The pest termites of South America: taxonomy, distributionand statusR. ConstantinoDepartamento de Zoologia, Universidade de Brasılia, Brasılia, Brazil

Ms. received: February 23, 2001; accepted: March 20, 2002

Abstract: A total of 77 species of termites have been reported as structural or agricultural pests in South America.These records are reviewed, with indication of the damage they cause, their pest status, distribution and sources ofinformation. Among these, 40 species are reported as structural pests, 53 species as agricultural pests and 15 species as

both. However, only 18 species are considered major pests and the status of most of the others is uncertain. Uncertaintyabout status is much higher among agricultural pests. Specific taxonomic problems are discussed and distribution mapsof the main pest genera and species are presented. The most important structural pests are the introduced Coptotermeshavilandi and Cryptotermes brevis and the native Nasutitermes corniger. Reticulitermes was introduced into Uruguay

and Chile, where it is the only serious pest. The main agricultural pests are Heterotermes spp., Nasutitermes spp.,Cornitermes spp., Procornitermes spp. and Syntermes spp., and the most affected crops are sugarcane, upland rice andeucalyptus.

1 Introduction

Termites are abundant and diverse in most parts ofSouth America, particularly in tropical lowland for-ests, savannas and grasslands. Several native termitespecies have been reported as structural or agriculturalpests, and some structural pests have been introducedfrom other regions. As natural habitats are graduallyreplaced by urban and agricultural space, new pests arediscovered, while the increasing trade with othercontinents facilitates the introduction of new pests.However, the information about the pest termites ofSouth America is still very limited. There are noestimates of the economic losses caused by termites orthe relative importance of each species. Agriculturaltermite pests are more numerous and much less studiedthan urban pests. Furthermore, termite research andexpertise in South America are highly concentrated inBrazil, whereas the fauna and termite problems oflarge countries such as Colombia are virtually un-known.There are about 400 termite species recorded for

South America (Constantino, 1998). Five families arepresent: Kalotermitidae (53 species), Rhinotermitidae(19), Serritermitidae (2), Termitidae (325), and Term-opsidae (1). Nasute termites (Nasutitermitinae) aredominant both in abundance and diversity, comprising54% of all species. At least six species were introducedfrom other regions and became established as urbanpests: Cryptotermes brevis, Cryptotermes dudleyi, andCryptotermes havilandi (Kalotermitidae) and Coptot-ermes havilandi, Reticulitermes lucifugus and Reticulit-ermes santonensis (Rhinotermitidae) (Constantino,

1998; Ripa and Castro, 2000). In general, the taxonomyof South American termites is far from complete, thereare many undescribed species, and very few generahave been properly revised.Termite damage in South America was first repor-

ted by European naturalists (Pohl, 1832; Hagen, 1855).The work of Silvestri (1903) in Argentina, Brazil andParaguay, of Holmgren (1906) in Bolivia and Peru, ofEmerson (1925) in Guyana, of Snyder (1926) in Boliviaand Brazil and of Araujo (1958a, b, 1977) in Brazil,established the groundwork for the scientific study ofSouth American termites. A list of the pest termitesreported in Brazil was presented by Silva et al. (1968),mostly based on the literature. Fowler and Forti

(1990) reviewed the termite problem in Brazil, pre-senting a highly incomplete and incorrect list ofspecies, which was severely criticized by Mill (1991,1992), who presented an overview of the termite pestsof Brazilian Amazonia. Torales et al. (1997) reviewthe Argentinian termite fauna and Torales (1998) theireconomic importance, especially in the CorrientesProvince. The termite pests of Uruguay were reviewedby Aber (1998), where an introduced Reticulitermesseems to be the only important species. Lefeuve (1990),surveyed the termites of French Guiana, indicatingsome that cause structural damage. The termiteproblem of Trinidad was reviewed by Adamson

(1946). For other countries, there are isolate recordsin publications dealing with taxonomy, control andbiology.In this paper, the taxonomy, distribution and status

of the pest termites of South America are reviewed,with new records.

JEN 126 (2002)

J. Appl. Ent. 126, 355–365 (2002)� 2002 Blackwell Verlag, BerlinISSN 0931-2048

U. S. Copyright Clearance Center Code Statement: 0931–2048/2002/2607–0355 $ 15.00/0 www.blackwell.de/synergy

2 Methods

Most pest records presented here come from a critical reviewof the literature, but some are based on the author’s own,unpublished data. The lists in tables 1–5 cover mainlandSouth America and Trinidad; other islands are not included.In some cases the identifications have been corrected byre-examination of relevant specimens. It is very difficult topresent an accurate list due to taxonomic uncertainties.Specific taxonomic problems are discussed in the text. Thedetermination of pest status in tables 1–5 is based onqualitative information. A species is considered a major pestif it consistently causes significant damage and a minor pest ifit causes damage only occasionally. A question markindicates that the information available is insufficient todetermine status. However, most of these species are prob-ably minor pests or not pests. Specimens examined aredeposited in the entomological collection of the University ofBrasılia. Taxonomic authorities and additional taxonomicdata can be found in Constantino (1998). Distribution maps(figs 1–7) were prepared with the Generic Mapping Tools(Wessel and Smith, 1998).

3 Taxonomy and distribution of pest termites

A total of 77 termite species reported as pests in SouthAmerica are presented in tables 1–5, corresponding to19% of all species. Among these, 40 species (10%) arereported as structural pests, 53 species (13%) asagricultural pests and 15 species (4%) as both. How-ever, only 18 species (4%) are considered major pestsand 14 (3.5%) minor pests. The status of the rest isuncertain and most are likely to be very minor or notpests. Uncertainty about status is much higher amongagricultural pests (73% uncertain) compared to struc-tural pests (40%).

3.1 Kalotermitidae

A few introduced Cryptotermes (fig. 1) are the mainstructural pests of this family. This genus was revisedby Bacchus (1987) but there are several undescribedspecies in South America and some may representminor pests. Cryptotermes domesticus has been recor-ded in Trinidad (Araujo, 1977), but this record has notbeen confirmed (Scheffrahn and Krecek, 1999). Thetime and place of introduction of these species isuncertain, but C. brevis has probably been present formore than 100 years.

