Beruflich Dokumente
Kultur Dokumente
c INRA/DIB-AGIB/ EDP Sciences, 2009 www.apidologie.org
DOI: 10.1051/apido/2009012
Review article
1
Universidade Federal do Cear, Departamento de Zootecnia CCA, Campus do Pici, CP 12168 Fortaleza,
CE 60.021-970, Brazil
2
Departamento de Ecologia, Instituto de Biocincias, Universidade de So Paulo 05508-900,
So Paulo, SP, Brazil
3
Departamento de Apicultura, Campus de Ciencias Biolgicas y Agropecuarias, Universidad Autnoma de
Yucatn. Apartado Postal 4-116 Mrida Yucatn CP 97100, Mexico
4
IMBIV (UNC - CONICET), Departamento de Diversidad Biologica y Ecologia, Facultad de Ciencias Exactas,
Fisicas y Naturales, Universidad Nacional de Cordoba, CC 495, 5000, Cordoba, Argentina
5
Departamento de Biologa, Facultad de Ciencias, Universidad Nacional de Colombia, Sede Bogot, Bogot,
D.C., Colombia
Abstract The Neotropics bee fauna is very rich with 5000 recognised species, including 33 genera (391
species) of Meliponini, but it is estimated to be at least three fold greater in species richness. Deforestation,
agriculture intensification and introduction/spread of exotic competing bee species are considered the main
threats to most indigenous species, although other less obvious causes can aect the populations of some
bee species locally. Eorts to conserve the native bee fauna include better knowledge of bee richness and
diversity (standardized surveys, larger bee collections and appropriate identification of bee species) and of
their population dynamics, raising of public and policy makers awareness, commercial applications of bee
products and services such as pollination and preservation of natural habitat.
1. BEE RICHNESS IN THE nomina nuda; and the existence of many ar-
NEOTROPICS eas/regions poorly sampled from where figures
on bee richness are certainly low.
It is well recognized that the Neotropics The recent publication of Moures cata-
bears a rich bee fauna (Moure et al., 2007). logue (Moure et al., 2007), the most com-
However, obtaining precise figures of this rich- prehensive work of the Neotropical Apoidea
ness is not easy due to a series of reasons such so far, has compiled most information avail-
as the spread of this information in thousands able and presented reliable data on numbers
of papers, books, dissertations and thesis; the and the geographic distribution of bee species.
lack of accurate information on valid names, The catalogue presents 5000 valid bee species
synonymy, homonymy, misidentifications and names but estimates an astonishing total num-
ber of 15 150 bee species in the Neotropics,
Corresponding author: B.M. Freitas, implying that all data available up to date on
freitas@ufc.br bee diversity in the region represents less than
* Manuscript editor: Mark Brown a third of actual species richness (Tab. I).
Table I. Number of Neotropical bee species by country. Numbers in brackets are the estimated total number
of species using the equation: y = 0.2889x + 1.2108; r2 = 0.984, where y = estimated total number of
species.
1.1. Mexico and Central America et al.s (2007) table on the number of Neotrop-
ical bee species by country, including also the
The apifauna of Mexico and Panama is estimates based on Ayala et al. (1993), Maes
probably the best studied of the countries that (1993) and Ayala (2006) of the whole of Mex-
form the Mesoamerica region. Unfortunately, ico (Tab. I).
data on the bee fauna in other countries of
Central America are scarce, and sampling ef-
forts have only recently started in Guatemala 1.2. South America
(Enriquez et al., 2003).
