2002 vergara&ayala beesof zap

Diversity, Phenology and Biogeography of the Bees (Hymenoptera: Apoidea) of Zapotitlán delas Salinas, Puebla, MexicoAuthor(s): Carlos H. Vergara and Ricardo AyalaSource: Journal of the Kansas Entomological Society, Vol. 75, No. 1 (Jan., 2002), pp. 16-30Published by: Allen Press on behalf of Kansas (Central States) Entomological SocietyStable URL: http://www.jstor.org/stable/25086035 .Accessed: 21/02/2011 13:18

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JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY 75(1), 2002, pp. 16-30

Diversity, Phenology and Biogeography of the Bees

(Hymenoptera: Apoidea) of Zapotitl?n de las Salinas, Puebla, Mexico

Carlos H. Vergara1 and Ricardo Ayala2

abstract: A faunistic study of the bees (Apoidea) of a semi-desert in the Transverse Neovol

canic Belt of central Mexico was carried out by systematic monthly collecting, during one year. A

total of 259 species, 69 genera and seven families were recorded; 44 are undescribed species, 46

are new records for the state of Puebla and, of these, 3 are also new records for Mexico. The great est diversity, at the familial level, was found for Apidae, with 33 genera and 92 species and, at the

generic level, for Lasioglossum Curtis with 29 species, and Perdita Smith, with 20 species. The

greatest monthly diversity was observed during September, when 57 genera were active. The main

biogeographic elements found, at the generic level, are of cosmopolitan affinity (22%), neotropical

affinity (20%) and of neartic affinity (13%). Our results agree with the prediction that the Tehuac?n

desert would present a number of species intermediate between the xeric regions of the US and the

mesic areas of the Mexican Plateau. Also, the number of species common to the deserts of Sonora

and Chihuahua, on the one hand, and Tehuac?n, on the other, is low, reflecting differences in the

floristic composition of the three regions. key words: Biogeography, bees, Hymenoptera, Apoidea, diversity

Introduction

Mexico has a rich fauna of bees, consisting of 1800 species and subspecies grouped in

144 genera and seven families. This number could rise to over 2000, once several taxa

have been revised and poorly collected regions of the country are better known (Ayala et

al., 1996). In global terms, the greatest diversity of bees is known from semi-desertic,

warm, extratropical regions of the world. The greatest regional number of species has been

recorded in deserts and shrublands of California and Arizona, but records from the corre

sponding adjacent regions of Mexico are lacking (Michener, 2000). Other deserts of Mex

ico such as the Tehuac?n-Cuicatl?n Valley, where the study region is included, remain

poorly known from the viewpoint of insects. Only 36 bee species had been found in 17

sites collected until 1993 (Ayala et al., 1993). Faunistic studies on bee communities in Mexico have been carried out in Chamela,

Jalisco (Ayala, 1988), in Sierra del Tigre, Jalisco (Estrada, 1992), in the Biosphere Reserve

of Sian Ka'an, Quintana Roo (Roubik et al., 1991), in San Gregorio, Guanajuato (God?nez,

1991), and in the Volc?n de Tequila, Jalisco (Fierros, 1998). This paper presents information on the species diversity, phenology and biogeography

of bees from the Valley of Zapotitl?n de las Salinas.

Methods and Materials

Study area

The Valley of Zapotitl?n de las Salinas is part of the Floristic Province of Tehuac?n

Cuicatl?n (Rzedowski, 1981). It is located between 8? 14' and 18?22'N and 97?26' and

1 Departamento de Qu?mica y Biolog?a, Universidad de las Am?ricas-Puebla, Santa Catarina M?rtir, Puebla,

72820 Mexico. [email protected] 2 Estaci?n de Biolog?a Chamela, Instituto de Biolog?a, Universidad Nacional Aut?noma de M?xico, Apartado

Postal 21, San Patricio, Jalisco, 48980 M?xico. [email protected]

VOLUME 75, ISSUE 1 17

97?34'W. The valley is drained by the Salado River, which is part of the Papaloapan system.

The climate, measured at the meteorological station of Zapotitl?n Salinas, is dry with summer rains, described as Bsohw" (w)(e)g in the Koppen classification, modified for

Mexico (Garcia, 1973). The vegetation of the Valley is xerophitic scrub, constituted by four vegetation units:

thorny shrub, tetechera [Neobuxbaumia tetetzo (A. Weber ex. K. Schum) Backeb (Cac

taceae) forest], cardonal forest [Cephalocereus hoppenstedii K. Scum (Cactaceae)] and

izotal [Beaucarnea gracilis Hart (Liliaceae) forest] (Zavala, 1982).