3.2 Rhinotermitidae

The important pest species belong to genera Coptot-ermes, Heterotermes and Reticulitermes (table 2).There are at least two native Coptotermes species inSouth America: Coptotermes niger and Coptotermestestaceus, but apparently under the second name thereis a complex of several similar species, most of whichare undescribed. Coptotermes havilandi was intro-duced from Asia and became established along thecoastal region of Brazil. It was first recorded in Rio in1923 (Araujo, 1970) and its range seems to have beencontinuously expansing since then (fig. 2). Six speciesof Heterotermes occur in South America and their T

able

1.Dry-woodtermites(Kalotermitidae)reportedaspestsinSouthAmerica.Someoftheseidentificationsaredoubtful;seetext.Thequestionmarkindicatesthatthe

peststatusofagivenspeciesisunknown,butmostoftheseareprobablyveryminorpests.AllKalotermitidaearewood-feeders

Species

Structural

peststatus

Agriculturaldamage

Speciesdistribution

References

Crops

Status

Comatermesperfectus

coffee

?Colombia,Peru,Trinidad,Venezuela

Weid

ner(1980)

Cryptotermesbrevis

major

widespread(introduced)

Araujo(1958a);B

andeir

aetal.(1998);L

efeuve(1990);S

ilvaetal.(1968)

Cryptotermescubicoceps

?fruittrees

Guyana,?Brazil

Mil

l(1991)

Cryptotermesdomesticus

?Trinidad(introduced)

Araujo(1977)

Cryptotermesdudleyi

major

Brazil,Colombia,Trinidad(introduced)

Araujo(1977);B

andeir

aetal.(1989)

Cryptotermeshavilandi

major

Brazil,Guianas(introduced)

Araujo(1958a,1977)

Cryptotermessp.

?Brazil

Constantin

o(unpublisheddata)

Eucryptotermeswheeleri

minor

Brazil

Constantin

o(1997)

Glyptotermespellucidus

?fruittrees

?Guyana,Venezuela,Brazil?

Mil

l(1991)

Neotermescastaneus

fruittrees

?Brazil?,Chile,Colombia,Trinidad,Venezuela

Bequaert(1925)

Neotermeschilensis

minor

Chile

Goetsch(1933);R

ipaandC

astro(2000)

Neotermesfulvescens

fruittrees

?Brazil,Argentina

Sil

vaetal.(1968)

Neotermesparaensis

fruittrees

?Brazil

Mil

l(1992)

Neotermeswagneri

fruittrees

?Brazil

Sil

vaetal.(1968);M

ill(1992)

Rugitermesoccidentalis

fruittrees

?Brazil,Argentina

Sil

vaetal.(1968);M

ill(1992)

Tauritermesvitulus

minor

Brazil

AraujoandF

ontes(1979)

Tauritermessp.

minor

Brazil

Constantin

o(unpublisheddata)

356 R. Constantino

taxonomy was revised by Constantino (2001). All ofthem are pests, but Heterotermes tenuis and Heterot-ermes longiceps are the most important (fig. 3).Reticulitermes lucifugus was reported as a majorintroduced pest in Uruguay by Aber and Fontes

(1993), who estimated that it was introduced in the1960s. More recently, Reticulitermes santonensis wasintroduced in Chile (Ripa and Castro, 2000). How-ever, according to Jenkins et al. (2000), R. santonensisis a synonym of R. flavipes and it is likely that bothintroductions into South America correspond to thesame species.

3.3 Termitidae

3.3.1 Apicotermitinae

Most species are harmless humus-feeders, and onlyAnoplotermes pacificus and a few unidentified speciesof Anoplotermes, Aparatermes and Grigiotermes(table 3) have been reported as pests of several crops(Mill, 1992; Czepak et al., 1993). The taxonomy ofthe Neotropical Apicotermitinae, all soldierless, ischaotic and species identification is impossible inmost cases.

3.3.2 Nasutitermitinae

These can be divided into true nasutes (table 4), whichare pantropical, and mandibulate nasutes (table 5),which are endemic to the Neotropics. The taxonomy ofNasutitermes, the most diverse nasute genus, is con-fused and species identification is difficult. Nasutiter-mes corniger, the most important pest of this genus,has often been misidentified. Nasutitermes araujoi,Nasutitermes globiceps, Nasutitermes tatarendae andpossibly N. costalis are all synonyms of N. corniger(Constantino, unpubl.). The species identified asN. nigriceps in South America is probably eitherN. acajutlae or N. macrocephalus. Nasutitermes acajut-lae was considered a synonym of N. nigriceps, but wasrevalidated by Thorne et al. (1994). The records ofN. rippertii (Silva et al., 1968) and of �Obtusitermes sp.�(Dietrich, 1989; Berti-Filho, 1995) in Brazil are mostlikely incorrect because these are outside their knowndistribution.Among the mandibulate nasutes (table 5), the most

important species are in the genera Cornitermes(fig. 5), Procornitermes and Syntermes (fig. 6). Thesewere revised by Emerson (1952), Cancello (1986,1989) and Constantino (1995) and identification isrelatively easy. Records of Cornitermes snyderi as anagricultural pest in Brazil are incorrect. Cancello

(1989) suggested that C. snyderi and Cornitermessilvestrii are synonyms and the first was informallyadopted as the valid name. However, if they aresynonyms, C. silvestrii is the older name and if not,the species that occurs in Brazil is C. silvestrii anyway(C. snyderi is originally from Bolivia). Syntermesnanus and Syntermes molestus were treated as a singlespecies for a long time (Constantino, 1995), and mostpublished records of S. molestus as an agriculturalpest apparently refer to S. nanus.T

able

2.Subterraneantermites(Rhinotermitidae)reportedaspestsinSouthAmerica.Someoftheseidentificationsaredoubtful;seetext.Thequestionmarkindicatesthat

thepeststatusofagivenspeciesisunknown,buttheseareprobablyveryminorpests.AllRhinotermitidaearewood-feeding

Species

Structural

peststatus

Agriculturaldamage

Speciesdistribution

References

Crops

Status

Coptotermeshavilandi

major

Brazil(introduced)

Araujo(1958a);

Costa-L

eonardoetal.(1999)

Coptotermesniger

major?

fruittrees,palm

trees

?Colombia,Brazil?