Mexico is considered to have intermediate Like in many other parts of the world, the
bee richness between that of the United States apifauna of South America has been under-
(ca. 4000) and those in countries from Cen- studied. There have been few surveys in most
tral America (ca. 600) probably as a result countries and where they do exist, such as
of its position in which elements of Neartic, in Brazil, Argentina and Colombia, they are
Mesoamerican and Neotropical origins are not evenly distributed, tending to concentrate
combined (Tab. I; Ayala et al., 1993). Within in some regions and leaving others uninves-
Mxico, there are striking dierences; the arid tigated. In Brazil, for example, we found 84
and semiarid regions of the north represent surveys carried out in 130 dierent localities,
some of the richest bee faunas in the world but 16 out of them were in 25 sites only in
whilst the Yucatan Peninsula and the high- the state of So Paulo. In contrast, the North
lands above 3000 m seem to be relatively region of Brazil remains unsampled and few
species poor (Ayala et al., 1993). However, surveys have been made in the Central-West
these patterns have been considered tenta- region. These two regions together cover ap-
tive since most regions apart from the north- proximately 4 700 000 km2 , representing more
ern border of Mexico with the United States than 50% of the Brazilian territory and include
have been scarcely surveyed, so the number important and extensive biomes such as the
of species may well be above 2000. For in- Amazon rainforest and the tropical wetland of
stance, in the Yucatan Peninsula, a recent study the Pantanal Matogrossense. Presently, Brazil
in the biosphere of Sian Kaan, in the state of has 1678 valid species names for bees (Moure
Quintana Roo has revealed the presence of 74 et al., 2007).
bee species (Roubik et al., 1991) and a sur- Besides uneven sampling, the lack of a
vey across the state of Yucatan reported 140 standardized methodology and type of data
species and the first report of Colletidae (50% collected are important constraints to a com-
more species than were reported in previous parison among surveys, but Silveira et al.
surveys; Contreras-Acosta, 1998). (2002) point out that the appropriate identi-
The most comprehensive reports of the bee fication of bee species is the major limita-
richness of Mexico and Central America are tion. This is why the great majority of studies
those of Ayala et al. (1993), Maes (1993) and have solely undertaken general comparisons
Ayala (2006). From these studies, the estima- based on species richness of Apoidea fami-
tions for the bee fauna of Mexico are of 8 fam- lies found within distinct areas of the same
ilies, 144 genera and ca. 1800 species. Four biome (e.g. Silveira and Campos, 1995) or
bee genera and eight subgenera are endemic. among biomes within the same geographic re-
The greatest number of species has been found gion (e.g. Jamhour and Laroca, 2004), mak-
in the following genera: Centris, Deltoptila, ing clear the urgent need of a synthesis of
Exomalopsis, Mesoxaca, Peponapis and Pro- this accumulated knowledge. Some attempts
toxea. Yet Moures catalogues (Moure et al., have been made by Pinheiro-Machado et al.
2007) did not include bee species in Northern (2002), for instance, who reviewed 46 surveys
states of Mexico, considering them as belong- made in Brazil, presenting information related
ing to the Neartic fauna. From this point of only to species richness of Apoidea families
view, Mexico would have 916 bee species of in dierent biomes. Biesmeijer et al. (2005)
the Neotropical fauna. Here, we present Moure synthesized data from eusocial Apidae from
4 B.M. Freitas et al.
27 surveys, making a presence/absence ma- and agriculture expansion are the main causes
trix to analyze interactions patterns and the of deforestation. Removal of native vegeta-
influence of factors such as altitude, latitude tion aects bee populations basically through
and habitat. Biesmeijer and Slaa (2006) also the loss/change of floral resources and nesting
compared data from 28 Brazilian bee surveys sites (Kremen et al., 2004).
