simplistic, and problems with 118,700 Evenhuis 1989
many of the assumptions made have been amply dem- Lepidoptera 120,000 Southwood 1978
onstrated (Stork 1988; May 1988; Thomas 1990). What 165,000 Laithwaite et al. 1975
has received less consideration, with regard to both 1 12,000 Arnett 1985
142,500 Holloway et al. 1987
these and other recent estimates of species numbers, is Hymenoptera 100,000 Southwood I978
how they compare with what else w e know about insect 1.10,000 Brown 1982a
species richness. Perhaps most significantly, the special- 103,000 Arnett 1985
ist knowledge of the taxonomic community at large,
much of it written but much of it not, has generally been
ignored. This is somewhat unfortunate, as it is through (Parker treats the Hemiptera [ = Heteroptera] and Ho-
the hands of this same community that the bulk of spec- moptera as two separate orders). Estimates of the num-
imens collected in the centers of faunal diversity has ber of described species in each of these groups have
passed and continues to pass. been made on a number of occasions (Table 1 ). Given
In this paper I draw together a large amount of infor- the difficulty in compiling such tallies the results for
mation about species richness, culled largely from the each order tend to demonstrate a reasonable level of
taxonomic community, to try and determine to what similarity, although Arnett’s (1985) figures tend to be
extent these data support estimates of tens of millions of lower, in some cases drastically, than those in other
insect species. Much of what follows is unashamedly studies. Why Arnett’s figures should be so different is
speculative; unless w e can ultimately count the earth’s unclear.
insects, all attempts to determine how many there are Markedly more Coleoptera have been described than
(including those already published) will remain open to species in any of the other orders. It is not possible to
this accusation. state categorically for these remaining orders which has
It is plain that one fundamental difficulty with global most described species. However, there are probably
species richness estimates is the lack of any agreed and more Lepidoptera described than Diptera o r Hy-
uniformly applied definition of what a species is. This menoptera, and the last two probably have roughly
complexity has been discussed elsewhere (Adis 1990; equal numbers of described species. On an annual basis,
May 1990) and will not be pursued here. Rather, I ac- increases in the numbers of species described and syn-
cept species concepts as currently used by workers on onymized in each of the four major orders may be
different groups, as well as estimates based on current highly variable (Table 2). The publication of only a few
perceptions as to how these concepts may change in the major works can have a profound effect on the figures
future. for any one year. Nonetheless, it is noteworthy that for
the 2 years for which data have been collated, a total of
about 5500 species has been described for the four or-
Described Species ders, and around a third of this number synonymized.
Indeed, the levels of synonymy are far higher than com-
Before considering how many insect species we don’t monly entertained.
yet know, w e need to be clear about how many have The proportion of species richness accounted for by
already been described. Estimates of the total to date are the Coleoptera, Diptera, Lepidoptera, and Hymenoptera
varied. Borror et al. (1981) and Arnett (1985) both give
figures of about 750,000, while Wolf (1987) gives one Table 2. Approximate numbers of species of Coleoptera,
Diptera, Lepidoptera, and Hymenoptera recorded as newly
of 874,161 including other arthropods. The 1976 cen- described or as newly synonymized in Zoological
sus of collections in the Department of Entomology of Record for two years.
the Natural History Museum, London (unpublished),
roughly estimated 827,000 known insect species of 1986-8 7 1988-89
which representatives of 445,000 were housed in the Descr.
~
Svnon. Descr. Svnon.
department. Coleoptera 2300 574 2134 580
Of the 2 8 insect orders recognized by Parker ( 1982), Diptera 1044 227 1126 202
only four contribute substantially to the total, the C o - Lepidoptera 943 274 785 353
Hymenoptera 1081 352 1349 788
leoptera, Diptera, Lepidoptera, and Hymenoptera

Conservation Biology
Volume 5, N o 3, September 1991
Gaston Global Insect Species Richnm 285