Araujo(1970);M

ill(1992);S

nyderandZ

etek(1946)

Coptotermestestaceus

minor

eucalyptus,cassava,

rubbertree

?Bolivia,Brazil,Colombia,Ecuador,

Guianas,Peru,Trinidad,Venezuela

Bandeir

a(1981);B

andeir

aetal.(1989);S

antosetal.(1990);

Sil

vaetal.(1968)

Dolichorhinotermeslatilabrum

?Bolivia,Brazil?

Mil

l(1991)

Heterotermesassu

minor

Brazil

Constantin

o(2001)

Heterotermesconvexinotatus

major

passionfruit,maize

?Venezuela,Colombia

Dom

inguez-G

ilandM

cP

heron(1992);C.J.R

osales(pers.comm.);

SnyderandZ

etek(1946)

Heterotermescrinitus

sugarcane,peachpalm

?Brazil,Guyana,Venezuela

Guagliu

mi(1962);M

ill(1992)

Heterotermessulcatus

minor

Brazil

Constantin

o(2001)

Heterotermeslongiceps

minor

sugarcane

major

Brazil,Argentina

Torales(1998),B

andeir

aetal.(1998);C

onstantin

o(unpubl.data)

Heterotermestenuis

minor

sugarcane,eucalyptus

major

widespread

Bandeir

aetal.(1989);C

onstantin

o(2001);L

efeuve(1990);

NovarettiandF

ontes(1998);S

ilvaetal.(1968)

Reticulitermessantonensis

major

major

Chile(introduced)

Rip

aandC

astro(2000)

Reticulitermeslucifugus

major

major

Uruguay(introduced)

AberandF

ontes(1993)

Rhinotermesmarginalis

minor

minor

Amazonia,Guianas

Bandeir

aetal.(1989);M

ill(1991)

Pest termites of South America 357

Table

3.TermitesofSubfamiliesApicotermitinae(soldierlesstermites)andTermitinaereportedaspestsinSouthAmerica.Someoftheseidentificationsaredoubtful;see

text.Thequestionmarkindicatesthatthepeststatusofagivenspeciesisunknown,buttheseareprobablyveryminorpests

Species

Structural

peststatus

Agriculturaldamage

Feedinghabit

Distribution

References

Crops

Status

APICOTERMITINAE

Anoplotermespacificus

eucalyptus,

variouscrops

?humus?

Argentina,Bolivia,Brazil,

Paraguay,Peru

Mil

l(1992);S

ilvaetal.(1968)

Aparatermessp.

rice

?leaforgrass-litter

Brazil

Czepaketal.(1993)

Grigiotermessp.

rice

?humus

Brazil

Czepaketal.(1993)

TERMITINAE

Amitermesamifer

minor

leaforgrass-litter

Argentina,Brazil,Paraguay

Bandeir

aetal.(1998);T

orales(1998)

Amitermesexcellens

?peanut,soybean,

cassava,cashew

?decayingwood

Brazil,Guyana

Bandeir

a(1998);M

ill(1992);S

ilvaetal.(1968)

Amitermesforeli

passionfruit

??

Colombia,Venezuela

Dom

inguez-G

ilandM

cP

heron(1992)

Amitermessp.

sugarcane

??

Brazil

NovarettiandF

ontes(1998)

Cylindrotermessp.

sugarcane,

eucalyptus

?wood

Brazil

NovarettiandF

ontes(1998);

Berti-

Fil

ho(1995)

Microcerotermesarboreus

minor

passionfruit

minor

wood

Amazonia,Guianas,Trinidad

Bandeir

aetal.(1989);D

om

inguez-G

iland

McP

heron(1992)

Microcerotermesexiguus

?wood

Brazil,Colombia,Trinidad,Venezuela

Guagliu

mi(1966);M

ill(1991)

Microcerotermesstrunckii

minor

fruittrees

minor

wood

Argentina,Brazil

Mil

l(1992);T

orales(1998)

Neocapritermesopacus

eucalyptus

major

intermediate

Argentina,Bolivia,Brazil,Ecuador,

Paraguay,Peru

Sil

vaetal.(1968);

Berti-

Fil

ho(1995)

Neocapritermesparvus

sugarcane

major

intermediate

Brazil,Paraguay

NovarettiandF

ontes(1998)

Termesfatalis

?fruittrees

?decayingwood

Amazonia,Guianas,Trinidad

Mil

l(1992)

Termessaltans

sugarcane

?decayingwood

Brazil,Argentina,Paraguay,Uruguay

Sil

vaetal.(1968)

358 R. Constantino

Table

4.Nasutetermites(Termitidae:Nasutitermitinae)reportedaspestsinSouthAmerica.Someoftheseidentificationsaredoubtful;seetext.Thequestionmarkindicates

thatthepeststatusofagivenspeciesisunknown,buttheseareprobablyveryminorpests

Species

Structural

peststatus

Agriculturaldamage

Feeding

habit

Distribution

References

Crops

Status

Convexitermesconvexifrons

minor

wood

Brazil,Peru

Constantin

o(unpubl.data)

Cortaritermesfulviceps

minor

wood;litter

Argentina,Brazil,Paraguay,Uruguay

Torales(1998)

Nasutitermesacangussu

?wood

Brazil

Bandeir

a(1998)

Nasutitermesaquilinus

minor

eucalyptus

wood

Argentina,Brazil,Paraguay

Laffontetal.(1998);T

orales(1998)

Nasutitermesbivalens

coffee

??

Brazil

Sil

vaetal.(1968)

Nasutitermesbrevioculatus

sugarcane

??