Bee diversity

Nine sites within the Valley were visited seven times, in 1-2 day trips between March

1994 and July 1996. Collecting was done at forty sites, including the nine visited initially, in three to five-day collecting trips, every month from July 1996 through June 1997. Bees

were observed and collected on flowers, or in other activities (flight, nesting, etc.). The collected specimens were sacrificed, pinned, labeled, and sent for determination to the au

thorities mentioned in the Acknowledgements section. One of us, Ricardo Ayala (Instituto de Biolog?a, UNAM, Estaci?n de Biolog?a Chamela), determined the Centridini. Vouch ers are deposited at the Entomological Collection, Universidad de las Am?ricas-Puebla,

Cholula, Puebla, Mexico.

A list of species was compiled, following the classification of Michener (2000). The dis tribution of species abundance was fitted to a truncated lognormal model (Pielou, 1975;

Magurran, 1988). A graph representing the accumulated number of species per unit of col

lecting effort (hours of collecting/person) was constructed.

Bee phenology

Information on the presence of each genus is presented for every month between July 1996 and June 1997, except for Neolarra Ashmead, since this genus was not collected dur

ing that period.

Biogeographic affinities

Biogeographic affinities were determined at the generic level, based on existing infor mation on the geographic distribution of the genera examined (Michener, 1979; Ayala,

1988; Michener et al., 1994; Michener, 2000).

Results

Bee diversity, phenology, and biogeographic affinities

A total of 3487 specimens were collected (169 in March 1994-July 1996; 3318 in July 1996-June 1997), and 259 species were found, grouped in 69 genera and 7 families. Af ter the genus name, its biogeographic affinity is indicated by the following conventions:

COS = cosmopolitan; HOL = holartic; NW = New World; NEA = neartic; NEO =

neotrop ical; MES = mesoamerican; SON = sonoran; AMP =

amphitropical. Also, information on the months when bees of the genus were collected is provided. One asterisk after the

species name indicates a new record for the State of Puebla. Two asterisks indicate a new record for Mexico.

The expression "sp." after generic names indicates one of two situations: 1. the species was already known to the respective specialist, even though a valid description for it did

18 JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

not exist at the time of determination. 2. the species was recognized as a separate taxo

nomic unit, but neither could it be assigned to any of the species known to the specialist nor could it be recognized as a new species. Species for which valid specific epithets were

not assigned, were sequentially numbered within subgenera.

Within each subgenus, species names were ordered in the following manner: 1. valid

species names followed by author and description year, in alphabetic order; 2. new

species; and, 3. species that were not assigned specific epithets (see previous paragraph.)

COLLETIDAE

COLLETINAE

1. Collet?s Latreille (COS except Indo-Australia; January, February, March, August, Sep tember, October)

Collet?s algarobiae Cockerell 1900

Collet?s macconelli Metz 1910

Collet?s ?ff. panamensis Michener 1954

Collet?s wootoni Cockerell 1897*

Collet?s new species 1

Collet?s new species 2

Collet?s sp.

DlPHAGLOSSINAE

Caupolicanini

2. Caupolicana Spinola (AMP; September) Caupolicana {Caupolicana) new species 1

3. Crawfordapis Moure (MES; September)

Crawfordapis new species 1

Hylaeinae

4. Hylaeus Fabricius (COS; January, February, March, April, July, August, September, Oc

tober)

Hylaeus sp. 1

Hylaeus sp. 2

Hylaeus sp. 3

Hylaeus sp. 4

Hylaeus sp. 5

Hylaeus sp. 6

Hylaeus sp. 7

ANDRENIDAE

Andreninae

5. Andrena Fabricius (HOL + African; August, September, October, November) Andrena {Callandrena) bilimeki LaBerge 1967

Andrena {Callandrena) dreisbachorum LaBerge 1967*

Andrena {Callandrena) inculta LaBerge 1967*

Andrena {Callandrena) limatula LaBerge 1967*

Andrena {Callandrena) perpunctata LaBerge 1967

Andrena {Callandrena) new species 1


Andrena (Callandrena) new species 2

Andrena (Callandrena) new species 3 Andrena (Callandrena) new species 4 Andrena (Callandrena) new species 5

Andrena (Callandrena) new species 6

Panurginae

Protandrenini

6. Protandrena Cockerell (NW; August, September) Protandrena (Heterosarus) fuscitarsis Timberlake 1975*

Protandrena (Heterosarus) gracilis Timberlake 1975*

Protandrena (Heterosarus) parvulus (Friese 1916) Protandrena (Heterosarus) new species 1

Protandrena (Heterosarus) new species 2

Protandrena (Protandrena) ?ff. punctulata Timberlake 1976 Protandrena (Protandrena) new species 1