will doubtless be at least as great for the sum of unde- Table 3. Estimates of numbers of described species for major
scribed and described species as it is for the latter alone. components of the Coleoptera, Diptera, Lepidoptera,
It is thus principally to these four orders we must look and Hymenoptera.
for an understanding of the magnitude of global insect Described
species richness. Sbb. Reference
Coleoptera:
Coleoptera Carabidae 30,000 Lawrence 1982
The Coleoptera currently comprise more than 150 fam- Staphylinidae 30,000 ...
Scarabaeidae 25,000 ...
ilies ( 168 in Crowson 1981; 153 in Lawrence 1982). In
Buprestidae 15,000
common with other taxa where similar patterns have Tenebrionidae 18,000
been sought (e.g., Anderson 1974; Dial & Marzluff Cerambycidae 35,000
1989), the distribution of described species among Chrysomelidae 35,000 ...
these families is highly skewed with the majority con- Curculionidae 50,000 ...
taining relatively few species. Indeed, just eight families Diptera:
Tipulidae 14,000 Alexander & Byers
contain about two-thirds of described beetles (Table 3). 1981
The bulk of undescribed species belong to the same 13,000 + Bickel 1982
eight families. However, their true relative sizes are un- Asilidae 5,000 Wood 1981
doubtedly very different from those in Table 3 (see be- Bombyliidae 4,000 + Bickel 1982
low). Dolichopodidae 6,000 Robinson & Vockeroth
1981
A number of attempts have been made to estimate the Syrphidae 6,000 Vockeroth &
global or regional size of beetle families using rates of Thompson 1987
species description. However, these have largely been Tachinidae 6,000 + Bickel 1982
unsuccessful because the curve of cumulative numbers 8,000 Wood 1987
of described species through time on which they have Lepidoptera:
Pyralidae 20,000 Holloway et al. 1987
been based is insufficiently defined for these purposes Noctuidae 2 1,000 Holloway et al. 1987
until it begins to asymptote (see debate between White Arctiidae 11,000 Watson & Goodger
1975, 1979; Frank & Curtis 1979; O’Brien & Wibmer 1986
1979). Holloway et al. 1987
The Staphylinidae (as defined by many modern work- Geometridae 20,000 Holloway et al. 1987
Hymenoptera:
ers to include Dasyceridae, Scaphidiidae, Pselaphidae, Apocrita 125,000 Brown 1982a
Empelus, and Silphidae [in major part]; more extensive Ichneumonoidea
than that of Lawrence (1982) and Table 3) form what is Braconidae 10,000 Gauld & Bolton 1988
almost certainly the largest beetle family, perhaps num- Ichneumonidae 14,816 Townes 1969
bering around 300,000 species (P. M. Hammond, per- Chalcidoidea 18,601 Noyes 1990
Vespoidea
sonal communication). The largest subfamily of sta- Formicidae 8,800 B. Bolton, pers. comm.
phylinids, the Aleocharinae, may alone comprise some E. 0. Wilson, pers.
100,000 species (Hammond 1975). The figure of comm.
300,000 represents an eight- to tenfold increase over Apoidea
the number of described species in the family, vastly Apidae 20,000 Michener 1974
Roubik 1989
greater than suggested increases for any other major
beetle families. Estimated increases for the pselaphids
alone are considerably smaller, with suggestions of two superfamily;however, neither comes close to the levels
to three (D. S. Chandler, C. Besuchet, pers. comm.), and of increase predicted for the Staphylinidae.Suggestions
four to five (P. M. Hammond, pers. comm.) times the of approximate doublings of described species numbers
8400 described species. have also been made for two further major beetle fam-
The Curculionidae, currently the insect family with ilies, the Carabidae (est. 60,000 spp., N. E. Stork, pers.
most described species, have been estimated as having a comm.) and the Chrysomelidae (est. 60,000-70,000
total of 85,000(O’Brien & Wibmer 1979) species (note: spp., Jolivet 1988). It is perhaps not unreasonable to
Curculionidae sensu O’Brien & Wibmer is not that of believe that some other of the better-worked large bee-
Lawrence [ 19821and Table 3), while the superfamily to tle families will show increases of a similar magnitude.
which they belong, the Curculionoidea ( 56,900 de- These include the Scarabaeidae, Buprestidae, Ceramby-
scribed spp.), has been estimated to number some cidae, and Elateridae, which although not included in
180,000 species (G. Kuschel, pers. comm.). These are Table 3, contains some 9000 described species
increases of between two to four times the number of (Lawrence 1982).
described species. The two estimates are difficult to rec- Several families not so far mentioned may be much
oncile as the Curculionidae comprise the majority of the larger than numbers of described species would indi-