Brazil,Bolivia,Paraguay

Sil

vaetal.(1968)

Nasutitermescallimorphus

?wood

Brazil

Bandeir

aetal.(1989)

Nasutitermescorniger

major

fruittrees

wood

widespread(Fig.7)

Bandeir

aetal.(1989);B

andeir

aetal.(1998);

Costa-L

eonardo(2000);S

nyderandZ

etek(1946);

Torales(1998)

Nasutitermescostalis

?fruittrees

?wood

?G

uagliu

mi(1966);M

ill(1992)

Nasutitermesephratae

minor

fruittrees

?wood

widespread

Lefeuve(1990);M

ill(1992);S

nyderandZ

etek(1946)

Nasutitermesguayanae

?wood

Brazil,Colombia,Guianas,Trinidad,

Venezuela

Bandeir

aetal.(1989)

Nasutitermesmeinerti

Ficussp.

??

Venezuela

Guagliu

mi(1966)

Nasutitermesnigriceps

minor

wood

?(seetext)

Bandeir

aetal.(1989);L

efeuve(1990);M

ill(1992)

Nasutitermesoctopilis

fruittrees

?wood

Guyana,BrazilianAmazonia

Mil

l(1992)

Nasutitermesperuanus

fruittrees

??

Bolivia,Brazil,Ecuador,Peru

Mil

l(1992)

Nasutitermessurinamensis

?fruittrees,maize

?wood

Amazonia,Guianas

Lefeuve(1990);M

ill(1992)

Subulitermessp.

eucalyptus

?humus

Brazil

Die

tric

h(1989)

Velocitermesglabrinotus

soybean,cassava

?grass-litter

Brazil

Mil

l(1992)

Pest termites of South America 359

3.3.3 Termitinae

Most species are humus-feeders and only Cylindroter-mes,Microcerotermes and a few species of other generaare wood-feeders and can damage wood (table 3). Thetaxonomy of most neotropical genera of this subfamily

Fig. 1. Distribution of introduced Cryptotermes speciesin South America. Cryptotermes brevis is present inmost urban areas within the grey zone. Sources: Araujo

(1977); Bandeira et al. (1989); Fontes and Araujo

(1999); Constantino (unpublished data)

Fig. 2. Distribution of Coptotermes species in SouthAmerica. Sources: Constantino (1998 and unpublisheddata); Costa-Leonardo et al. (1999)

Table

5.Mandibulatenasutetermites(Termitidae:Nasutitermitinae)reportedaspestsinSouthAmerica.Someoftheseidentificationsaredoubtful;seetext.Thequestion

markindicatesthatthepeststatusofagivenspeciesisunknown,buttheseareprobablyveryminorpests

Species

Agriculturaldamage

Feedinghabit

Distribution

References

Crops

Status

Armitermeseuamignathus

eucalyptus

?intermediate

Bolivia,Brazil,Paraguay

Sil

vaetal.(1968)

Cornitermesbequaerti

pasture

?intermediate

Brazil

Fernandesetal.(1998)

Cornitermescumulans

pasture,sugarcane,

eucalyptus

major

intermediate

Argentina,Brazil,Paraguay

Fernandesetal.(1998);N

ovarettiandF

ontes(1998);

Sil

vaetal.(1968)

Cornitermessilvestrii

sugarcane,pasture

major

grass

Bolivia,Brazil

Guagliu

mi(1971);F

ernandesetal.(1998);S

ilvaetal.(1968)

Embiratermessp.

eucalyptus

?intermediate

Brazil

Die

tric

h(1989)

Procornitermesaraujoi

eucalyptus,rice

major

intermediate

Brazil

Sil

vaetal.(1968)

Procornitermesstriatus

eucalyptus,rice,pineapple

?intermediate

Argentina,Brazil,Paraguay,Uruguay

Sil

vaetal.(1968)

Procornitermestriacifer

sugarcane,rice,coffee,

eucalyptus,maize

major

intermediate

Argentina,Bolivia,Brazil

Sil

vaetal.(1968)

Rhynchotermessp.

eucalyptus

?leaf-orgrass-litter

Brazil

Die

tric

h(1989)

Syntermesgrandis

sugarcane

?grass-litter

Bolivia,Brazil,Guianas

Guagliu

mi(1971);S

ilvaetal.(1968)

Syntermesinsidians

eucalyptus

?grass-litter

Brazil

Fonseca(1949)

Syntermesmolestus

sugarcane,rice

?leaf-orgrass-litter

Amazonia,Guianas

Guagliu

mi(1971);M

ill(1992)

Syntermesnanus

eucalyptus,rice

major

grass-litter

Argentina,Brazil,Paraguay

Fonseca(1949)

Syntermesobtusus

pasture

?grass-litter

Argentina,Bolivia,Brazil,Paraguay

Maric

onietal.(1976);S

ilvaetal.(1968);

Valerioetal.(1998)

360 R. Constantino

is confused and species identification is difficult.Among the economically important ones, only Neo-capritermes was taxonomically revised and is relativelyeasy to identify (Krishna and Araujo, 1968).

4 Urban pests

Urban pests are fewer and relatively better known thanthe agricultural ones. With the exception of N. cornigerin a few cities, the most important species wereintroduced: Cryptotermes brevis, Coptotermes havilandiand Reticulitermes spp. Cryptotermes brevis can befound in most cities except in southern and colderones, and has been estimated to be the most important

Fig. 3. Distribution of Heterotermes (grey area)and Reticulitermes (triangles) in South America.Heterotermes tenuis occurs in most of the grey area.Sources: Aber and Fontes (1993); Constantino (2001);Ripa and Castro (2000)

Fig. 4. Distribution of Nasutitermes (grey area) inSouth America. Sources: Bandeira et al. (1989),(1998); Costa-Leonardo (2000); Torales (1998);Torales et al. (1997); Constantino (unpublished data)

Fig. 5. Distribution of Cornitermes (grey area) inSouth America. Sources: Cancello (1989) and Con-

stantino (unpublished data)

Fig. 6. Distribution of Syntermes (grey area) in SouthAmerica. Source: Constantino (1995)

Pest termites of South America 361

termite pest in several of them. Coptotermes havilandicauses much more serious damage, but its distributionis more limited. In addition to the structural damage,termites may also affect urban trees and their moundsmay be a nuisance in some areas. Trees hollowed byCoptotermes are a potential cause of serious accidents.Coptotermes testaceus has been occasionally reportedas causing severe structural damage in some cities, butit seems to be relatively rare in urban areas. Severalspecies of Nasutitermes are able to damage wood andN. corniger seems to be the most important structuralpest of this group, having adapted well to the urbanenvironment. It occurs from Mexico to northernArgentina, and is mentioned as a major, widespreadurban pest in South America here for the first time. Asummary of the problems reported in some cities ispresented below. The locations of these cities areindicated in fig. 7.

