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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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BEE FAUNAS (HYMENOPTERA: APOIDEA) OF SIXNATURAL 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, aredescribed and analyzed. Atotal of 130 bee species was recorded belonging to 50 genera and five fam-ilies, of which 32 are new records for Yucatan State. The genera with the highest species richnesswere Megachile (20 species), Lasioglossum (12 species), Coelioxys (11 species, most of these arecleptoparasites ofMegachile), Ceratina (9 species) andAugochlora (8 species). Species richness byNPA was 30 species in El Palmar; 55 in Dzibilchaltn; 58 in Kabah; 60 in Dzilam; 69 in Tabi; and79 in Yalahau. Bee faunal composition in the NPAs, biogeographic affinities and the importance ofwild 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 flow-

ering 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 diver-

sity 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 Bio-

sphere Reserve in Jalisco state (Ayala, 2004); Mapimi Biosphere Reserve in Dur-

ango 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 fau-

nal 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.

530 ENTOMOLOGICAL NEWS

______________________________

1 Received on September 23, 2008. Accepted on November 9, 2008.2 Departamento de Zoologa, Campus de Ciencias Biolgicas y Agropecuarias, Universidad Auto-nma de Yucatan. Apartado Postal 4-116 Colonia Itzimna 97100 Mrida, Yucatan, Mxico. E-mails:(ER-N) [email protected] (VM-R) [email protected] (HD-G) [email protected]

3 Estacin de Biologa Chamela, Instituto de Biologa, Universidad Nacional Autonma de Mxico.Apartado Postal 21,48980 San Patricio, Jalisco, Mxico. E-mail: [email protected].

Mailed on December 13, 2010


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(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 morpho-

species belonging to taxonomically unrevised genera, meaning they are unrec-

ognized 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 collect-

ed 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 nat-

ural 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 polli-

nation studies, and to promote bee conservation and management in NPAs.

METHODS

An examination was made of wild bee specimens on deposit in the RegionalEntomological Collection of the Universidad Autonoma de Yucatan (CER-

UADY) 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, Jalis-

co, Mexico (IBUNAM-Chamela) and the American Museum of Natural History

(New York, USA) (AMNH) (with the support of Dr. John Ascher), review of bib-

liographic material (published and unpublished catalogs from AMNH) and con-

sultation with experts in specific groups (see Acknowledgements). The Meli-

poninae classification was according to Camargo and Pedro (2008).

Volume 120, Number 5, November and December 2009 531


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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

Yalahau 2040'37", 2034'59" N;

8910'49", 8915'00" W 5683.28 TDF AW0

Dzilam 2126'33", 2124'51" N;

8847'54", 8847'49" W 61706.83 SFF BS1

El Palmar 2055'00", 2111'00" N;

9000'00", 9022'30" W 50177.39 CDV BS0

Kabah 2014'12", 2013'22" N;

8939'34", 8938'03" W 949.76 SDTF AW0

Tabi 2014'47", 2014'59" N;

8930'26", 8932'29" W 1355.74 SDTF AW0

Veg = Vegetation types, TDF = Tropical Deciduous Forest, SFF = Seasonally FloodedForest, 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.


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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 sub-

strates 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 pre-

sented 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 rep-

resent 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 gen-

era and species) in the six studied NPAs was the Apidae, followed by Megachili-

dae, 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 S1 S2 S3 S4 S5 S6 TSE

Family ColletidaeSubfamily Colletinae

Colletes arizonensis Stephen, 1954 SO G - - X - X X 8Colletes punctipennis maurus

Stephen, 1954 SO G - X - - X - 4Subfamily Diphaglossinae

Tribe CaupolicaniniPtiloglossa eximia (Smith, 1861) SO G - - X - - - 1

Subfamily HylaeinaeHylaeus sp1 SO C X - - - - X 3Hylaeus sp2 SO C - - - - X X 2Hylaeus sp3 SO C - - - X - - 1Hylaeus sp4 SO C - - - - X - 1Hylaeus sp5 SO C - X - - - - 1

Family AndrenidaeSubfamily Panurginae

Calliopsis hondurasica Cockerell, 1949 SO G - - - - - X 6Pseudopanurgus crenulatus



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(Cockerell, 1905) SO G X - X - X X 35Family Halictidae