ConservationBiology
Volume 5, No. 3, Scptmbcr 1991
286 Global Insect Species Richness
cate. Recent work in tropical forests suggests that
groups such as the mordellids ( 1200 described species,
Lawrence 1982) may be more diverse than previously
suspected (Hammond 1990). However, none of these
taxa is likely to alter radically any impressions of the
overall size of the order gained from looking at those
families with most described species.
Taking an overall ratio of described to undescribed
species, somewhere between that suggested for sta-
phylinids and those for other major families seems a
reasonable approach to estimating the size of the order.
Assuming 350,000 described beetle species, figures
range from about a million ( 3 X described spp.) to
about three million (9 X described spp.). Such figures
are of a similar magnitude to the 2 to 5 million estimate
of Hammond (1975).
Diptera
The Diptera are the best cataloged of the major insect
orders, with volumes available for each of the major
biogeographic regions (incl. Delfinado & Hardy 1973,
1975, 1977; Crosskey 1980a; McAlpine 1981, 1987;
Soos 1 9 8 4 4 ; Evenhuis 1989). Despite this, there has
been very little consideration of the probable size of
components of the order. In comparison with families of
the other three major insect orders, even the largest of
the 123 or so dipteran families (Bickel 1982) do not
comprise particularly large numbers of described spe-
cies (Table 3 ) . Some families, such as those containing
biting flies (e.g., Culicidae, Tabanidae) and those with
many large or obvious species (e.g., Asilidae, bombyli-
idae, Syrphidde ), are markedly better known than oth-
ers. In none of the reasonably well known groups is the
species total likely to be vast. The Tipulidde comprise
the largest family, about 10 percent of described flies. A
substantial number remain to be named, with Alexander
and Alexander (1970) suggesting that not more than
half of the Neotropical species have been described, but
the group is among the better known.
In terms of true species numbers some of the families
with relatively few described species are among the
largest. The Tachinidae, Cecidomyiidae ( 3000-4000 de-
scr. spp., Harris 1980; Gagne 1981), Phoridae (3000
+
descr. spp., R. H. L. Disney, pers. comm.; 2500 , Peter-
son 1987) and Mycetophilidae (3000 described spp.,
Vockeroth 1981 ) are particularly diverse. Crosskey
(1980b) states that “there is little doubt” that Tachi-
nidde is the largest family of living Diptera, though he
gives no indication as to how large. If it parallels some
other parasitoid families (e.g., Ichneumonidae, Bra-
conidae) it may number 50,000 species or more. Gagne
(cited in Thompson 1990) estimates that there may be
some 15,000 species of cecidomyiid in the Nearctic re-
gion alone, some 35 percent of the Diptera estimated to
occur there (Thompson 1990). Thompson (1990)
Conservation Biology
Volume 5 , No. 3, September 1991
Gaston
seems to believe that this is too high, but whatever the
precise figure, cecidomyiids comprise one of the largest
families of Diptera in North America. If their patterns of
richness are repeated in the tropics, as some believe,
then they must rate among the largest of insect families.
The Phoridae are another group which though clearly
very diverse in the tropics remain extremely poorly
known. Frfty thousand may be a reasonable guess at
their global richness (R. H. L. Disney, pers. comm.), but
at this stage it can be little more than a guess.
Extracting some overall picture of the size of the or-
der from these scattered remarks is obviously virtually
impossible. Colless and McAlpine ( 1970) regard “the
world total of species, described and undescribed spe-
cies, [as] probably being at least 150,000,”while Hardy
(1977, cited in McAlpine 1979) estimates a global total
of 200,000 species of flies. If groups such as the phorids
and cecidomyiids are as diverse globally as detailed
study in temperate regions and limited observations
from tropical ones suggest, then such a doubling of de-
scribed species is very conservative. However, even
doubling Hardy’s estimate fails to generate a figure of
half a million species of flies.
Lepidoptera
The Lepidoptera are generally regarded as the best col-
lected and studied of the four major insect orders. How-
ever, statements regarding the probable size of the or-
der or of its major components do not seem to have
been made. It comprises about 120 families ( 1 37 in
Munroe 1982; 113 in Holloway et al. 1987), of which
just four each contain more than 10,000 described spe-
cies (Holloway et al. 1987). Using Holloway et al.3
(1987) estimates these families total about a third of
described species of Lepidoptera (Table 3). The Noctu-
idae, Arctiidae, and Geometridae are all “macro-
lepidopteran” (superfamilies Bombycoidea + Geo-
metroidea + Noctuoidea + Papilionoidea) families,
and account for the bulk (>60%) of the 80,000 species
(Watson & Goodger 1986) estimated to have been de-
scribed in this group of Lepidoptera. Both these and the
other macrolepidopteran families have been reasonably
well collected in the tropics and it is doubtful that fur-
ther study will more than double their sum total. Rates
of synonymy are reasonably high in many revisions of
these taxa in tropical regions (e.g., Holloway 1983),
markedly reducing any net gain in species numbers pro-
duced through more intensive collecting efforts.
The high frequency of synonyms that remain to be
identified presents a complication to estimating the
probable size of some lepidopteran groups. A pattern
already experienced with some groups, and likely to be
repeated in others, is that the number of recognized
species peaks and then actually declines with time (e.g.,
Danainae: Ackery & Vane-Wright 1984), as rates of syn-
onymy overtake rates of description.
Gaston
As with all the major insect orders, in general, the
lepidopteran g r o u p s w i t h small body size ( t h e
“microlepidoptera”) are more poorly known than those
that are larger. Here, collecting in tropical regions still
yields a high proportion of undescribed species, partic-
ularly among pyraloids and gelechioids.
Discussion with several lepidopterists suggests that an
upper bound to the overall size of the order of 500,000
species is reasonable, and that a figure somewhat lower
is quite probable. Based it seems upon projections of 1
to 10 million insect species in total, and the same ratio
of all Lepidoptera species to all insect species as is true
of described species, Hodges (1976) states that there
may be as many as 280,000 to 1,400,000 Lepidoptera
species.
Hymenoptera
More serious consideration of total numbers of species
has been given to Hymenoptera than to any of the other
three major insect orders. Here, the vast majority of
species lie in the Apocrita, with an estimated 125,000
described species (Brown 1982a; a figure in excess of
some estimates of the size of the order as a whole; see
Table 1). Two superfamilies account for a substantial
proportion of apocritans, and probably for most of total
hymenopteran species richness. These, the Ichneu-
monoidea (Ichneumonidae + Braconidae ) and the Chal-
cidoidea (21 families; Gauld & Bolton 1988), have a
combined total of around 45,000 described species (Ta-
ble 3). Estimates for the probable total species richness
of both groups are available in the literature. Townes
(1969) offers a figure of 60,500 ichneumonids, while
van Achterberg (1984) suggests a figure of at least
40,000 braconids, and Mason (1979) at least 50,000.
These are increases by factors of four to five over num-
bers of described species and total around 100,000 ich-
neumonoids. Noyes ( 1978) estimates that the Chalci-
doidea also number around 100,000 species and sees no
reason drastically to alter this conclusion at present 0.
Noyes, pers. comm.). This also represents an increase of
five over described species numbers of the order.
These estimates provide a minimum figure for the
Apocrita, and hence for Hymenoptera as a whole, of
around 200,000 species. This agrees with the sugges-
tions of Brown ( 1982a) and Masner ( 1979) and is not
too dissimilar to Gauld and Bolton’s (1988) minimum
number of 250,000 Hymenoptera. However, to improve
upon this figure w e need to add to it estimates for the
other major superfamilies, namely the Cynipoidea, Proc-
totrupoidea, Vespoidea, and Apoidea. Gauld and Bolton
(1988) estimate there are some 20,000 species in the
Cynipoidea, an increase by a factor of six over described
species. Determining the size of the other groups is
more problematical, with little information available for
the groups in their entirety. Total numbers of ant spe-
Global Insect Species Richness 287
cies (Vespoidea: Formicidae) are estimated to be be-
tween about double (15,300 spp., B. Bolton, pers.
comm.) and quadruple (30,000 spp., E. 0.Wilson, pers.
comm.) the numbers of described species. Roubik
( 1989) tentatively suggests that there are twice as many
bee species (Apoidea: Apidae; defined as per Gauld &
Bolton 1988) as described, giving some 40,000 species,
while Gauld & Bolton (1988) give a figure of 25,000-
30,000,The combined total of species in the Vespoidea
and Apoidea is unlikely to reach 100,000 species, and
while the Proctotrupoidea may be relatively large it is
smaller than either the Ichneumonoidea or the Chalci-
doidea.
In sum, adding in the more minor superfamilies not
previously mentioned, the Hymenoptera could reason-
ably be argued to number around half a million species.
I. I). Gauld (pers. comm.) believes 600,000 to be a max-
imal figure.
The Relative Sizes of the Major Orders
In considering the possible richness of the four major
orders, evidence based on their relative sizes has been
purposely avoided. Although the question of which is
the more speciose order is a long-standing one, it has
seen little in the way of resolution. Interestingly, data on
the numbers of described species and estimates of the
true richness of Coleoptera, Diptera, Lepidoptera, and
Hymenoptera in various regions of the world suggest
different conclusions. In general, in each region there
are more described beetles than species of any of the
other orders. For Diptera, Lepidoptera, and Hy-
menoptera there is no consistent tendency for one to be
more diverse than the others (Table 4a). In contrast,
when estimates of described and undescribed species
are considered, Hymenoptera is clearly believed to be
the most diverse order in several temperate regions,
with Diptera also being more speciose than Coleoptera
(Table 4b). There are consistently fewer lepidopteran
species estimated than any of the other three major or-
ders, with them typically numbering about half the spe-
cies of beetle. The primary exceptions to these gener-
alizations are the Australian regions, which have long
been regarded as atypical in their patterns of species
richness.
There is no information upon which to base similar
comparisons of the relative richness of the major insect
orders for tropical regions. Storks (in press) limited
canopy fogging in Brunei found more Hymenoptera and
Diptera species than Coleoptera on several trees (Table
5). However, without more information on the relative
beta-diversity of the orders and how representative such
samples are of regional faunas these results are impos-
sible to interpret. The absence of good data makes ex-
trapolation of the results in Table 4 to the global situa-
Conservation Biology
Volume 5 , No. 3, September 1991
288 Global Insect Species Richness Caston