4.1 Brazil

4.1.1 Belem

The largest city in Amazonia, with about 1.2 millionpeople. It differs from most Brazilian cities in thatwood houses are very common, especially among low-income families. Bandeira et al. (1989) studied the

wood pests of this city, recording 12 termite species.Nasutitermes spp. caused more than 50% of thedamage, and the single most important pest termitewas N. corniger. Coptotermes cf. testaceus was alsocommon and very destructive. Cryptotermes dudleyiwas the only common dry-wood termite there. Theyalso mention Heterotermes tenuis, Rhinotermes mar-ginalis and Microcerotermes arboreus as minor pests.

4.1.2 Joao Pessoa

A large city in the north-east of Brazil, with about500 000 people. Bandeira et al. (1998) recorded 10wood pests. The most important were Cryptotermesbrevis and Nasutitermes corniger (identified as Nasut-itermes sp. B, near N. tatarendae) followed by Hete-rotermes longiceps (listed as Heterotermes sp.). Minorpests include Amitermes amifer, Microcerotermes sp.and other Nasutitermes spp.

4.1.3 Brasılia

A large city built in the savanna (Cerrado) of centralBrazil about 40 years ago, with nearly 500 000 peopleand more than 2 million in the metropolitan region.More than 20 termite species live in the urban area ofBrasılia (Constantino and Dianese, 2001). Cryptotermesbrevis is the most important wood pest, followed byHeterotermes longiceps and H. tenuis. Cornitermescumulans builds large mounds, which are consideredan aesthetic problem in lawn and gardens. Syntermesnanus is a large termite which is abundant, can damageliving vegetation and its soldiers can bite people anddomestic animals.

4.1.4 Sao Paulo

The largest city in South America, with about 17million people. At least 20 termite species occur in thecity (Fontes, 1995; Constantino, unpubl.). The mostimportant pests are Coptotermes havilandi and Crypt-otermes brevis. Several other species have been repor-ted as pests, probably minor, including Nasutitermessp., Eucryptotermes wheeleri, Heterotermes assu andone undescribed Tauritermes.

4.2 Argentina: Corrientes

According to Torales (1998), the most important woodpest is Nasutitermes corniger, followed by Heteroter-mes longiceps. Minor pests include Amitermes amifer,Microcerotermes strunckii and Cortaritermes fulviceps.

4.3 Uruguay: Montevideo

According to Aber (1998), the only important termitepest in this city is Reticulitermes lucifugus, which wasintroduced in the 1960s. Before that time, termiteproblems were probably very minor in Uruguay.Bacchus (1987) reports the presence of Cryptotermesbrevis in Uruguay from a sample collected in 1920, butthis species is not mentioned by Aber and its currentstatus there is uncertain.

Fig. 7. Main pest termites in some South American cit-ies. Sources: Bandeira et al. (1989) (Belem); Bandeira

et al. (1998) (Joao Pessoa); Constantino (unpublisheddata) (Coro, Brasılia and Sao Paulo); Fontes (1995)(Sao Paulo); Torales (1998) (Corrientes); Aber

(1998) (Montevideo). For Brasılia and Sao Paulo, thenumber of species corresponds to all termites found inthese cities, most of them not pests. Introduced speciesare in bold face

362 R. Constantino

4.4 Venezuela: Coro

One of the oldest cities built by the Spanish in SouthAmerica, it has many historic buildings under officialprotection. These buildings are constantly attacked bytermites (Constantino, unpubl.). The wide adobe wallscontain straw, creating a very favourable environmentfor termites. The most common species found in thosebuildings were Amitermes foreli, Nasutitermes cf.acajutlae and Heterotermes convexinotatus. Amitermesforeli is a soil termite not expected to be found in ahouse, but it seems to live well inside the adobe wall.

5 Agricultural pests

The most affected crops in South America are sugar-cane, upland rice, and eucalyptus. A termite effect onpastures is controversial. Other crops affected includemaize, cotton, peanuts, soybean, coffee, cassava, fruittrees and some vegetables such as carrots. Termitedamage includes: (1) injury to roots; (2) injury to leavesand stems; (3) injury to woody tissue by kalotermitids;(4) injury to woody tissue by arboreal nesting speciessuch as Nasutitermes and Microcerotermes; (5) hol-lowing of tree trunks by Coptotermes. The mostimportant pests belong to genera Heterotermes, Na-sutitermes, Cornitermes, Procornitermes and Synter-mes. Besides injuring living plants, most Cornitermesbuild large mounds, which are considered a mechanicalproblem for the use of machinery in agriculture, reducethe useful soil surface and provide shelter for snakes,scorpions and spiders, among other problems. Fer-

nandes et al. (1998) claim that, at least in pastures,Cornitermes spp. are best defined as aesthetic pests,because farmers dislike their presence but they causelittle or no real damage to grass production. Coptot-ermes testaceus can damage cassava (Bandeira, 1981)and living trees (Santos et al., 1990). However, a largenumber of termite species is present in agrosystems,their identification is often uncertain, and the faunavaries among different regions. These are all nativespecies and, unlike the urban pests, they are associatedwith their natural biomes.The largest biome of South America is the Amazon,

with nearly 6 million km2 (33%). Agriculture in thisregion is mostly small-scale shifting cultivation andextensive pastures. There are also natural savannas inthe Amazon, which have been used as natural pasturesand, more recently, to grow soybean. Nasutitermesspp., Coptotermes testaceus and Heterotermes tenuismay cause variable damage to crops such as cassava,maize and fruit trees. Some pastures show a highdensity of Cornitermes mounds, which are considered aproblem. However, this is a large and heterogeneousregion and studies on the impact of termites inAmazonian agriculture are very limited.The second biome in South America is the Cerrado,

a kind of savanna which occupies about 2 million km2

(12%). Termites are extremely abundant and diverse inthis region, which is the most important agriculturalfrontier in Brazil. Large areas of Cerrado have beenreplaced by agrosystems, and the termites already

present may cause severe damage to some crops.Syntermes nanus, for example, which is abundant inthe Cerrado, is a pest of eucalyptus, upland rice andother crops. Fonseca (1949) reported that, in a localityin the State of Sao Paulo, Brazil, termites of the genusSyntermes killed up to 70% of the young eucalyptustrees in a Cerrado area. Cornitermes spp., Procornit-ermes spp. and Heterotermes spp. are also common inthe Cerrado.The third biome in size is the Brazilian Atlantic