Subfamily HalictinaeTribe AugochloriniAugochlora (Augochlora) albiceps

Friese, 1925 PS W - - - - X - 1Augochlora (Augochlora) nigrocyanea

Cockerell, 1897 PS W - - - X X X 5Augochlora (Augochlora) smaragdina

Friese, 1917 PS W X X X X X X 32Augochlora (Oxystoglosella) aurifera

Cockerell, 1897 EU G - X X - X X 110Augochlora (Oxystoglosella)

cordiaefloris Cockerell, 1907 EU G X X X X X X 79

Augochlora sp1 UK UK - - - X - - 1Augochlora sp2 UK UK X - X X X X 24Augochlora sp3 UK UK - - - X - - 1Augochlorella bracteata Ordway, 1966 EU G - - X - - X 2Augochlorella neglectula (Cockerell,

1897) EU G X X X - X X 16Augochlorella pomoniella (Cockerell,

1915) EU G X X X - X X 14Augochloropsis (Paraugochloropsis)

metallica (Fabricius, 1793) PS G X X X - X X 36Augochloropsis sp1 UK G X X X - X X 22Augochloropsis sp2 UK G X - X - X X 19Augochloropsis sp3 UK G - - - - X X 8Caenaugochlora (Caenaugochlora)

gemmella (Cockerell, 1912) PS G - - X - X - 6Pseudaugochlora graminea

(Fabricius, 1804) PS G - - X - X X 7Caenohalictus amatitlana (Cockerell,

1912) PS G - - X - X - 74Temnosoma smaragdinum Smith, 1879 CL - - - X - - - 1

Tribe HalictiniAgapostemon (Notagapostemon) nasutus

Smith, 1853 PS G - - - X - - 2Halictus (Odontalictus) ligatus

Say, 1837 EU G - - X - X - 4Halictus (Seladonia) hesperus

Smith, 1862 EU G - - - - X - 2Lasioglossum (Dialictus) sp1 UK UK X X X X X X 99Lasioglossum (Dialictus) sp2 UK UK X X X X X X 104Lasioglossum (Dialictus) sp3 UK UK X X X X X X 46Lasioglossum (Dialictus) sp4 UK UK - - X - X X 7Lasioglossum (Dialictus) sp5 UK UK X X - - X X 23Lasioglossum (Dialictus) sp6 UK UK - X - - - - 1Lasioglossum (Dialictus) sp7 UK UK - - X X X - 9Lasioglossum (Dialictus) sp8 UK UK - - - X X - 2Lasioglossum (Dialictus) sp9 UK UK - - - X - - 1Lasioglossum (Dialictus) sp10 UK UK - X - - - - 1Lasioglossum (Dialictus) sp11 UK UK - - - X - - 2Lasioglossum (Evylaeus) sp1 UK UK - - - - X X 7

Family MegachilidaeSubfamily Megachilinae

Tribe AnthidiniAnthidiellum (Loyolanthidium) apicale



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(Cresson, 1878) SO C - - X - X X 10Anthodioctes (Anthodioctes) gualanensis

(Cockerell, 1912) SO C - X X - X - 3Hypanthidium (Hypanthidium) mexicanum

(Cresson, 1878) SO C X X X - X X 16Stelis (Dolichostelis) costalis

Cresson, 1872 SO - - - X - - - 1Tribe Megachilini

Coelioxys (Acrocoelioxys) aztecaCresson, 1878 CL - X X X - X X 13

Coelioxys (Acrocoelioxys) otomitaCresson, 1878 CL - - X - - - - 1

Coelioxys (Acrocoelioxys) sp1 CL - X - X - - - 2Coelioxys (Cyrtocoelioxys) sp1 CL - X X - - - X 4

Coelioxys (Cyrtocoelioxys) speculiferaCockerell, 1931 CL - - X - - - - 2

Coelioxys (Glyptocoelioxys) cf.vituperabilis Holmberg, 1903 CL - - X - - - - 1

Coelioxys (Glyptocoelioxys) sp1 CL - - - - - X - 2Coelioxys (Haplocoelioxys) sp1 CL - X X - - - - 2Coelioxys (Neocoelioxys) similima

Smith, 1854 CL - X X - - X X 5Coelioxys (Neocoelioxys) cf. similima

Smith, 1854 CL - - - X - - - 2Coelioxys (Rhinocoelioxys) sp1 CL - - - - - X - 1Megachile (Acentron) albitarsis

Cresson,1872 SO C X X X - - X 31Megachile (Austromegachile) sp1 SO C - - - - X - 4Megachile (Chelostomoides) otomita