Table 4. Relative numbers of Coleoptera, Diptera, Lepidoptera, either the Diptera or Hymenoptera, resulting in differ-
and Hymenoptera species in the insect faunas of different ent interpretations of the relative sizes of the orders
regions, presented as proportions of the numbers of coleopteran dependent upon whether described or estimated spe-
species (i.e., Coleoptera = I), (a) for known species and ( b ) for
estimated total numbers of species. A small part of the variation cies numbers are used. Examination of the years of de-
between regions can be attributed to whether Strepsiptera are scription of species in the major orders in the British
treated as oart of the Coleootera or not. fauna demonstrate this from a historic perspective (Fig.
Dipi. Lep, Hym Refmence
1). Many beetle species were described early on, at a
time when relatively few species from the other orders
( a ) Known
N. America 0 83 0.48 0.74 Kosztarab & Schaefer had been recognized. Historically, the disproportionate
1990 interest in Coleoptera extends to the tropics also. Many
Canada 105 0.70 0.89 Danks 1979 of the great collectors of these regions from the last
Norway 0 94 0.64 0.94 Aagaard & Hagvar
1987 century, such as Bates and Wallace, collected large num-
Hungary 0 85 0.54 1.67 Mahunka 1986 bers of Coleoptera and Lepidoptera (mostly macrolepi-
Switzerland 0 37 0.64 0.43 Sauter 1974 doptera), and assembling collections of both these
Britain 150 0.63 1.60 Stubbs 1982
Israel 1 00 0.77 1.00 A Freidberg, pers groups was a frequent colonial pastime. In contrast, rel-
comm , Kugler 1990 atively few tropical specimens of Diptera and Hy-
Hawaii 0 81 0.69 0.47 Howarth, 1990 menoptera found their way to taxonomists until early
Japan 0 58 0.57 0.45 Chu 1989
Taiwan 0 64 0.77 0.51 Maa 1956 this century.
Australia 0 27 0.74 0.52 CSIRO 1991 If the estimates of the sizes of the major orders made
Tasmania 0 39 0.64 0.24 P B McQuillan, earlier are approximately valid then w e must conclude
pers comm
New Zealand 0 4? 0.34 0.09 Watt 1975 that, despite such differential collecting, strong latitu-
( b ) Estimated dinal gradients in the relative species richness of the
N. America 0 77 0.49 1.03 Taylor 1983 major insect taxa exist such that there are more beetle
11-16 0.53 1.37 Kosztarab & Schaefer
1990 species globally than any other order. If not, then the
Canada 159 0.74 1.83 Danks 1979 estimates of numbers of Coleoptera may be too large,
Switzerland 164 0.64 1.64 Sauter I974 those of Diptera and Hymenoptera considerably too
Britain 146 0.61 1.73 Taylor 1983
Australia 0 30 0.65 0.54 Taylor 1983 small, or, most probably, both. Even if the rank sequence
Tasmania 0 51 0.66 0.51 P. B. McQuillan,
pers comm.
7000