Forest, which has already been mostly destroyed andreplaced by cropland and urban areas. Termites areless abundant and diverse in comparison with both theAmazon and the Cerrado. The termite pests presentare not very different from those of the Cerrado ingeneric composition, but the species may differ. Neo-capritermes opacus is a dominant species in some areasand a pest of some crops.The Caatinga, a xerophytic open forest or savanna

of north-eastern Brazil, is not an important agricul-tural region, and its termite fauna is poorly known,with estimates of more than 60% of undescribedspecies. The Chaco, a xerophytic open forest of Bolivia,Paraguay and northern Argentina, is also not import-ant for agriculture, and information on its fauna isvery limited. It seems to be similar to that of theCerrado. The colder, southern region which includesthe Pampa, Patagonia and temperate forests seems tobe relatively free of termite problems. The forests ofnorth-eastern Colombia and northern Venezuela areconnected to those of Panama, with similar termitefauna. The main pests are Nasutitermes corniger,N. ephratae and Heterotermes convexinotatus. Thetermite fauna of the Llanos (grassland) of Venezuelaseems to be composed mainly of mound-buildingNasutitermes and Velocitermes, but their importancein agriculture is unknown.

6 Discussion

Introduced structural pest termites are becoming amajor problem in South America. Cryptotermes brevisbecame the ubiquitous house termite a long time ago.Coptotermes havilandi is still expanding its range and islikely to eventually invade most of tropical SouthAmerica. Temperate South America, which was relat-ively free from termite problems, now faces theintroduction of Reticulitermes spp., which are likelyto invade Argentina, Paraguay and southern Brazil.Native urban pests are, in general, less important.

Exceptions are Nasutitermes corniger, which is themost important structural pest in some places (Ban-

deira et al., 1989; Torales, 1998) and some Heteroter-mes. New records of minor pests have been relativelycommon in many cities, and some of these mayeventually become important. Heterotermes assu, forexample, seems to have recently invaded cities from theneighbouring forests (Constantino, 2001).The termite problem in tropical agriculture was

reviewed by Wood (1996). Compared to Africa andthe Indo-Malayan region, where some Macrotermit-inae are major agricultural pests (Wood, 1996),

Pest termites of South America 363

termites are a lesser problem in South America. TheMacrotermitinae are absent in the New World, andthe major agricultural pests are Nasutitermitinae andRhinotermitidae. In general, knowledge about pesttermites in agriculture is more limited in SouthAmerica than in the Old World tropics. In additionto these differences, the major patterns are the same asthose mentioned by Wood (1996) and the mostaffected crops are also the same. Apparently, in somecases termites do not become pests because pesticidesused to control other pests reduce their populations.For this reason, the termite problem seems to be moresevere in small-scale subsistence cultivation wherepesticides are rarely used, such as reported by Mill

(1991) in the Amazon.Pest status is much easier to determine for structural

pests because they cause direct damage. Damage inagriculture and forestry, however, is usually indirect.Many authors assume that a species is a pest if it isobserved injuring a living plant, or worse, just becauseit is present in the agrosystem. More accurately, itshould be considered a pest only if it causes economicdamage, which is rarely estimated. Termites can injureroots, stems and leaves, but this may or may not resultin real loss of production. Damage by a given speciesmay be too rare to consider it a pest. Sometimes it isalso difficult to distinguish consumption of dead tissuefrom attack to living tissue. For these reasons, manyreports of termite problems are false.Taxonomy has been a major impediment to the

study and control of termites in South America due tothe large number of species, high proportion ofundescribed or poorly described species and lack ofproper taxonomic training of many authors. Very fewspecialists are able to identify neotropical termitespecies and many of the economically importantgroups lack proper taxonomic revision, resulting in alarge number of incorrect, doubtful or incompleteidentifications. The use of invalid or unavailable namesand misspellings are also very common. Under theseconditions it is very difficult to accurately store andretrieve information on the biology and economicimportance of each species. Several doubtful pestrecords have been repeatedly cited and even includedin entomological textbooks. A proper pest recordshould contain the following information: (1) anestimate of the economic damage; (2) the geographicdistribution of the problem; (3) how the species wasidentified; (4) place where voucher specimens aredeposited. Without voucher specimens, it is impossibleto verify any published identification.

Acknowledgements

This work was partially supported by a grant from theBrazilian National Council for the Development of Science(CNPq no. 301423⁄87–3). I thank Og DeSouza for reviewingthe manuscript.

References

Aber, A., 1998: Termite subterranea, xilofaga del Uruguay.Reticulitermes lucifugus (Isoptera, Rhinotermitidae).

In: Cupins: O desafio do conhecimento. Ed. by Fontes,L. R.; Berti-Filho, E. Piracicaba: FEALQ, pp. 437–451.

Aber, A.; Fontes, L. R., 1993: Reticulitermes lucifugus(Isoptera, Rhinotermitidae), a pest of wooden structures,is introduced into the South American Continent. Socio-biology 21, 335–339.

Adamson, A. M., 1946: Termites in Trinidad and Tobago.Trop. Agric. 23, 221–223.

Araujo, R. L., 1958a: Contribuicao a biogeografia dostermitas de Minas Gerais, Brasil. Arq. Inst Biol. 25,

219–236.Araujo, R. L., 1958b: Contribuicao a biogeografia dos

termitas de Sao Paulo, Brasil. Arq. Inst Biol. 25, 185–217.Araujo, R. L., 1970: Termites of the Neotropical Region. In:

Biology of Termites. Ed. by Krishna, K.; Weesner, F. M.New York: Academic Press, pp. 527–576.