Cresson, 1878 SO C - - X - X - 15Megachile (Chelostomoides)

quadridentata Mitchell, 1930 SO C - X - - - X 3Megachile (Chelostomoides) sp1 SO C - - X - - - 1Megachile (Chelostomoides) sp2 SO C - - X - X - 2Megachile (Chelostomoides) texensis

Mitchell, 1956 SO C X - - - - - 1Megachile (Leptorachis) sp1 SO C - X X - X X 9Megachile (Melanosarus) nigripennis

Spinola, 1841 SO C X - - - - X 2Megachile (Neochelynia) chichimeca

Cresson, 1878 SO C X X X X X X 38Megachile (Neochelynia) coelioxoides

Cresson, 1878 SO C - - X - X - 5Megachile (Neochelynia) sp2 SO C X X - - - X 27Megachile (Neochelynia) sp3 SO C - X - - - X 4Megachile (Pseudocentron) inscinta SO C X X X - - X 6Megachile (Pseudocentron) sp1 SO C - X - X - - 2Megachile (Pseudocentron) sp2 SO C - - - X - X 3Megachile (Pseudocentron) sp3 SO C X - - - - X 3Megachile (Sayapis) zaptlana

Cresson, 1878 SO C X X - X - X 24Megachile (Sayapis) frugalis

Cresson, 1872 SO C - X - - X - 2Megachile (Sayapis) sp1 SO C X - - - - - 1

Tribe OsmiiniAshmeadiella (Ashmeadiella) cf.

bequaerti Cockerell, 1931 SO C - - - - - X 10



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Heriades (Neotrypetes) variolosuspurpurascens (Cockerell 1931) SO C X X - X X X 88

Family ApidaeSubfamily Anthophorinae

Tribe CentridiniCentris (Heterocentris) analis

(Fabricius, 1804) SO C X - - - - - 3Centris (Heterocentris) nitida Smith,

1874 SO C X X - - X - 9Centris (Heterocentris) sp. nov. SO C X X - - - X 13Centris (Heterocentris) trigonoides

Lepeletier, 1841 SO C X X - - - X 47Centris (Trachina) eurypatana

Snelling, 1984 SO G - X - - - - 1

Tribe EmphoriniDiadasia (Dasiapis) tropicalis

(Cockerell, 1918) UK G - - - - - X 3Tribe EuceriniFlorilegus (Florilegus) condignus

(Cresson, 1878) SO G X - - - - X 5Gaesischia (Gaesischiana) exul Michener,

LaBerge and Moure, 1955 UK G - - - - X X 11Melissodes (Melissodes) tepaneca

Cresson, 1878 SO G X X X X - X 78Svastra (Epimelissodes) nitida

(LaBerge, 1956) SO G - - - - - X 1Tribe ExomalopsiniAncylocelis apiformis (Fabricius, 1793) SO G - - - - - X 3Exomalopsis (Exomalopsis) analis

Spinola, 1853 SO G - - X - X X 37Exomalopsis (Exomalopsis) boharti

Timberlake, 1980 SO G X - - - X X 35Exomalopsis (Exomalopsis) mellipes

Cresson, 1878 SO G - - - - X X 16Exomalopsis (Exomalopsis) similis

Cresson, 1865 SO G - X X X X X 44Paratetrapedia (Lophopedia) apicalis

(Cresson, 1878) UK G - X X - - X 3Paratetrapedia (Paratetrapedia) moesta

(Cresson, 1878) UK G X X X - X X 119Paratetrapedia (Xanthopedia) swainsonae

(Cockerell, 1909) SO G - - X - X X 71Tribe Osirini

Osiris semiatratus Shanks, 1987 CL - - - X - - X 2Tribe Tetrapedini

Tetrapedia maura Cresson, 1878 SO C - - - - - X 1Subfamily Nomadinae

Tribe EpeoliniEpeolus sp1 CL - - - - - X - 2Triepeolus cameroni

(Meade-Waldo, 1913) CL - - - X - - - 2Subfamily Xylocopinae

Tribe CeratininiCeratina (Calloceratina) amabilis

Cockerell, 1931 PS W X X X - X X 38Ceratina (Calloceratina) itzarum



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Cockerell, 1931 PS W X X X - X X 11

Ceratina (Calloceratina) sp1 UK W - X - X - X 17

Ceratina (Calloceratina) sp4 UK W X X - - - X 7Ceratina (Calloceratina) sp5 UK W - - - - X - 2