tion difficult. There may be strong latitudinal trends in

the relative diversity of these taxa of which we are (I)

largely unaware. Danks (198S), for example, shows that .-a,

u
in more northerly regions Diptera appear to become :SO00
proportionately better represented, while Coleoptera v)

Y-
become less so. Attempts to understand such patterns o
will be greatly confounded by the differential collect- 0
ability of, and interest in, the major orders. It is clear C

from Table 4 that Coleoptera are in many regions con- 3000

sidered to be proportionately far better described than .-
-
+
U

Table 5. Relative numbers of Coleoptera, Diptera, Lepidoptera,

and Hymenoptera species in canopy-fogging catches kom ten
zv
3

Bornean trees, presented as proportions of the numbers of 1000

coleopteran species (i.e. Coleoptera = 1). Data from
Stork (1991).
I I I 1 t
Tree Dip. Lep. Hym.
7870 1890 7970
Shorea johorensis 1.02 0.1 1 1.38
0.68 0.07 1.11 Year
... 0.87 0.06 0.83
... 0.86 0.09 1.11 Figure 1. Numbers of species of Coleoptera, Diptera,
Shorea macrophylla 0.98 0.13 1.64 Lepidoptera, and Hymenoptera on the British faunal
... 0.77 0.11 0.99
Pentaspadon motleyi 1.77 0.32 1.07
list that were described in different years, and are
... 1.23 0.20 1.58 currently recognized Years are for names in usage,
Castanopsis sp. 1.39 0.19 1.97 and thus include some junior primary homonyms.
Indet. 1.OO 0.15 0.79
Data from Kloet G Hincks (1972, 1976, 1977, 1978).

