Bee Faunas (Hymenoptera: Apoidea) of Six Natural

Protected Areas in Yucatan, Mexico

Author(s): Enrique Reyes-Novelo, Virginia Mlndez-Ramrez,
Ricardo Ayala, and Hugo Delfn-Gonzlez
Source: Entomological News, 120(5):530-544. 2009.
Published By: The American Entomological Society
DOI: http://dx.doi.org/10.3157/021.120.0510
URL: http://www.bioone.org/doi/full/10.3157/021.120.0510

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530

ENTOMOLOGICAL NEWS

BEE FAUNAS (HYMENOPTERA: APOIDEA) OF SIX

NATURAL PROTECTED AREAS IN YUCATAN, MEXICO1
Enrique Reyes-Novelo,2 Virginia Mlndez-Ramrez,2 Ricardo Ayala,3
and Hugo Delfn-Gonzlez2
ABSTRACT: The wild bee fauna of six natural protected areas (NPAs) in Yucatan State, Mexico, are
described and analyzed. A total of 130 bee species was recorded belonging to 50 genera and five families, of which 32 are new records for Yucatan State. The genera with the highest species richness
were Megachile (20 species), Lasioglossum (12 species), Coelioxys (11 species, most of these are
cleptoparasites of Megachile), Ceratina (9 species) and Augochlora (8 species). Species richness by
NPA was 30 species in El Palmar; 55 in Dzibilchaltn; 58 in Kabah; 60 in Dzilam; 69 in Tabi; and
79 in Yalahau. Bee faunal composition in the NPAs, biogeographic affinities and the importance of
wild bee conservation in the region are discussed.
KEY WORDS: Bees, Apoidea, diversity, Hymenoptera, Yucatan

Wild bees require floral resources (e.g. pollen, nectar, oils and resins) for their
proper development and reproduction. Bees are also a key element in terrestrial
ecosystems because they provide the ecological service of pollination by visiting
the flowers of cultivated and wild plants, thus playing a vital role in agricultural
production and the conservation of vegetal communities. At least 67% of flowering plants require pollinators, mainly bees (Kevan, 1999; Cane and Tepedino,
2001; Williams et al., 2001; Potts et al., 2005; Michener, 2007).
Their importance to ecosystem maintenance makes it vital that wild bee diversity be researched and conserved, primarily in natural protected areas (NPAs),
where scarce diversity data has been generated to date. Mexico contains 161
NPAs (accounting for approx. 11.56% of the country; CONANP, 2007), but bee
species have been researched in only four of them: Chamela-Cuixmala Biosphere Reserve in Jalisco state (Ayala, 2004); Mapimi Biosphere Reserve in Durango state (Lopez-Mendoza, 2003); Sian Kaan Biosphere Reserve in Quintana
Roo state (Roubik et al., 1991; Cairns et al., 2005); and Ria Lagartos Biosphere
Reserve in Yucatan state (Novelo-Rincon et al., 2003).
Over 19,500 described bee species are recognized worldwide (Ascher and
Pickering 2009), of which nearly ten percent are recorded in Mexico (1,767
species in 144 genera; Moure et al., 2007; Ascher and Pickering 2009). Bee faunal studies done in Mexico to date include Godinez-Garcia (1991) in Guanajuato
state; those of Hinojosa-Diaz (1996 and 2003) in Mexico City and Morelos state;
Vergara and Ayala (2002) in Puebla state; Lopez-Mendoza (2003) in Durango
state; Novelo-Rincon et al. (2003) in Yucatan state; and Godinez-Garcia et al.
______________________________
1

Received on September 23, 2008. Accepted on November 9, 2008.

Departamento de Zoologa, Campus de Ciencias Biolgicas y Agropecuarias, Universidad Autonma de Yucatan. Apartado Postal 4-116 Colonia Itzimna 97100 Mrida, Yucatan, Mxico. E-mails:
(ER-N) enrique.reyes@uady.mx (VM-R) virmelen@uady.mx (HD-G) gdelfin@uady.mx

Estacin de Biologa Chamela, Instituto de Biologa, Universidad Nacional Autonma de Mxico.