Ceratina (Calloceratina) sp6 UK W - - - - X - 1

Ceratina (Ceratinula) cf. arizonensis

Cockerell, 1898 PS W X X - X X X 21

Ceratina (Ceratinula) sp2 UK W - - - - X - 1

Ceratina (Zadontomerus) cf. nautlana

Cockerell, 1897 PS W X X X X X X 144

Tribe Xylocopini

Xylocopa (Neoxylocopa) mexicanorum

Cockerell, 1912 PS W X X X X - X 14

Xylocopa (Neoxylocopa) nautlanaCockerell, 1904 PS W - - - - - X 2

Xylocopa (Schonnherria) muscaria

(Fabricius, 1775) PS W X - - - - X 3

Xylocopa (Stenoxylocopa) micheneri

Hurd, 1978 PS W - X - X - - 2

Subfamily Euglossinae

Eufriesea mexicana (Mocsry, 1897) SO C - - - - - X 1

Euglossa (Euglossa) viridissima Friese,

1899 PS C X - X - - X 9

Eulaema (Apeulaema) polychroma

(Mocsry, 1899) PS C X X - - - X 10

Exaerete smaragdina (Gurin-Mneville,

1845) CL - X X X - X X 21

Subfamily Meliponinae

Cephalotrigona zexmeniae (Cockerell,

1912) EU C X X X - X X 128

Frieseomelitta nigra (Cresson, 1878) EU C X X X X X X 2289

Lestrimelitta niitkib Ayala, 1999 CL C X - X - X X 7

Nannotrigona perilampoides

(Cresson, 1878) EU C X X X - X X 202

Partamona bilineata (Say, 1837) EU C X - - - - - 1

Plebeia frontalis (Friese, 1911) EU C X - X - X X 82

Plebeia moureana Ayala, 1999 EU C - X - - - - 155Trigona fulviventris Gurin-

Mneville, 1845 EU C - X X X X X 296

Trigona fuscipennis Friese 1900 EU C - X - - - - 7

Trigonisca pipioli Ayala, 1991 EU C X X X X X X 1632

Total of specimens examined 6785

Total of species 55 60 58 30 69 79 152



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The highest species richness was observed inMegachile (20 species, 15.4%),

Lasioglossum (12 species, 9.23%), Coelioxys (11 species. 8.46%), Ceratina (9

species, 6.9%) andAugochlora (8 species, 6.1%); that is, 10% of the genera con-

tained 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 2. Overall native bee genera and species richness by family in six Natural ProtectedAreas from Yucatan, Mexico.

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


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(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 mega-

chilids than halictids, while in the center of the state (Yalahau) these two fami-

lies had approximately equal numbers of species. In the south, Halictidae was

more speciose than Megachilidae. Halictidae was also more speciose in El Pal-

mar, 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 un-

known. This pattern held at all the NPAs save for El Palmar, where no clep-

toparasitic species were recorded and most of their species belong to the un-

known life style category (Fig. 4).

Overall, most of the recorded species are cavity-nesting, followed by ground-

nesting 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 ground-

nesting 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 mor-

phospecies in fauna reports in Mexico (> 55% of records) prevents any closecomparison 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 differ-

ences 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 fac-tor.

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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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 formany 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 (Godinez-

Garcia, 1991; Roubik et al., 1991; Estrada, 1992; Fierros-Lopez, 1996; Hinojosa-

Diaz, 2003; Ayala, 2004). In these studies, however, Halictidae is more speciose

than Megachilidae, which contrasts with the higher species content for Mega-

chilidae observed here. Differences in species composition are mainly due to the

absence of species belonging toLasioglossum s. str. and the greater richness ofspecies of Coelioxys andMegachile in Yucatan (most of them at Northern). This

greater richness is possibly because these bees do not need soil for nesting. Des-

pite 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; Estra-

da, 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 report-

ed by Novelo et al. (2003). This is interesting in terms of functional diversitysince 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).


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


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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 ele-

ments 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 environ-

mental fragmentation and/or changes in land use in bee communities. This


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


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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 impor-

tance of bees in the reproduction of many flowering plant species. This is partic-

ularly the case for dry tropical forest vegetation, which is recognized as one of

the vegetation types in the tropical Americas most threatened by human activi-

ties, 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 ver-

sions 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 nomencla-ture, and to Jerome G. Rozen Jr. for granting facilities at AMNH. Antonio Aguiar helped with iden-

tifying 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 CONA-

CYT-SEMARNAT (2004-C01-180/A-1) to Virginia Melendez Ramirez.

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