Conservation Biology
Volume 5, No. 3, September 1991
Gaston
of sizes of the orders is approximately correct, it seems
likely that the absolute numbers of species are more
similar than the estimates imply.
Other Large Orders
Following Parker ( 1 9 8 2 ) , t h e Hemiptera ( = Het-
eroptera) and Homoptera are regarded here as each
having ordinal status. However, if they were treated as a
single order, they would join the Coleoptera, Diptera,
Lepidoptera, and Hymenoptera in comprising a substan-
tial proportion of described insect species richness.
There are around 35,000 described Hemiptera in 74
families (Slater 1982; 33,000 described spp. in Hodkin-
son & Casson, 1991) and 45,000 described Homoptera
in 55 families (Kosztarab 1982; 48,500 described spp. in
Hodkinson & Casson, 1991), totaling about 10 percent
of described insects. J. A. Slater (pers. comm.) estimates,
conservatively, that the total number of species in the
Hemiptera is around three times the number of de-
scribed species. The largest family in the order, the Mir-
idae, numbers some 10,000 described species (Slater
1982) and is likely to account for much of the final total.
The Homoptera is also dominated by a single family, the
Cicadellidae. This currently numbers 15,000-20,000
described species (Kosztarab 1982, Hamilton 1984),
but Hamilton (1984, commented on by Abdul-Nour
1984) calculates it may actually be as large as 30,000-
80,000 species. This is an increase by a factor of no
more than about five. Another large group of homopter-
ans, the superfamily Coccoidea (5000 + described spp.,
Kosztarab 1982; 6000 described spp., Miller & Koszt-
arab 1979), may c o n t a i n “ p e r h a p s as many
undescribed” species as described (Miller & Kosztarab
1979).
Hodkinson and Casson ( 1991) use the proportion of
undescribed to described species of Hemiptera and Ho-
moptera in samples collected in Sulawesi to estimate
that there may be a world total of 184,000-193,000
species in these orders. These figures are encouragingly
consistent with the factors of increase already men-
tioned.
So How Many Insects Are There?
Regardless of the relative size of the major orders, esti-
mates of 30 or 50 million insect species (see Introduc-
tion) seem incompatible with our present understand-
ing of the richness of the major insect taxa as outlined
here. Among the assessments of the size of various su-
perfamilies and families, and what they predict about
the sizes of the major orders, there is little to encourage
belief in such large numbers. Rather, a figure of less than
10 million seems more tenable, and one of around five
million feasible.
If true, this means that a tenth or more of the world’s
Global lnsect Species Richness 289
insects have already been described. Virtually all of the
estimates of total species numbers cited for individual
families and superfamilies have involved increases over
described species of less than tenfold, and most are con-
siderably less. This is illustrated in Figure 2, which in-
cludes estimates for a heterogeneous collection of other
insect groups that have not previously been mentioned.
There is no strong tendency for the ratio of the esti-
mated total numbers of species to numbers of described
species to increase with the number of described spe-
cies. Large superfamilies and families do not seem to be
believed to be consistently proportionately more poorly
known.
In a similar vein it is interesting to contrast estimated
numbers of species in insect groups in the Nearctic, the
best known of the biogeographic regions, with estimates
of the global richness of those same groups (Fig. 3). For
many groups about ten times as many species are esti-
mated to exist worldwide than are believed to occur in
the Nearctic. In all cases the factor is less than one hun-
dred times. Given an upper estimate of around 150,000
described and undescribed species of insects in North
America alone (Kosztarab & Schaefer 1990) this rela-
tionship predicts more than a million insects world-
wide. The lack of estimates of numbers of additional
species in the rest of the Nearctic, however, limits the
usefulness of these figures.
The insects can be regarded as one of three major
radiations that have generated the bulk of the earth’s
species, the others being the nematodes and the fungi.
Hawksworth (Table 1 in diCastri & Younes 1990) has
recently proposed that around 8 percent of species of
fungi have been described. Applied to the insects this
proportion generates an estimate of around 10 million
species, which, considering that the insects are likely to
be better known than the fungi, provides another hint
that they number less than this figure.
Interestingly, prior to Erwin’s (1982) paper, most es-
timates of the numbers of insect species were of a sim-
ilar magnitude to that implied here. Sabrosky (1952)
cites guesses of 2.5 to 10 million species of insects,
Brues et al. ( 1954) suggest figures of 3.75 to 7.5 million
insect species, and there are several estimates of total
animal species numbers of less than 10 million (summa-
rized in May 1990).
Additional Evidence
Some additional, more circumstantial evidence exists
which, while being insufficient by itself, tends to sup-
port the idea that insect species numbers may not be as
vast as has been claimed recently.
(i) The principal justification for dramatic increases
in the magnitude of estimates of the numbers of insect
species has been the discovery of high species richness
Conservation Biology
Volume 5, No. 3, September 1991
290 Global Insect Species Richness
7 importance of this finding depends upon the bulk of
c cies of trees. The last point has been discussed at length
o w
0 2 4 of significant numbers of unknown canopy species in
Log. estimuted no. o f
described species
Figure 2. Relationship between the total numbers of
extant species estimated for a variety of insect or-
ders, superfumilies, and families, and the number of
species ulreudy described (both variables logIo trans-
formed). Data for Odonutu (Corbet 1979; Duvies &
Tobin 1984, 1985), Triductyloideu ( K K Guntbw,
pers. comm.), Dermupteru (Nickle & Sukui 1990),
Embiopteru (Ross 1982), Psocopteru (C A! Smitbers,
pers. comm.), Anopluru (Kim et ul. 19901, Tbysuno-
pteru (L. M o u n d , pers. comm.), Cicadellidae (Koszt-
urab 1982; Hamilton 19841, Scbizopteridue (Sluter
19821, Psylloideu (0.Hollis, pers. comm.1, Apbididue