Apartado Postal 21,48980 San Patricio, Jalisco, Mxico. E-mail: barajas@servidor.unam.mx.
Mailed on December 13, 2010

Volume 120, Number 5, November and December 2009

531

(2004) in the Sierra Madre Oriental. The most thoroughly studied state in the
country is Jalisco (Ayala, 1988; 2004; Estrada, 1992; Fierros-Lopez, 1996).
Many of the species reported in these studies are new to science or morphospecies belonging to taxonomically unrevised genera, meaning they are unrecognized in the Mexican fauna. Clearly, further research is needed to adequately
describe the diversity of Apoidea in Mexico.
As part of a broad bibliographic revision, Ayala et al. (1996) cited 67 species
in Yucatan, and later published a revision of the Meliponini in Mexico, adding
three new species (Ayala, 1999). The most extensive list of bee species collected in the region was published by Novelo-Rincon et al. (2003), who reported 140
species. They only identified 65 to the species level, but these included 37 new
species records for the state, thus increasing the total number of identified
species to 107.
The objective of the present study was to characterize the apifauna in six natural protected areas (NPAs) in Yucatan State. This will serve as a contribution to
the knowledge of wild bees in southeast Mexico and to generate species richness
data for these NPAs that will serve as a baseline for future ecological and pollination studies, and to promote bee conservation and management in NPAs.
METHODS
An examination was made of wild bee specimens on deposit in the Regional
Entomological Collection of the Universidad Autonoma de Yucatan (CERUADY) collected from six NPAs: Dzibilchaltn National Park; Dzilam State
Reserve; El Palmar State Reserve; Kabah State Park; Lagunas de Yalahau State
Park; San Juan Bautista Tabi Scenic, Historic, Cultural and Natural Protected
Area and its Sacnicte Annex (Table 1, Figure 1). Specimen collection was done
as part of faunal diversity research in the NPAs of Yucatan during 2005 and 2006.
Collections were made using entomological aerial nets, Malaise traps, yellow
containers and McPhail traps baited with honey diluted to 50%.
Genus-level determination of specimens was made based on Micheners
(2007) keys, and species-level identification was done by comparison with bee
collections at the Chamela Biological Station, UNAM Institute of Biology, Jalisco, Mexico (IBUNAM-Chamela) and the American Museum of Natural History
(New York, USA) (AMNH) (with the support of Dr. John Ascher), review of bibliographic material (published and unpublished catalogs from AMNH) and consultation with experts in specific groups (see Acknowledgements). The Meliponinae classification was according to Camargo and Pedro (2008).

532

ENTOMOLOGICAL NEWS

Table 1. Basic information on the natural protected areas (PAs) in Yucatan, Mexico.

Protected area

Latitude/Longitude

Surface (ha)

Veg

Climate

Dzibilchaltn

2104'26", 2106'00" N;
8934'51", 8936'50" W

539.43

TDF

AW0

2040'37", 2034'59" N;
8910'49", 8915'00" W

5683.28

TDF

AW0

2126'33", 2124'51" N;
8847'54", 8847'49" W

61706.83

SFF

BS1

2055'00", 2111'00" N;
9000'00", 9022'30" W

50177.39

CDV

BS0

2014'12", 2013'22" N;
8939'34", 8938'03" W

949.76

SDTF

AW0

2014'47", 2014'59" N;
8930'26", 8932'29" W

1355.74

SDTF

AW0

Yalahau
Dzilam
El Palmar
Kabah
Tabi

Veg = Vegetation types, TDF = Tropical Deciduous Forest, SFF = Seasonally Flooded
Forest, CDV = Coastal Dune Vegetation; SDTF = Semi-Deciduous Tropical Forest; AW0
= Warm subhumid with lowest humidity percentage; BS1 = Dry with medium humidity
percentage; BS0 = Dry with lowest humidity percentage.

Figure 1. Location of the six Natural Protected Areas from Yucatan, Mexico.