in order to explain patterns of species rich- The Amazon rainforest is the largest ex-
ness, niche width and association among taxa panse of forest in the world (5.5 million
of eusocial Apidae. These diculties bring km2 ) and is located to the north of the South
much discrepancy: in Colombia Nates-Parra American continent, being shared by 9 coun-
(2005) reports approximately 600 identified tries: Brazil, Bolivia, Ecuador, Peru, Colom-
bee species to the country, but Moures cata- bia, Venezuela, Suriname, French Guiana and
logue (Moure et al., 2007) recognizes only 398 Guiana. Presently, the Amazon rainforest suf-
valid names, although estimates around 900 fers the highest rates of deforestation in the
species in Colombia. Americas. The largest part of it is located in
The Meliponini tribe (Hymenoptera: Ap- Brazil (60%) and covers almost half of that
idae), eusocial bees native to the New country. Brazil is also there where deforesta-
World, exhibit remarkable bee richness in tion has been most intense; 756 km2 of for-
the Neotropics. They form perennial colonies, est were cleared in August 2008 alone, an area
which inhabit most of the tropical Ameri- much larger than Singapore (641 km2 ), for
cas, ranging from the lowlands of Mexico to example. From January to August 2008 de-
the northwest of Argentina and Montevideo, forestation in Brazil amounted to 5.681 km2
Uruguay, including some Caribbean islands, (INPE, 2008). According to the Brazilian Na-
from sea level to altitudes up to 4000 m in the tional Institute of Space Research (INPE) and
Bolivian Andes (Camargo and Pedro, 2007). the World Wildlife Fund (WWF), 17.1% of the
According to Camargo and Pedro (2007), there Brazilian Amazon rainforest had been cleared
are 33 exclusively Neotropical valid genera to date (WWF, 2008). Fearnside (2008) has
of Meliponini, including one recently extinct identified the reasons for the intense deforesta-
species and a total of 391 valid species names. tion of Amazonia. Wood extraction, cattle pas-
ture and, more recently, expansion of soybean
plantations towards the North of the Brazil
2. THREATS TO BEES IN THE are the main causes of deforestation. In other
NEOTROPICS South American countries, clearing of land for
agriculture is more prevalent (WWF, 2008).
There are many threats to native bees in the Like in the Amazon rainforest, the cer-
Neotropics, all directly or indirectly related to rado vegetation of Central Brazil, which in-
human activities such as habitat fragmentation cludes the tropical wetland of the Pantanal
and habitat loss, unsustainable honey hunting, Matogrossense, is being rapidly replaced by
biological invasions, and intensive use of her- grassland and soybean plantations. Recently
bicides and pesticides. However, these threats Morton et al. (2006) and Souza (2006) used
can be pooled into three major categories: de- satellite images to demonstrate fast cerrado de-
forestation, agriculture intensification and in- forestation and the land use after deforesta-
troduction/spread of exotic species tion. The eects on the bee fauna both in the
Amazon rainforest as well in the cerrado in a
short span of time can only be roughly esti-
2.1. Deforestation mated, but predictions for the future are poten-
tially catastrophic because they cover large ar-
Neotropical ecosystems along with those eas, including complete river drainage systems
from Southeast Asia (Oldroyd and Nanork, and impact global carbon and water cycles,
2009) are among the most strongly impacted energy balance and climate (Goudie, 2001;
by deforestation. Logging, firewood gathering, WWF, 2008). These are the regions least sam-
charcoal production, clearing for cattle pasture pled in Brazilian bee collections and many bee
Bee status in the Neotropics 5
Table II. Average amount of forest reduction per year in Mexico and Central America (FAO, 2003).
species are probably still to be discovered and of Hymenoptera in fragmented and continuous
described. There is the imminent risk of losing forest for two consecutive years (Galetto et al.,
bee species before their discovery. 2007). A common pattern for these two Chaco
In the Americas, the regions with the sec- regions was that feral honeybees (Apis mellif-
ond highest tropical deforestation rates are era) were more frequent on small fragments
Mexico and Central America which altogether (Aizen and Feinsinger, 1994; Galetto et al., un-
lost an average of 1.3% (285 000 hectares) of publ. data).