( R Blackman, pers. comm.), Cqlonidue (S. A
Slipinski, pws. comm.), Geotrupidue (H. F. Howden,
pers. cornm.), Eucnemidue CI; Muon4 pers. comm.),
Pselupbidue (D. S. Cbundler, C Besucbet, pers.
comm.), Cbysomelidae ('Joliuet, 1988),Alticinue
(D. G. Furtb, pers. comm.), Rbysodidue (R 7: Bell,
pers. comm), Trogidue (C H. Scboltz, pers. comm.),
Curculionoideu (G. Kuscbel, pers. comm.), Curabidue
( N I? Stork, pers. comm.), Stapbylinidue (P. M. Hum-
mond, pers. cornm.), Dasyceridue (I. Lobl, pers.
comm.), Inopeplidue and Passundridue (S. A Slipin-
ski, pers. comm.), Antbribidue (B. D. Valentine, pers.
comm.), Strepsiptera (Brown, 1982b), Mecopteru
(G. W Byers, pers. comm.), Sipbonupteru (Kim et ul.
1990), Drosopbilidue (22. Grimaldi, pers. comm.),
Pboridae (R. H. L. Disney, pers. comm.), Tricbopteru
(P. Barnur4 pers comm.; Scbmid 19681, Cynipoideu
(Guuld & Bolton 19881,Bruconidue (Mason 1979;
Acbterberg 1984; Gauld & Bolton 19881,Icbneu-
monidue (Townes 1969), Omyridue ( P Hunson,
Conservation Biology
Volume 5 , No. 3, September 1991
Gaston
in the canopy of tropical forests (Erwin 1 9 8 3 ~ )The .
canopy insects being undescribed, their being largely
restricted to the canopy, and their exhibiting a high
degree of specialization in occurrence on particular spe-
elsewhere (Stork 1988; May 1990), it being generally
concluded that levels of specificity may not be particu-
larly high. Unfortunately, few data are available on the
proportion of canopy species that have been described.
Although samples have been obtained by fogging forest
canopies in several tropical regions ( e g , Erwin 1983b;
Erwin & Scott 1980; Stork 1988, 1991; Stork & Bren-
dell 1990) in all cases our general knowledge of the
insect faunas of these areas is rudimentary. Thus, we
have no context in which to place claims of large num-
bers of undescribed arboreal species, as virtually all in-
sects found in these regions, be they from the canopy or
not, are undescribed. There has been no demonstration
areas where the insect faunas are thought to be reason-
ably well known.
The latter test is problematical, because tropical in-
sect faunas are so poorly studied. Nonetheless, it could
very well fail for the simple reason that a high propor-
tion of insect species sampled from the canopy are also
found elsewhere. Brown and Hodkinson (1989) sum-
marize some of &heevidence, and Hammond ( 1990) has
recently shown that some 85 percent of beetle species
collected at canopy level (using fogging, malaise traps,
and baited traps) in the enormous sampling program in
Dumoga-Bone National Park, Sulawesi, were also caught
using ground-level trapping methods (e.g., flight-
intercept traps, malaise traps, litter sampling). This gen-
eral picture should perhaps come as no surprise.
Ground-level traps are often set at forest edges and in
tree-fall gaps, which may be habitats more similar to
forest canopies than the understory itself and thus will
catch canopy species. Moreover, many species probably
pass only part of the day, season, or life cycle in the
canopy. Lower levels of the forest may have to be visited
for reproduction, acquisition of particular resources
( e g ,parasitoids may visit understory plants for nectar),
or to pass less favorable periods (e.g.,Rees [ 19831found
that the proportion of Hemiptera flying at different
heights in a forest was related to the amount of rainfall).
Equally, many species seem to use the canopy for rea-
sons unassociated with feeding (e.g., Stork 1987,
1991). Furthermore, many insects may have to make
pets. comm.), Pelicinidue (I D. Gaul4 pers. comm),
Cbalcidoideu (Noyes 1978, 19901, Formicidue (B.
Bolton, E 0. Wilson, pers. comm), Vespidae (J M.
Curpenter, pers. comm.), and Apidae (Micbener 1974;
Guuld & Bolton 1988; Roubik 1989).
Gaston
/ /
0-l
I I
0 2 4 fact have extensive geographical ranges. Richards
Log. e s t i m a t e d no. of
N e a r c t i c species
Figure 3. Relationship between the total numbers of
extant species estimated for a variety of insect or-
ders, superfamilies, and families, and the number of
species estimated to occur in the Nearctic (incl. Mex-
ico) (both variables log,,J transformed). Data (with
sources as in the legend to Figure 2 unless o t h m i s e
indicated) for Tridactyloidea, Tbysanoptera, Psyl-
loidea, Aphididae, Ceylonidae, Eucnemidae, Pse-
laphidae, Alticinae, Rhysodidae, Trogidae, Dasycer-
idae, Inopeplidae, Pussandridae, Anthribidae,
Mecop tera, Drosophilidae, Phoridae, Trichoptera,
Rhopalocera (R. I. Vane-Wright, pers. comm.), Ich-
neumonidae (Townes 1969;I. D. Gauld pers. comm.),
Pelecinidae, Chalcidoidea ( ‘ Noyes, pers. comm.),
J
Formicidae, and Vespidae.
extensive movements around tropical forests to find
widely scattered resources produced in the high-species
richness, low-abundance patterns typical of such habi-
tats. Thus, specialist herbivores, for example, may have
to travel large distances to find host plants, and their
enemies likewise to find them. Some groups are clearly
adapted for high mobility (e.g., Gauld & Gaston, ms),
and it is far from clear that movements are best made in
the canopy. Even for species in which the majority of
individuals spend most of their time in the treetops,
specimens will undoubtedly turn up elsewhere.
In summary, the proportion of tropical forest insects
that are canopy specialists is unclear but their contribu-
tion to global richness estimates may be far less than has
been suggested.
(ii) Although the alpha diversity of insect species may
be very high in tropical forests, truly vast numbers of
species can only be considered realistic if the geograph-
ical ranges of most are small (beta diversity is high).
Global Insect Species Richness 291
Unfortunately, some taxonomic work on tropical insect
groups tends to lend support to a belief in the predom-
inance of localized distributions which may yet prove
misplaced. In short, revisionary works that concern lo-
cal faunas and fail to consider the possibility that the
species may have previously been described from places
many thousands of miles away may lead to artefactual
impressions of high levels of local endemicity. Although
clearly much greater undertakings, regional and partic-
ularly global revisions are much to be preferred. Ideally,
these should involve specimens from a large number of
localities, as it is only then that patterns of morpholog-
ical variation can be clearly seen and the dangers of