Volume 120, Number 5, November and December 2009

533

Species distributions and biogeographical affinities were determined based on

data proposed by Ayala et al. (1996), Michener (1979 and 2007), Ascher et al.
(2008) and AMNH literature and specimen records of species new to Yucatan.
Grouping by lifestyles was done according to the four categories proposed by
Michener (1974) and Roubik (1989): eusocial; solitary; cleptoparasitic; and
parasocial (i.e. all species exhibiting different social levels, such as communal,
quasisocial and semisocial). Nesting sites were assigned according to the criteria
of Michener (2007): Cavity for bees that use preexisting holes in different substrates as rocks, soil banks, tree branches, etc., ground for excavating their nests
in the soil, and finally wood for the ones that excavate soft or hard wood for their
nests. The lifestyle and nesting sites of many species are still unknown, and in
some cases they were inferred based on data for taxonomically related species
(except for most of the morphospecies). The ecological aspects of the data presented here will be analyzed elsewhere.
RESULTS
A total of 6,785 specimens was examined from five families, 50 genera and
130 species, of which 32 were new records for the region (Table 2). These represent 34% of the genera and 7.4% of the bee fauna recorded in Mexico. Species
richness by NPA was 30 species in El Palmar; 55 in Dzibilchaltn; 58 in Kabah;
60 in Dzilam; 69 in Tabi; and 79 in Yalahau. The richest family (in terms of genera and species) in the six studied NPAs was the Apidae, followed by Megachilidae, Halictidae, Colletidae and Andrenidae (Fig. 2).
Table 2. Bee species of six protected areas in Yucatan. LS = Lifestyle: PS =
Parasocial, SO = Solitary, EU = Eusocial, CL = Cleptoparasite, UK= Unknown.
Nest: S = Soil, C = Preexisting cavity, W = Wood. S1 = Dzibilchaltun, S2 =
Dzilam, S3 = Kabah, S4 = El Palmar, S5= Tabi, S6 = Yalahau. TSE = Total of
specimens examined, * New records for Yucatan.
Superfamily Apoidea: Clade Anthophila (Bees) LS Nest
Family Colletidae
Subfamily Colletinae
Colletes arizonensis Stephen, 1954
Colletes punctipennis maurus
Stephen, 1954
Subfamily Diphaglossinae
Tribe Caupolicanini
Ptiloglossa eximia (Smith, 1861)
Subfamily Hylaeinae
Hylaeus sp1
Hylaeus sp2
Hylaeus sp3
Hylaeus sp4
Hylaeus sp5
Family Andrenidae
Subfamily Panurginae
Calliopsis hondurasica Cockerell, 1949
Pseudopanurgus crenulatus

S1

S2

S3

S4

S5 S6 TSE

SO

SO

SO

SO
SO
SO
SO
SO

C
C
C
C
C

X
-

X
-

X
X
-

X
X
-

3
2
1
1
1

SO

534

ENTOMOLOGICAL NEWS

Superfamily Apoidea: Clade Anthophila (Bees)

(Cockerell, 1905)
Family Halictidae
Subfamily Halictinae
Tribe Augochlorini
Augochlora (Augochlora) albiceps
Friese, 1925
Augochlora (Augochlora) nigrocyanea
Cockerell, 1897
Augochlora (Augochlora) smaragdina
Friese, 1917
Augochlora (Oxystoglosella) aurifera
Cockerell, 1897
Augochlora (Oxystoglosella)
cordiaefloris Cockerell, 1907
Augochlora sp1
Augochlora sp2
Augochlora sp3
Augochlorella bracteata Ordway, 1966
Augochlorella neglectula (Cockerell,
1897)
Augochlorella pomoniella (Cockerell,
1915)
Augochloropsis (Paraugochloropsis)
metallica (Fabricius, 1793)
Augochloropsis sp1
Augochloropsis sp2
Augochloropsis sp3
Caenaugochlora (Caenaugochlora)
gemmella (Cockerell, 1912)
Pseudaugochlora graminea
(Fabricius, 1804)
Caenohalictus amatitlana (Cockerell,
1912)
Temnosoma smaragdinum Smith, 1879
Tribe Halictini
Agapostemon (Notagapostemon) nasutus
Smith, 1853
Halictus (Odontalictus) ligatus
Say, 1837
Halictus (Seladonia) hesperus
Smith, 1862
Lasioglossum (Dialictus) sp1
Lasioglossum (Dialictus) sp2
Lasioglossum (Dialictus) sp3
Lasioglossum (Dialictus) sp4
Lasioglossum (Dialictus) sp5
Lasioglossum (Dialictus) sp6
Lasioglossum (Dialictus) sp7
Lasioglossum (Dialictus) sp8
Lasioglossum (Dialictus) sp9
Lasioglossum (Dialictus) sp10
Lasioglossum (Dialictus) sp11
Lasioglossum (Evylaeus) sp1
Family Megachilidae
Subfamily Megachilinae
Tribe Anthidini
Anthidiellum (Loyolanthidium) apicale