its rainforests each year between 2000 and In Colombia, a red book of terrestrial in-
2003 (FAO, 2003). However, rates of forest vertebrates (Amat et al., 2007) mentions 10
lost are substantially dierent between coun- bee species as threatened, including species of
tries (see Tab. II). Meliponini and the genus Aglae, Eufrisea and
In Mexico for instance, rapid industrializa- Bombus. Habitat loss and fragmentation due
tion and uncontrolled population growth over to deforestation are the main threats to native
the last few decades have had a strong im- bees from the Pacific coast to the Andes, the
pact on the countrys environment and left Colombian Amazon rainforest and the Caribe
less than 10 percent of its original tropi- (Gonzles and Nates-Parra, 1999; Nates-Parra,
cal rainforests standing. Today Mexicos rain- 2005; Bonilla-Gmez, 2007). But there are
forests are limited to southeastern regions other causes of concern, such as unsustainable
along the Gulf of Mexico and the southern honey hunting in the Andes, Amazon, Ori-
state of Chiapas. These forests are most threat- noquia and Caribe that are aecting popula-
ened by subsistence activities, especially fuel tions of the stingless bees Melipona favosa and
wood collection, land clearance for agricul- Melipona eburnea (Nates-Parra, 2007) and,
ture and livestock, illegal logging and poach- particularly, Parapartamona imberbis which
ing (Gomez-Pompa and Kaus, 1999). is endemic to high mountains of the Colom-
During the last 30 years, great pressure on bian Andes that are now being converted for
natural woodlands has occurred in central and potato crops and cattle breeding areas. An-
northern Argentina, mainly due to the expan- other four Parapartamona species, P. brevip-
sion of crop fields for soybean, leading to high ilosa, P. caliensis, P. vittigera and P. zonata
rates of deforestation (Zak et al., 2004; Grau are also endangered, although their distribu-
et al., 2005). The Chaco is the second largest tion also covers the Ecuadorian Andes (Nates-
dry forest biome in South America and a sig- Parra, 2007).
nificant proportion of it was aected by de-
forestation. For this region, similar declines in
abundance and diversity of solitary and social 2.2. Agriculture intensification
bees were registered in fragmented and non-
fragmented landscapes of Tucumn in NW Ar- Agriculture expansion and its intensifi-
gentina (Aizen and Feinsinger, 1994, 2003). A cation throughout the Americas have been
dierent pattern was observed in Chaco wood- pointed out as a major threat to bees (Kremen
lands of central Argentina, because no signifi- et al., 2002; Silveira, 2004). Besides being
cant dierences were registered in the richness one of the three main causes of deforestation
6 B.M. Freitas et al.
along with logging and clearance for cattle In Argentina, Chaco and Aizen (2006) deter-
pasture, large-scale arable agriculture also re- mined whether forest patches of Yungas (NW
duces plant and animal diversity dramatically, Argentina) act as a source of pollinators for
diminishing bee nesting and feeding opportu- grapefruit (Citrus paradisi); they registered a
nities and kills adult and larval bees by using total of 50 insect species on grapefruit flowers,
agrochemicals and by ploughing the soil. including 22 (44%) species from six families
The density of stingless bee nests in the of bees (Apidae, Andrenidae, Anthophoridae,
southern state of Yucatan in Mexico was sig- Chrysididae, Halictidae and Megachilidae).
nificantly higher in undisturbed areas com- The feral Africanized honeybee A. mellifera
pared to those where agriculture and ranch- contributed more than 90% of the visits to
ing have decimated the forests (Santos-Leal, grapefruit flowers, representing the main pol-
2006). In coconut plantations (Cocos nucifera) linator, probably because of its capability to
where the terrain is cleared of other plants, persist in agricultural landscapes (Chaco and
it has been reported that bee richness is low Aizen, 2006). Interestedly, the frequency of
apparently due to reduced nest site availabil- visits to grapefruit flowers decreased by more
ity and foraging resources limited to only than twofold as distance to the forest in-
this plant species (Melndez et al., 2004). creased and the flower-visiting fauna became
In Rio Grande do Sul, the extreme south of reduced. Chaco and Aizen (2006) concluded
Brazil, Melipona bicolor schencki is threat- that negative forest edge eects on flower-
ened (Blochtein and Marques, 2003), mainly visiting insects inside grapefruit plantations
due to the intense deforestation and increas- are widespread in the increasingly deforested
ing plantation of Pinus for paper production. landscape of NW Argentina.
These bees need large trees for nesting (Witter Two studies of passion fruit (Passiflora
et al., 2008) and an undisturbed landscape. edulis cf. flavicarpa) pollination carried out
Meanwhile, in Cear, the Northeastern part of in NE (Freitas and Oliveira-Filho, 2001) and
Brazil, Melipona quinquefasciata, a ground- SE (Camillo, 1996) Brazil found increments
nesting stingless bee, is endangered due to from 92 to 700% after the introduction of 25
firewood gathering and agricultural expansion Xylocopa frontalis nest/ha. Comparisons be-
which destroy the ecosystem needed for them tween these two studies made by Freitas and
to nest as well as their floral host plants (Lima- Oliveira-Filho (2003) suggest that the num-
Verde and Freitas, 2002; Eardley et al., 2006). ber of wild X. frontalis in the southern area
There are no precise or reliable figures of was much lower than that of the northern area,
how much land has been converted annually mainly due to the lack of appropriate nest
into agriculture. In Brazil, for instance, an in- sites in the highly agricultural landscape of SE
crement of between 1.2 to 2.7% in cropped Brazil.