Araujo, R. L., 1977: Catalogo dos Isoptera do Novo Mundo.Rio de Janeiro: Academia Brasileira de Ciencias.

Araujo, R. L.; Fontes, L. R., 1979: Notes on the Neotropicalgenus Tauritermes, with a new species from Brazil(Isoptera, Kalotermitidae). Revta bras. Ent. 23, 29–34.

Bacchus, S., 1987: A taxonomic and biometric study of thegenus Cryptotermes (Isoptera: Kalotermitidae). Trop.Pest Bull. 7, 1–91.

Bandeira, A. G., 1981: Ocorrencia de cupins (Insecta,Isoptera) como pragas de mandioca em Bujaru. Para.Acta Amazonica 11, 149–152.

Bandeira, A. G., 1998: Danos causados por cupins naAmazonia Brasileira. In: Cupins: O desafio do conheci-mento Ed. by Fontes, L. R.; Berti Filho, E. Piracicaba:FEALQ, pp. 87–98.

Bandeira, A. G.; Gomes, J. I.; Lisboa, P. L. B.; Souza, P. C. S.,1989: Insetos pragas de madeiras de edificacoes em Belem– Para. (Boletim Tecnico no. 101). Belem: EMBRAPA-CPATU.

Bandeira, A. G.; Miranda, C. S.; Vasconcellos, A., 1998:Danos causados por cupins em Joao Pessoa, Paraıba –Brasil. In: Cupins: O desafio do conhecimento. Ed. byFontes, L. R.; Berti Filho, E. Piracicaba: FEALQ, pp.75–85.

Bequaert, J., 1925: Neotermes injurious to living guava tree,with notes on other Amazonian termites. Ent. News 36,289–294.

Berti-Filho, E., 1995: Cupins e florestas. In: Aspectos atuaisda biologia e controle de cupins. Ed. by Berti-Filho, E.;Fontes, L. R. Piracicaba: FEALQ, pp. 127–140.

Cancello, E. M., 1986: Revisao de Procornitermes Emerson(Isoptera, Termitidae, Nasutitermitinae). Papeis AvulsosZool. (Sao Paulo) 36 (19), 189–236.

Cancello, E. M., 1989: Revisao de Cornitermes Wasmann(Isoptera, Termitidae, Nasutitermitinae). Ph.D. Thesis.Sao Paulo, Brazil: University of Sao Paulo.

Constantino, R., 1995: Revision of the neotropical termitegenus Syntermes Holmgren (Isoptera: Termitidae). Univ.Kansas Sci. Bull. 55, 455–518.

Constantino, R., 1997: Notes on Eucryptotermes with a newspecies from central Amazonia (Isoptera: Kalotermiti-dae). Sociobiol. 30, 125–131.

Constantino, R., 1998: Catalog of the living termites of theNew World (Insecta: Isoptera). Arq. Zool. 35, 135–231.

Constantino, R., 2001: Key to the soldiers of South AmericanHeterotermes with a new species from Brazil (Isoptera:Rhinotermitidae). Insect Syst. Evol. 31, 463–472.

Constantino, R.; Dianese, E. C., 2001: The urban termitefauna of Brasılia, Brazil. Sociobiol. 38, 323–326.

Costa-Leonardo, A. M., 2000: Biological data for theneotropical termite Nasutitermes globiceps (Isoptera:Termitidae: Nasutitermitinae). Sociobiol. 36, 63–78.

364 R. Constantino

Costa-Leonardo, A. M.; Barsotti, R. C.; Camargo-Dietrich,C. R. R., 1999: Review and update on the biology ofCoptotermes havilandi (Isoptera, Rhinotermitidae).Sociobiol. 33, 339–356.

Czepak, C.; Ferreira, E.; Nogueira, S. B., 1993: Identificacaoe quantificacao de cupins rizofagos na cultura do arroz desequeiro. Pesq. Agropec. Bras. 28, 871–875.

Dietrich, C. R. R. C., 1989: Ocorrencia de cupins (Insecta:Isoptera) em reflorestamento de Eucalyptus spp. Master’sThesis. Sao Paulo, Brazil: University of Sao Paulo.

Dominguez-Gil, O. E.; McPheron, B. A., 1992: Arthropodsassociated with passion fruit in western Venezuela.Florida Entomol. 75, 607–612.

Emerson, A. E., 1925: The termites from Kartabo, BarticaDistrict, Guyana. Zoology 6, 291–459.

Emerson, A. E., 1952: The neotropical genera Procornitermesand Cornitermes (Isoptera, Termitidae). Bull. Am. Mus.Nat. Hist. 99 (8), 475–540.

Fernandes, P. M.; Czepak, C.; Veloso, R. S., 1998: Cupinsde montıculos em pastagens: prejuızo real ou pragaestetica? In: Cupins: O desafio do conhecimento. Ed. byFontes, L. R.; Berti-Filho, E. Piracicaba: FEALQ,pp. 187–210.

Fonseca, J. P., 1949: Experiencias de combate quımico acupins subterraneos no Horto Floresta de Guaranı. Arq.Inst Biol. 19, 57–88.

Fontes, L. R., 1995: Cupins em areas urbanas. In: Aspectosatuais da biologia e controle de cupins. Ed. by Berti-Filho,E.; Fontes, L. R. Piracicaba: FEALQ, pp. 57–75.

Fontes, L. R.; Araujo, R. L., 1999: Os cupins. In: Insetos eoutros invasores de residencias. Ed. by Mariconi, F. A.M. Piracicaba: FEALQ, pp. 35–90.

Fowler, H. G.; Forti, L. C., 1990: Status and prospects oftermite problems and control in Brazil. Sociobiol. 17, 45–56.

Goetsch, W., 1933: Die chilenischen Termiten. Zool. Jahrb.Abt. Syst. 64, 225–243.

Guagliumi, P., 1962: Las plagas de la cana de azucar enVenezuela. Caracas: Ministerio de Agricultura y Cria.Centro de Investigacion Agronomica.