LS Nest S1
SO G
X

S2
-

S3
X

S4
-

S5
X

S6 TSE
X
35

PS

PS

PS

32

EU

110

EU G
UK UK
UK UK
UK UK
EU G

X
X
-

X
-

X
X
X

X
X
X
X
-

X
X
-

X
X
X

79
1
24
1
2

EU

16

EU

14

PS
UK
UK
UK

G
G
G
G

X
X
X
-

X
X
-

X
X
X
-

X
X
X
X

X
X
X
X

36
22
19
8

PS

PS

PS
CL

G
-

X
X

X
-

74
1

PS

EU

EU
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK

G
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK
UK

X
X
X
X
-

X
X
X
X
X
X
-

X
X
X
X
X
-

X
X
X
X
X
X
X
-

X
X
X
X
X
X
X
X
X

X
X
X
X
X
X

2
99
104
46
7
23
1
9
2
1
1
2
7

Volume 120, Number 5, November and December 2009

Superfamily Apoidea: Clade Anthophila (Bees) LS Nest S1

(Cresson, 1878)
SO C
Anthodioctes (Anthodioctes) gualanensis
(Cockerell, 1912)
SO C
Hypanthidium (Hypanthidium) mexicanum
(Cresson, 1878)
SO C
X
Stelis (Dolichostelis) costalis
Cresson, 1872
SO
Tribe Megachilini
Coelioxys (Acrocoelioxys) azteca
Cresson, 1878
CL
X
Coelioxys (Acrocoelioxys) otomita
Cresson, 1878
CL
Coelioxys (Acrocoelioxys) sp1
CL
X
Coelioxys (Cyrtocoelioxys) sp1
CL
X
Coelioxys (Cyrtocoelioxys) speculifera
Cockerell, 1931
CL
Coelioxys (Glyptocoelioxys) cf.
vituperabilis Holmberg, 1903
CL
Coelioxys (Glyptocoelioxys) sp1
CL
Coelioxys (Haplocoelioxys) sp1
CL
X
Coelioxys (Neocoelioxys) similima
Smith, 1854
CL
X
Coelioxys (Neocoelioxys) cf. similima
Smith, 1854
CL
Coelioxys (Rhinocoelioxys) sp1
CL
Megachile (Acentron) albitarsis
Cresson,1872
SO C
X
Megachile (Austromegachile) sp1
SO C
Megachile (Chelostomoides) otomita
Cresson, 1878
SO C
Megachile (Chelostomoides)
quadridentata Mitchell, 1930
SO C
Megachile (Chelostomoides) sp1
SO C
Megachile (Chelostomoides) sp2
SO C
Megachile (Chelostomoides) texensis
Mitchell, 1956
SO C
X
Megachile (Leptorachis) sp1
SO C
Megachile (Melanosarus) nigripennis
Spinola, 1841
SO C
X
Megachile (Neochelynia) chichimeca
Cresson, 1878
SO C
X
Megachile (Neochelynia) coelioxoides
Cresson, 1878
SO C
Megachile (Neochelynia) sp2
SO C
X
Megachile (Neochelynia) sp3
SO C
Megachile (Pseudocentron) inscinta
SO C
X
Megachile (Pseudocentron) sp1
SO C
Megachile (Pseudocentron) sp2
SO C
Megachile (Pseudocentron) sp3
SO C
X
Megachile (Sayapis) zaptlana
Cresson, 1878
SO C
X
Megachile (Sayapis) frugalis
Cresson, 1872
SO C
Megachile (Sayapis) sp1
SO C
X
Tribe Osmiini
Ashmeadiella (Ashmeadiella) cf.
bequaerti Cockerell, 1931
SO C
-