area to 2009 is expected, adding an additional Intensive agriculture and ranching im-
544 to 1276 thousand hectares to the 47.31 plies a collateral use of agrochemicals to
million hectares cultivated in 2008 (Conab, reach desirable pH and amounts of nutri-
2008). However, it is not possible to estimate ent in the soil as well as the use of chem-
what percentage of this area will be from new icals for pest and disease control leading to
clearings. soil degradation and accumulation of harm-
Ironically, most plant species cultivated by ful levels of toxic substances which may af-
man are dependent to some extent on biotic fect the survival of bees (Ramalho et al.,
pollinators, from which bees are the most im- 2000; Caldas and Sousa, 2000). Pesticides
portant group (Klein et al., 2007). Many crops are widely used in the Neotropics. In Mex-
could be susceptible to declines in pollinator ico, Central America and some countries of
abundance and diversity of wild pollinators South America, strict control on the use of
and increasingly frequent collapses of man- many chemicals, including several banned in
aged honey bee (Apis mellifera) populations Europe and the United States, is lacking.
(Aizen et al., 2008b). Nevertheless, few em- Agriculture in Latin America relies heavily on
pirical data are available for the Neotropics. pesticide imports and countries like Mexico,
Bee status in the Neotropics 7
Argentina and Brazil are large pesticide im- 2003). Two separate surveys of bee species
porters (Tansey, 1995; Pinheiro and Freitas, visiting cultivars of Cucumis moschata and
unpubl. data). Some pesticides banned some- Capsicum chinense in Yucatn showed that
where else are still permitted in Mexico, for the use of pesticides reduced the number of
example, DDT, aldrin, clordane, and cloroben- species and the total numbers of individuals af-
zolate (Alvarado-Mejia et al., 1994). The sit- ter spraying with malathion and diazinon (Xiu
uation in Central America is similar. In Costa Ancona, 2007).
Rica and Nicaragua, extensive use of very haz-
ardous pesticides continues and so does poi-
soning of bees with organophosphates, car- 2.3. Exotic species
bamates, endosulfan and paraquat. However,
in 2000, the ministers of health of the seven Deliberate or accidental introduction of ex-
Central American countries agreed to ban or otic bee species or other organisms that might
restrict twelve pesticides (Wesseling et al., interact directly or indirectly with local popu-
2005). In Brazil, there are restrictions and reg- lation can be hazardous to a native bee fauna
ulations to the use of agrochemicals, but little (reviewed in Stout and Morales, 2009).
enforcement. The classic example of an exotic animal in-
Insecticides are potentially the most dam- troduced to the Americas is the case of Apis
aging agrochemicals for bees (De La Ra mellifera. European races of this species, na-
et al., 2009). Although their immediate lethal tive to the Old World, have been brought to
eects are usually pointed out as the major various countries of the Americas by European
problem to bees, in many cases sub-lethal ef- settlers since the XVI century. In 1956, A. mel-
fects are more dangerous. They do not cause a lifera scutellata, an African race, was intro-
prompt death of the bee or (for social species) duced to Brazil and spread wildly throughout
colony but aect their sensory and neuromo- the continent from Argentina to the USA, ex-
tor skills. Pesticides may be applied directly or cept Chile, in what has been called African-
consumed in food, disrupting sensory, naviga- ization (Seeley, 1986; Winston, 1987).