Protandrena (Protandrena) new species 2

Protandrena (Protandrena) new species 3

Protandrena (Protandrena) sp. 1

Protandrena (Protandrena) sp. 2

Protandrena (Pterosarus) bidentis (Cockerell 1986)* Protandrena (Pterosarus) ?ff. solidaginis Roberson 1893 Protandrena (Pterosarus) sp. 1

Protandrena (Pterosarus) sp. 2




7. Pseudopanurgus Cockerell (NEA + MES; July, September) Pseudopanurgus tomentosus Timberlake 1973

Pseudopanurgus new species 1

Perditini

8. Macrotera Smith (NEA; April, May, June, July, August, September, October, Decem

ber) Macrotera (Cockerellula) azteca Timberlake 1954 Macrotera (Cockerellula) new species 1 Macrotera (Macrotera) bicolor (Smith 1853) Macrotera (Macroteropsis) parkeri Timberlake 1980

9. Perdita Smith (SON; February, March, April, May, June, July, August, September, Oc

tober, November, December)

Perdita (Epimacrotera) new species 1

Perdita (Perdita) aculeata Timberlake 1968 Perdita (Perdita) annectens Timberlake 1968* Perdita (Perdita) aff. caerulescens Timberlake 1964 Perdita (Perdita) decemnotata Timberlake 1962 Perdita (Perdita) aff. luciae Cockerell 1899 Perdita (Perdita) maerens Timberlake 1964* Perdita (Perdita) maesta Timberlake 1964


Perdita {Perdita) marginata Timberlake 1964*

Perdita {Perdita) aff. mexicanorum Cockerell 1896

Perdita {Perdita) oaxacana Timberlake 1964

Perdita {Perdita) obscurifacies Timberlake 1968*

Perdita {Perdita) aff. pernitens Timberlake 1980

Perdita {Perdita) pueblana Timberlake 1964

Perdita {Perdita) spheralceae balteata Timberlake 1964

Perdita {Perdita) new species 1

Perdita {Perdita) new species 2

Perdita {Perdita) sp. 1



10. Calliopsis Smith (AMP; September)

Calliopsis {Calliopsima) kucalumea Shinn 1967

Oxaeinae

11. Protoxaea Cockerell and Porter (AMP; September, October) Protoxaea {Mesoxea) nigerrima (Friese 1912)*

HALICTIDAE

Halictinae

Halictini

12. Agapostemon Gu?rin-M?neville (NW; January, February, March, April, May, June,

July, August, September, October, November, December)

Agapostemon {Agapostemon) cockerelli Crawford 1901

Agapostemon {Agapostemon) sp. 1

Agapostemon {Agapostemon) sp. 2

13. Caenohalictus Cameron (NEO; February, July, September, November, December) Caenohalictus sp. 1

Caenohalictus sp. 2

Caenohalictus sp. 3

14. Halictus Latreille (HOL + African; January, February, April, June, July, August, Sep tember, October, November, December)

Halictus {Odontalictus) ligatus Say 1837*

Halictus {Seladonia) lutescens Friese 1921

Halictus {Seladonia) sp. 1

15. Lasioglossum Curtis (COS; January, February, March, April, May, June, July, August,

September, October, December)

Lasioglossum {Dialictus) sp. 1










Lasioglossum (Dialictus) sp. 10











Lasioglossum (Evylaeus) sp. 1



Lasioglossum (Lasioglossum) sp. 1






16. Sphecodes Latreille (COS; September)

Sp he code s sp. 1

Sphecodes sp. 2

AUGOCHLORINI

17. Augochlora Smith (NW; January, February, March, April, June, July, August, Sep tember, December)

Augochlora (Augochlora) nigrocyanea Cockerell 1897*

Augochlora (Augochlora) sp. 1

Augochlora (Augochlora) sp. 2

Augochlora (Oxystoglossella) sp. 1

18. Augochlorella Sandhouse (NW; January, February, April, July, August, September, Oc

tober, December)

Augochlorella neglectula (Cockerell 1897) 19. Augochloropsis Cockerell (NW; April, June, July, August, September, December)

Augochloropsis (Augochloropsis) ignita (Smith 1861)*

Augochloropsis (Paraugochloropsis) metallica (Fabricius 1793)

Augochloropsis (Paraugochloropsis) vesta (Smith 1853) 20. Neocorynura Schrottky (NEO; September)

Neocorynura (Neocorynura) discolor (Smith 1879)* 21. Pseudaugochlora Michener (NEO; January, March, August, September, December)

Pseudaugochlora gram?nea (Fabricius 1804)*

MELITTIDAE

Dasypodainae

Dasypodaini

22. Hesperapis Cockerell (NEA + African; September) Hesperapis (Disparapis) sp. 1


MEGACHILIDAE

Megachilinae

LlTHURGINI

23. Lithurgus Berthold (COS; March, April, May)

Lithurgus {Lithurgopsis) planifrons (Friese 1908)