535

S2
-

S3
X

S4
-

S5
X

S6 TSE
X
10

16

13

X
X

X
-

1
2
4

X
X

X
-

1
2
2

X
-

2
1

X
-

X
-

X
-

31
4

15

X
-

X
X

X
-

3
1
2

1
9

38

X
X
X
X
-

X
X
-

X
X
-

X
-

X
X
X
X
X

5
27
4
6
2
3
3

24

X
-

X
-

2
1

10

536

ENTOMOLOGICAL NEWS

Superfamily Apoidea: Clade Anthophila (Bees) LS Nest S1

Heriades (Neotrypetes) variolosus
purpurascens (Cockerell 1931)
SO C
X
Family Apidae
Subfamily Anthophorinae
Tribe Centridini
Centris (Heterocentris) analis
(Fabricius, 1804)
SO C
X
Centris (Heterocentris) nitida Smith,
1874
SO C
X
Centris (Heterocentris) sp. nov.
SO C
X
Centris (Heterocentris) trigonoides
Lepeletier, 1841
SO C
X
Centris (Trachina) eurypatana
Snelling, 1984
SO G
Tribe Emphorini
Diadasia (Dasiapis) tropicalis
(Cockerell, 1918)
UK G
Tribe Eucerini
Florilegus (Florilegus) condignus
(Cresson, 1878)
SO G
X
Gaesischia (Gaesischiana) exul Michener,
LaBerge and Moure, 1955
UK G
Melissodes (Melissodes) tepaneca
Cresson, 1878
SO G
X
Svastra (Epimelissodes) nitida
(LaBerge, 1956)
SO G
Tribe Exomalopsini
Ancylocelis apiformis (Fabricius, 1793) SO G
Exomalopsis (Exomalopsis) analis
Spinola, 1853
SO G
Exomalopsis (Exomalopsis) boharti
Timberlake, 1980
SO G
X
Exomalopsis (Exomalopsis) mellipes
Cresson, 1878
SO G
Exomalopsis (Exomalopsis) similis
Cresson, 1865
SO G
Paratetrapedia (Lophopedia) apicalis
(Cresson, 1878)
UK G
Paratetrapedia (Paratetrapedia) moesta
(Cresson, 1878)
UK G
X
Paratetrapedia (Xanthopedia) swainsonae
(Cockerell, 1909)
SO G
Tribe Osirini
Osiris semiatratus Shanks, 1987
CL
Tribe Tetrapedini
Tetrapedia maura Cresson, 1878
SO C
Subfamily Nomadinae
Tribe Epeolini
Epeolus sp1
CL
Triepeolus cameroni
(Meade-Waldo, 1913)
CL
Subfamily Xylocopinae
Tribe Ceratinini
Ceratina (Calloceratina) amabilis
Cockerell, 1931
PS W
X
Ceratina (Calloceratina) itzarum

S2

S3

S4

S5

S6 TSE

88

X
X

X
-

9
13

47

11

78

37

35

16

44

119

71

38

Volume 120, Number 5, November and December 2009

Superfamily Apoidea: Clade Anthophila (Bees) LS Nest

Cockerell, 1931
PS W
Ceratina (Calloceratina) sp1
UK W
Ceratina (Calloceratina) sp4
UK W
Ceratina (Calloceratina) sp5
UK W
Ceratina (Calloceratina) sp6
UK W
Ceratina (Ceratinula) cf. arizonensis
Cockerell, 1898
PS W
Ceratina (Ceratinula) sp2
UK W
Ceratina (Zadontomerus) cf. nautlana
Cockerell, 1897
PS W
Tribe Xylocopini
Xylocopa (Neoxylocopa) mexicanorum
Cockerell, 1912
PS W
Xylocopa (Neoxylocopa) nautlana
Cockerell, 1904
PS W
Xylocopa (Schonnherria) muscaria
(Fabricius, 1775)
PS W
Xylocopa (Stenoxylocopa) micheneri
Hurd, 1978
PS W
Subfamily Euglossinae
Eufriesea mexicana (Mocsry, 1897)
SO C
Euglossa (Euglossa) viridissima Friese,
1899
PS C
Eulaema (Apeulaema) polychroma
(Mocsry, 1899)
PS C
Exaerete smaragdina (Gurin-Mneville,
1845)
CL
Subfamily Meliponinae
Cephalotrigona zexmeniae (Cockerell,
1912)
EU C
Frieseomelitta nigra (Cresson, 1878)
EU C
Lestrimelitta niitkib Ayala, 1999
CL C
Nannotrigona perilampoides
(Cresson, 1878)
EU C
Partamona bilineata (Say, 1837)
EU C
Plebeia frontalis (Friese, 1911)
EU C
Plebeia moureana Ayala, 1999
EU C
Trigona fulviventris GurinMneville, 1845
EU C
Trigona fuscipennis Friese 1900
EU C
Trigonisca pipioli Ayala, 1991
EU C

537

S1
X
X
-

S2
X
X
X
-

S3
X
-

S4
X
-

S5 S6 TSE
X X
11
- X
17
- X
7
X
2
X
1

X
-

X
-

X
-

X
X

X
-

21
1

144

14

10

21

X
X
X

X
X
-

X
X
X

X
-

X
X
X

X 128
X 2289
X
7

X
X
X
-

X
X

X
X
-

X
X
-

X
X
-

X
X
X

X
X

X
X

X
X

X 296
7
X 1632

Total of specimens examined

Total of species

202
1
82
155

6785
55

60

58

30

69

79

152

538

ENTOMOLOGICAL NEWS

Figure 2. Overall native bee genera and species richness by family in six Natural Protected
Areas from Yucatan, Mexico.