tional and kin recognition abilities. Sublethal The introduction of A. mellifera to the New
eects can be dicult to notice and impact World has had a positive social impact through
more on bee diversity or population levels than honey hunting and beekeeping; Mxico, Ar-
easily detectable lethal eects (Freitas and gentina and Brazil are ranked among the great-
Pinheiro, unpubl. data). For example, many in- est honey producers in the world (Freitas et al.,
secticides are absorbed by pollen grain lipids 2007). In Mexico, the worlds third largest
and can intoxicate brood of solitary bees or honey producer, Apis mellfera colonies reach
cause brood and young bee mortality for a some of the highest densities anywhere in the
long time in colonies of social bees (Loper and world, in particular in the tropical areas of the
Ross, 1982). Gulf Coast and the Yucatan Peninsula. More-
The eect of pesticides on bee popula- over, since the 1980s, the numbers of honey
tions has not been documented in South Amer- bee colonies in tropical Mexico and Central
ica, but in Mexico and Central America some America have probably increased at least two
social bees are currently under threat such fold due to the colonization of feral colonies
as Melipona beecheii (Kerr, 2002) and Bom- of Africanized bees (Roubik, 1989; Quezada-
bus medius (Gonzlez-Acereto, unpubl. data). Eun and May-Itza, 2001). Similar patterns are
Although several poisoning incidents are re- also observed in parts of South America. Also,
ported in colonies of A. mellifera annually, Apis mellifera may have contributed as pol-
no direct link with the use of pesticides has linator to increase some production of some
been established. However, recently a study crops in the Americas, as shown by Freitas and
in Yucatan found that permethrine at doses Paxton (1998) and Roubik (2002) to cashew
used for control of mosquitoes during the rainy and coee, respectively.
season is highly poisonous for honey bees Large populations of A. mellifera may also
and stingless bees (Valdovinos-Nez et al., be socially impacting stingless beekeeping. In
8 B.M. Freitas et al.
NE Brazil reductions in the number of sting- of bumble bees that had been used for pol-
less bee beekeepers and colonies in the 1970s lination are non-native to Mexico and Cen-
and 1980s have been attributed to honeybees. tral America: Bombus impatiens and Bombus
The decrease of stingless beekeeping with terrestris (Winter et al., 2006; Palma et al.,
Melipona beecheii in the Yucatan peninsula 2008a, b). They began to be imported in 1994
has also been attributed to Africanized bees, (Velthuis and Van Doorn, 2006) but shipments
although no negative influence is reported by of European B. terrestris were banned into
stingless beekeepers of Veracruz along the Mexico between 1995 and 1996 due to par-
Mexican Gulf Coast who cultivate Scaptotrig- asite infections (Winter et al., 2006). How-
ona pectoralis (Quezada-Eun, 2005). ever, colonies of North American B. impatiens
The eect of such numbers of honey bees are still allowed into Mexico and Guatemala
on the native apifauna and flora is not known. (Morales, 2007). Therefore, there is a po-
There has been much discussion of the po- tential risk of naturalization of exotic bum-
tential negative eects caused by A. mellifera ble bees, as has already occurred in other
on local populations of native bee species by countries worldwide (Colla et al., 2006). This
competition for floral resources and by occu- is a serious threat to native bumble bees
pation of nesting sites of highly social sting- that can be infected with diseases and par-
less bees by honey bees (Roubik, 1989; Freitas asites and hybridized to an unknown extent
et al., 2007; reviewed in Stout and Morales, with their foreign counterparts (Cuadriello-
2009). Aguilar and Salinas-Navarrete, 2006; Velthuis
Monitoring pollinator visitation in French and van Doorn, 2006; reviewed in Stout and
Guiana over a period of 17 years, at a site Morales, 2009).