Guagliumi, P., 1966: Insetti e aracnidi delle pianti comuni delVenezuela segnalati nel periodo 1938–63. Rel. Monogr.Agr. Subtr. Trop. 86, 1–391.

Guagliumi, P., 1971: Entomofauna della canna da zucheronel nord-est del Brasile. Riv. Agric. Subtr. Trop. 65,

49–66.Hagen, H. A., 1855: Monographie der Termiten. Linn.

Entom. 10, 1–144.Holmgren, N., 1906: Studien uber sudamerikanische Term-

iten. Zool. Jahrb. Abt. Syst. 23, 521–676.Jenkins, T. M.; Verkerk, R. H. J.; Forschler, B. T., 2000:

DNA sequence data used to validate species taxonomyand origin of UK subterranean termite (Isoptera: Rhin-otermitidae) infestation. In: XXI International Con-gress of Entomology, Foz do Iguacu, Brazil. Ed byGazzoni, D. L. Londrina: Embraers, p. 524.

Krishna, K.; Araujo, R. L., 1968: A revision of theneotropical genus Neocapritermes (Isoptera, Termitidae,Nasutitermitinae). Bull. Am. Mus. Nat. Hist. 138,

84–138.Laffont, E. R.; Torales, G. J.; Arbino, M. O.; Godoy, M. C.;

Porcel, E. A.; Coronel, J. M., 1998: Termites asociadas aEucalyptus grandisW. Hill ex maiden en el Noroeste de laprovincia de Corrientes (Argentina). Rev. Agric. Piraci-caba 73, 201–214.

Lefeuve, P., 1990: A propos des termites de GuyaneFrancaise. Bois For. Trop. 224, 59–64.

Mariconi, F. A. M.; Franco, J. F.; Bleicher, R.; Domiciano,N. L., 1976: Combate aos ninhos do cupim Syntermes sp.com inseticidas. An. Soc. Ent. Brasil 5, 200–204.

Mill, A. E., 1991: Termites as structural pests in Amazonia,Brazil. Sociobiol. 19, 338–348.

Mill, A. E., 1992: Termites as agricultural pests in Amazo-nia, Brazil. Outlook Agric. 21, 41–46.

Novaretti, W. R. T.; Fontes, L. R., 1998: Cupins: uma graveameaca a cana-de-acucar no nordeste brasileiro. In:Cupins: O desafio do conhecimento. Ed. by Fontes, L.R.; Berti-Filho, E. Piracicaba: FEALQ, pp. 163–172.

Pohl, J. E., 1832: Reise Im Innern Von Brasilien, Vol. 1.Vienna.

Ripa, R.; Castro, L., 2000: Presencia de la termita subterra-nea Reticulitermes santonensis De Feytaud (Isoptera:Rhinotermitidae) en la comuna de Quillota. In: XXIIChilean Congress of Entomology. Ed by SociedadeChilean de Entornologia.Valdivia: Sociedade Chilena deEntomologia, pp. 82.

Santos, G. P.; Zanuncio, J. C.; Anjos, N.; Zanuncio, T. V.,1990: Danos em povoamentos de Eucalyptus grandis pelocupim-do-cerne Coptotermes testaceus Linnee, 1785(Isoptera: Rhinotermitidae). Revista Arvore 14, 155–163.

Scheffrahn, R. H.; Krecek, J., 1999: Termites of the genusCryptotermes Banks (Isoptera: Kalotermitidae) from theWest Indies. Insecta Mundi 13, 111–171.

Silva, A. G. A.; Goncalves, C. R.; Galvao, D. M.; Goncal-

ves, A. J. L.; Gomes, J.; Silva, M. N.; Simoni, L., 1968:Quarto catalogo dos insetos que vivem nas plantas doBrasil, seus parasitas e predadores. Rio de Janeiro:Ministerio da Agricultura.

Silvestri, F., 1903: Contribuzione alla conoscenza dei Termitie Termitofili dell’America Meridionale. Redia 1, 1–234.

Snyder, T. E., 1926: Termites collected on the MulfordBiological Exploration to the Amazon Basin 1921–22.Proc. U.S. Nat. Mus. 68, 1–76.

Snyder, T. E.; Zetek, J., 1946: The termite fauna of the CanalZone, Panama, and its economic significance. In: Termitesand Termite Control. Ed. by Kofoid, C. A. Berkeley, CA:University of California Press, pp. 342–346.

Thorne, B. L.; Haverty, M. I.; Collins, M. S., 1994:Taxonomy and biogeography of Nasutitermes acajutlaeand N. nigriceps in the Caribbean and Central America.Ann. Entomol. Soc. Am. 87, 762–770.

Torales, G. J., 1998: Rol de los isopteros en la Argentina. In:Cupins: O desafio do conhecimento. Ed. by Fontes, L. R.;Berti-Filho, E. Piracicaba: FEALQ, pp. 413–435.

Torales, G. J.; Laffont, E. R.; Arbino, M. O.; Godoy, M. C.,1997: Primera lista faunıstica de los isopteros de laArgentina. Rev. Soc. Entomol. Argent. 56, 47–53.

Valerio, J. R.; Santos, A. V.; Souza, A. P.; Maciel, C. A. M.;Oliveira, M. C. M., 1998: Controle quımico e mecanicode cupins de montıculo (Isoptera: Termitidae) em pasta-gens. An. Soc. Ent. Brasil 27, 125–131.

Weidner, H., 1980: Termiten aus Kolumbien nach Beobach-tungen von German O. Valenzuela und Fritz Schremmer.Anz. Schadlingskde., Pflanzenschutz, Umweltschutz 53,

65–69.Wessel, P.; Smith, W. H. F., 1998: New, improved version of

Generic Mapping Tools released. EOS Trans. Am.Geophys. U. 79, 579.

Wood, T. G., 1996: The agricultural importance of termites inthe tropics. Agr. Zool. Rev. 7, 117–155.

Author’s address: Reginaldo Constantino, Departamento deZoologia, Universidade de Brasılia, Brasılia, DF, 70910–900,Brazil. E-mail: constant@unb.br

Pest termites of South America 365