The highest species richness was observed in Megachile (20 species, 15.4%),
Lasioglossum (12 species, 9.23%), Coelioxys (11 species. 8.46%), Ceratina (9
species, 6.9%) and Augochlora (8 species, 6.1%); that is, 10% of the genera contained 46% of the species. The remaining species were divided among 11 genera
with two to five species each and 34 genera with just one species each. Eighty
(61.5%) species and 45 (34.6%) morphospecies were identified (13 of them were
unisexual), while five (3.8%) species varied slightly from the original species
descriptions, so were identified as near or affinis to the closest named species.
Richness in each NPA exhibited differences in the species composition of each
family. Yalahau contained the highest taxonomic richness in terms of generic and
species totals, followed by Tabi, Dzilam, Kabah, Dzibilchaltun and El Palmar

Figure 3. Native bee genera and species richness in six Natural Protected Areas from
Yucatan, Mexico.

Volume 120, Number 5, November and December 2009

539

(Fig. 3). Apidae was the richest family in most of the NPAs, with the exceptions
of El Palmar and Tabi, where the Halictidae were better represented.
The areas of deciduous vegetation in northern Yucatan contained more megachilids than halictids, while in the center of the state (Yalahau) these two families had approximately equal numbers of species. In the south, Halictidae was
more speciose than Megachilidae. Halictidae was also more speciose in El Palmar, which had the lowest overall richness, but a significant number of exclusive
species (Table 2).
The most frequent lifestyle in all the NPAs was solitary (39%), followed by
parasocial and cleptoparasitic (14% each), eusocial (12%), and 21% remain unknown. This pattern held at all the NPAs save for El Palmar, where no cleptoparasitic species were recorded and most of their species belong to the unknown life style category (Fig. 4).
Overall, most of the recorded species are cavity-nesting, followed by groundnesting and wood-nesting species. This pattern varied when analyzed by NPA,
with Dzibilchaltun and Dzilam having more cavity-nesting species than Tabi,
Kabah, Yalahau and El Palmar; the latter having a higher proportion of groundnesting species (Fig. 5).
DISCUSSION
Previously published species counts for Yucatan (Ayala et al., 1996; Ayala,
1999; Novelo-Rincon et al., 2003) contained counts of 107 species and many
specimens classified as morphospecies. In the present study, 130 species were
found of which 86 were identified and 44 classified as morphospecies. Of the 86
identified species, 32 are new records, which, when added to the previously
reported 107 species, make for a total of 139 listed bee species in Yucatan.
However, given the number of morphospecies found in previous reports and the
present study, it is quite possible that the regional bee fauna consists of closer to
200 species. Unfortunately, the high percentage of species classified as morphospecies in fauna reports in Mexico (> 55% of records) prevents any close
comparison of regional faunal composition. For example, Novelo-Rincon et al.
(2003) reported 64 species from coastal dune vegetation, 71 in dry forest and 104
in semi-deciduous tropical forest. The number of species identified for each of
these zones in the present study differs from their report, probably due to differences in sampling effort and collection methods (e.g. Novelo-Rincon et al., 2003
used no traps).
According with Heithaus (1974) bee community structure and composition
depends on vegetation structure and composition, so we assume that most of the
differences in bee composition among the NPAs could be explained by this factor.
Overall richness among the six NPAs is lower than reported for other regions
in Mexico: 346 species were identified in Morelos (Hinojosa-Diaz, 2003); 259 in
Puebla (Vergara and Ayala, 2002); 238 in Jalisco (Ayala, 2004); and 180 in Hi-

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Figure 4. Native bee lifestyle proportions within each Natural Protected Area from
Yucatan, Mexico.