where urban expansion brought some distur- In temperate forests of the southern An-
bance, Roubik (1996) pointed out a 100% in- des of Argentina, the native Bombus dahlbomii
crease in the proportion of honeybees visits to declined while foragers of the exotic Euro-
flowers, whereas native bees visits per flower pean Bombus ruderatus (introduced via New
diminished proportionally to honeybees domi- Zealand) increased along a gradient of anthro-
nance in forest plots. pogenic habitat alteration (Morales and Aizen,
Roubik (2000) also noted that a generalist 2002; Aizen and Feinsinger, 2003; Morales,
species as the Africanized honeybee, which 2007). A second invader, Bombus terrestris,
demands huge resources supply will not al- was recently registered for this biogeographic
ways invade all types of ecosystem. He did not region, probably coming from Chile where
find any Africanized honeybee in a preserved it was introduced for pollination purposes
area of lowland forest in Ecuador, meaning (Torreta et al., 2006). There are concerns of
that dierent communities are not equally vul- its spread towards Uruguay, Paraguay, Bo-
nerable to invasion. livia and southern Brazil. Exotic pollinators
Negative interactions of A. mellifera with can modify basic quantitative parameters of
solitary bees are less documented but do also plant-pollinator webs, such as the strength
exist (Murray et al., 2009; Stout and Morales, of interactions, influencing the basic founda-
2009), like the displacement of solitary bees tions of the architecture of mutualistic net-
such as Peponapis limitaris and Partamona bi- works (Aizen et al., 2008a) with putative
lineata from floral resources (Pinkus-Rendon consequences for the preservation of plant and
et al., 2005) and honey bees removing all animal populations (Stout and Morales, 2009).
pollen available in passion fruit flowers, pre-
venting pollen harvesting by Xylocopa bees
(Freitas and Oliveira-Filho, 2001). 2.4. Other threats
Apart from honey bees, exotic bumble-
bees seem to be the major concern nowa- Heavy metals are chemical elements of rel-
days in the Neotropics. Pollination in green- ative density higher than 5 g/cm3 found in high
houses has considerably increased in Mexico concentrations in waste areas (biosolids and
(Palma et al., 2008a,b), but the two species mining) or places of intense use (industrial
Bee status in the Neotropics 9
weisen die hchsten Entwaldungsraten in Ameri- siderurgia como fonte de micronutrientes e de con-
ka auf, aber auch in den Chaco-Wlder der ar- taminante para plantas de milho, Pesq. Agropec.
gentinischen und kolumbianischen Anden schrei- Bras. 35, 14831491.
tet die Entwaldung fort (Tab. II). Die Ausbreitung Aizen M.A., Feisinger P. (1994) Habitat fragmenta-
und Intensivierung der Landwirtschaft wird viel- tion, native insect pollinators, and feral honeybees
fach als die wichtigste Bedrohung fr Bienenar- in Argentine chaco serrano, Ecol. Appl. 4, 378
ten angesehen. Sie fhrt zu einer Verringerung der 392.
Artendiversitt bei Tieren und Pflanzen, verringert
Aizen M.A., Feisinger P. (2003) Bees not to be?
das Angebot an Nistmglichkeiten und Futterquel-
Responses of insect pollinator faunas and flower
len, und durch das Ausbringen von Pestiziden und pollination to habitat fragmentation, in: Bradshaw
Pflgen der Bden werden sowohl Bienenlarven als G.A., Marquet P.A. (Eds.), How landscapes
auch Adulte gettet. Ironischerweise sind die mei- change: Human disturbance and ecosystem dis-
sten Kulturpflanzen mehr oder weniger stark auf ruptions in the Americas, Springer-Verlag. Berlin,
die Prsenz biotischer Bestuber angewiesen, wobei pp. 111129.
die Bienen die wichtigste Gruppe darstellen. Auch
die Einfhrung fremder Bienenarten und anderer Aizen M.A., Morales C.L., Morales J.M. (2008a)
exotischer Organismen, die mit der lokalen Bie- Invasive mutualists erode native pollination webs,
nenfauna in Wechselwirkung treten, kann die ein- PLOS Biol. 6, 396403.
heimische Bienenfauna beeintrchtigen. Die Ein- Aizen M.A., Garibaldi L.A., Cunningham S.A., Klein
fhrung von Apis mellifera in die Neue Welt zur A.M. (2008b) Long-term global trends in crop
Steigerung der Honigproduktion und die von exo- yield and production reveal no current pollination
tischen Hummelarten fr Bestubungszwecke hat shortage but increasing pollinator dependency,
zu Bedenken Anlass gegeben ber die Konkurrenz Curr. Biol. 18, 15721575.
mit einheimischen Bienen um Futter- und Nistmg- Alvarado-Mejia J., Cobos-Gasca V., Gonzalez-
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