assigning specimens of one species to two or more
names avoided.
It is evident that many tropical insect species do in
( 1978), for example, states that Neotropical vespids
(one of the better known insect groups in the tropics)
tend to be very widely distributed at the species level,
apparently to a greater extent than are the birds. Simi-
larly, at least some Neotropical dolichopodids (Diptera)
have been recorded from a wide range of localities
(Robinson 1970), and three largely unrelated genera of
thrips (Thysanoptera) have representatives that have
been recorded at localities stretching from New York
and Iowa to southern Brazil (Mound 1972, 1976;
Mound & O’Neill 1972). Bourek (1988) comments on
how widespread some of the chalcidoids have been
found to be, while Janzen (1988) points out that of the
2000 macrolepidoptera species recorded for Santa Rosa
National Park, Costa Rica, more than 80 percent had
previously been described, but from other localities in
their “large” ranges. Indeed, it seems likely that, as with
other animal taxa such as birds and mammals, the bulk
of the insect species of lowland tropical regions will
have comparatively large ranges, while those of high
altitudes will show high degrees of local endemicity.
Although the data are as yet limited, it is interesting to
observe that some lowland tropical sites appear to con-
tain very high proportions of the species in the fauna of
the region to which they belong, though how represen-
tative these sites are remains to be seen. It should also
be noted more generally that many insect species have
ranges that extend into both temperate and tropical re-
gions.
Knowledge of the pattern of range sizes among trop-
ical insects could provide a crucial test of global species
richness estimates. Although our level of understanding
is as yet insufficient, such examples as these may add
some balance to pictures of the tropics in which every
patch of habitat is populated by its own endemics.
(iii) The potential role of differences in the propor-
tions of the major insect orders at different latitudes in
shaping species richness estimates has already been
mentioned. Nonetheless, it bears emphasis that the
(:onservation Biology
Volume 5, N o 3, September 1991
292 Global Insect Species Richness
number of insect groups for which rigorous data on the
relationship between latitude and species numbers are
available is small (examples include termites [ Isoptera],
Collins 1989; treehoppers [ Homoptera: Membracidae],
Wool & Olmstead 1984; semiaquatic bugs [Herniptera:
Gerromorpha], Andersen 1982; swallowtails [ Lepi-
doptera: Papilionidae], Scriber 1973, 1984; ichneu-
monids, Janzen 1981; Gauld 1986>. The ichneumonids
have attracted a greal deal of attention as a large group
whose richness does not increase in tropical regions
relative to temperate ones, and in this context there has
been much comment on the problems of demonstrating
latitudinal changes in the richness of insect taxa (e.g.,
Hespenheide 1978; Morrison et al. 1979). This raises
the interesting question of how many other groups have
extratropical peaks of species richness. We know of
some already (e.g., aphids [ Homoptera: Aphididae],
Dixon et al. 1987;Byrrhidae, Haliplidae, Silphidae, Cryp-
tophagidae, Lathridiidae [all Coleoptera], P. M. Ham-
mond, pers. comm.; bombyliids (Diptera), Hull 1973;
bees, Michener 1979; sawflies [ Hymenoptera: Sym-
phyta]) and the presence of yet more may further re-
duce estimates of global species numbers of insects.
Gauld and Gaston (ms) suggest that there are reasons to
believe that braconids and tachinids, for example, may
not show dramatic increases in species richness in trop-
ical regions.
(iv) As a final and more anecdotal observation, the
proportion of undescribed species that many insect tax-
onomists working with rain forest material meet seems
insufficient to just@ estimates of vast species numbers.
This point has been made by Monteith (1990) in his
criticism of K. L. Kitching’s comments on the species
richness of the insects associated with the canopy of
Australian rain forest. It is also evidenced by Gauld’s
( 1991 ) failure to continue to find many new species to
add to his total of 2000 ichneumonids for the whole of
Costa Kica. This despite about 90 percent of the species
being undescribed at the outset of the project and this
being one of the largest families of insects.
In Conclusion
This paper relies heavily upon the assessments of indi-
vidual systematists of the size of their specialist groups
in concluding that insect species probably do not num-
ber several tens of millions. The accuracy of such esti-
mates is undoubtedly highly variable, depending partic-
ularly upon the experience of the various systematists of
tropical faunas. Minimum estimates are more readily
constructed than are upper limits to species numbers,
and it is plausible that figures for at least some groups
are conservative. Nonetheless, they uniformly fail to
support Erwin’s (1982, 1988) estimates of insect spe-
cies richness. Moreover, they provide the only alterna-
Conservation Biolo&y
Volume 5, No. 3, September 1991
Gaston
tive to date to calculations similar in nature to those of
Erwin, and with equally questionable assumptions.
Although 1 hope it is self-evident, it should perhaps be
stated plainly that concluding that there are fewer insect
species than some have suggested provides no justifica-
tion for devaluing biodiversity, worrying less about its
fate, or lessening efforts to protect it. Whether there are
five, ten, fdteen, or more million species of insects in no
way alters their individual significance, nor the signifi-
cance of the crisis with which they are currently faced.
Acknowledgments
This paper has only proven feasible through the willing-
ness of many systematists to share their ideas as to the
sizes of their specialist groups. To all of them I am very
grateful. I alone can be held responsible for the ways in
which their insights have been used here. This work has
profited greatly from discussions with, and the com-
ments of, Kees van Achterberg, Barry Bolton, Henry Dis-
ney, Sian Gaston, Ian Gauld, Peter Hammond, Paul Han-
son, Dave Hollis, Chris Lyal, Bob May, Laurence Mound,
John Noyes, Gaden Kobinson, Nigel Stork, and Dick
Vane-Wright. Paul Barret kindly helped collate some of
the data.
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