dalgo (Godinez-Garcia et al., 2004). This agrees with Ayala et al. (1993 and
1996), who state that bee species richness in Yucatan is low compared to other
regions of Mexico, possibly due to subsoil conditions that restrict nesting for
many species (Roubik, 1989; Michener, 2007), as occurs in Florida (Pascarella
et al., 2000), and the young geological age of the Yucatan Peninsula.
Apidae had the highest species richness among the families recorded in the six
NPAs, which agrees with faunal studies from other regions in Mexico (GodinezGarcia, 1991; Roubik et al., 1991; Estrada, 1992; Fierros-Lopez, 1996; HinojosaDiaz, 2003; Ayala, 2004). In these studies, however, Halictidae is more speciose
than Megachilidae, which contrasts with the higher species content for Megachilidae observed here. Differences in species composition are mainly due to the
absence of species belonging to Lasioglossum s. str. and the greater richness of
species of Coelioxys and Megachile in Yucatan (most of them at Northern). This
greater richness is possibly because these bees do not need soil for nesting. Despite these differences, the fact that most of the genera recorded here contained
few species and most of the species belonged to a very few genera is similar to
other Mexican faunal studies (Godinez-Garcia, 1991; Roubik et al., 1991; Estrada, 1992; Fierros-Lopez, 1996; Hinojosa-Diaz, 2003; Ayala, 2004).
Lifestyle proportions were similar between the NPAs, but different as far as
species composition, which agrees with data for native bee communities reported by Novelo et al. (2003). This is interesting in terms of functional diversity
since bee lifestyles are a summary expression of how bees use the resources in
their medium, both for food and nest construction and permanence. There were
more ground-nesting species at the Southern NPAs, possibly because of deeper
soils than in the Northern (Bautista et al., 2005).

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541

Figure 5. Native bee nesting strategy proportions within each Natural Protected Area
from Yucatan, Mexico.

Previous reports of bee species geographic distribution (Ayala et al., 1993

Ayala et al., 1996; Michener, 1979 and 2007) indicate the bee fauna of Yucatan
to be mainly Neotropical with some elements broadly distributed in the Americas
(Florilegus condignus and Augochloropsis metallica). Some Neotropical elements with distribution in the Antilles have also been identified (Exomalopsis
similis, Temnosoma smaragdinum, and one species previously known only in
Jamaica, Paratetrapedia swainsonae). Another group contains species of
Nearctic affinity but with distribution reaching to southern Mexico and some as
far as Central America (Colletes arizonensis, the three Augochlorella species,
Heriades variolosus, Megachile albitarsis, Melissodes tepaneca and Stelis
costalis). Finally, there is a group of species endemic to the Yucatan Peninsula
such as Ceratina itzarum (founded in all the NPAs except El Palmar),
Ashmeadiella bequaerti and Osiris semiatratus (founded in Yalahau and Kabah).
Considering only those species identified here to the species level, the six
studied NPAs contain 62% of the known bee species in Yucatan. This is vital
information since many tropical NPAs lack data on the organisms they are
intended to protect, particularly on the most diverse animals: insects (Rodrigues
and Gaston, 2002; Samways, 2007). Increasing anthropogenic impact on tropical
ecosystems (particularly dry ones) makes generation of data like that produced
here even more urgent because it is indispensable for developing conservation
and sustainable management strategies for these areas (Sanchez Azofeifa et al.,
2005).
The data generated here on the bee fauna of six NPAs in Yucatan, Mexico,
will form the foundation for more in-depth research on the impact of environmental fragmentation and/or changes in land use in bee communities. This

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information can be utilized in the creation of natural resources management

plans with the broadest possible coverage which take into account the importance of bees in the reproduction of many flowering plant species. This is particularly the case for dry tropical forest vegetation, which is recognized as one of
the vegetation types in the tropical Americas most threatened by human activities, and the complex dynamic of which requires specific conservation strategies
(Bullock et al., 1995; Sanchez Azofeifa et al., 2005).

ACKNOWLEDGMENTS
The authors thank Victor Parra and Jorge Navarro for their valuable contributions to early versions of this manuscript. Special thanks to John S. Ascher (AMNH) for valuable help in identifying
bee species and providing unpublished data on Yucatan bee distribution, taxonomy, and nomenclature, and to Jerome G. Rozen Jr. for granting facilities at AMNH. Antonio Aguiar helped with identifying the Paratetrapedia species and Molly Rightmyer with identifying the Triepeolus species.
Roger Cauich and Laura Meneses helped in the development of this research. Enrique Reyes-Novelo
received a CONACYT postgraduate scholarship (171289). This research was supported by CONACYT-SEMARNAT (2004-C01-180/A-1) to Virginia Melendez Ramirez.

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