Lithurgus {Lithurgopsis) new species 1

Osmiini

24. Ashmeadiella Cockerell (NEA; February, March, April, May, June, July, August, Sep tember, October)

Ashmeadiella {Ashmeadiella) new species 1

Ashmeadiella {Ashmeadiella) cactorum (Cockerell 1897)* 25. Atoposmia Cockerell (SON; December)

Atoposmia {Eremosmia) tequila (Peters 1972) 26. Heriades Spinola (COS except South America and Australia; April, May, June)

Heriades {Neotrypetes) new species 1



27. Osmia Panzer (HOL; September, October) Osmia {Diceratosmia) azteca Cresson 1878

Osmia {Diceratosmia) new species 1

Anthidiini

28. Anthidiellum Cockerell (COS; May, September) Anthidiellum {Anthidiellum) aff. apicale (Cresson 1878)

29. Anthidium Fabricius (COS except Australia; June, July, September, October) Anthidium {Anthidium) maculifrons Smith 1854

Anthidium {Anthidium) maculosum Cresson 1878*

30. Anthodioctes Holmberg (NEO; September) Anthodioctes {Anthodioctes) sp. 1

31. Dianthidium Cockerell (NEA; August, September) Dianthidium {Adanthidium) anophrys Griswold and Michener 1988

Dianthidium {Adanthidium) new species 1

Dianthidium {Dianthidium) new species 1

Dianthidium {Mechanthidium) macrurum (Cockerell 1913)* 32. Paranthidium Cockerell and Cockerell (NEA; September)

Paranthidium {Rapanthidium) new species 1

33. Stelis Panzer (COS except Australia; May, October) Stelis {Dolichostelis) costaricencis (Friese 1921)

34. Trachusa Panzer (COS except Australia; September, October) Trachusa {Ulanthidium) mitchelli Michener 1948

Megachilini

35. Coelioxys Latreille (COS; February, September, December)

Coelioxys {Acrocoelioxys) azteca Cresson 1878*


Coelioxys (Boreocoelioxys) pratti Crawford 1914

Coelioxys (Neocoelioxys) menthae Cockerell 1897

Coelioxys (Synocoelioxys) new species 1

36. Megachile Latreille(COS; January, February, March, April, May, June, July, August,

September, October, November, December)

Megachi

Megachi

Megachii

Megachi

Megachii

Megachi

Megachi

Megachi

Megachi

Megachi

Megachh

Megachi

Megachi

Megachi

Megachi

Megachii

Megachi Megachn

\e (Argyropile) aff. rossi Mitchell 1943

\e (Argyropile) aff. paralella Smith 1853

]e (Chelostomoides) chilopsidis Cockerell 1900* ]e (Chelostomoides) otomita Cresson 1878*

]e (Chelostomoides) aff. subexilis Cockerell 1903

]e (Chelostomoides) texensis Mitchell 1976 \e (Chelostomoides) new species 1

]e (Cressoniella) zapoteca Cresson 1878

]e (Leptorachis) petulans Cresson 1878*

'e (Leptorachis) new species 1

]e (Lit?megachile) lippiae Cockerell 1900*

]e (Pseudocentron) aff. inscita Mitchell 1930

\e (Sayapis) aff. frugalis Cresson 1872

]e (Say apis) frugalis p s eudofrugalis Mitchell 1937*

]e (Sayapis) inimica sayi Cresson 1878

]e (Sayapis) zaptlana Cresson 1878

}e (Ty lome gachile) toluca Cresson 1878 e (Xanthosarus) augustini Cockerell 1905**

APIDAE

Xylocopinae

Xylocopini

37. Xylocopa Latreille (COS except Australia; January, February, March, April, May, June,

July, August, September, October, November, December)

Xylocopa (Neoxylocopa) varipuncta Patton 1879

Xylocopa (Notoxylocopa) tabaniformis melanosoma O'Brien and Hurd 1965

Xylocopa (Stenoxylocopa) micheneri Hurd 1978

Xylocopa (Xylocopoides) cyanea Smith 1874

Ceratinini

38. Ceratina Latreille (COS; January, February, March, April, July, August, September, December)

Ceratina (Zadontomerus) eximia Smith 1862*

Ceratina (Zadontomerus) ignara Cresson 1878*

Ceratina (Zadontomerus) nautlana Cockerell 1897*

Ceratina (Zadontomerus) new species 1






NOMADINAE

NOMADINI

39. Nomada Scopoli (COS; August, September) Nomada sp. 1

Nomada sp. 2

Nomada sp. 3

Epeolini

40. Doeringiella Holmberg (HOL + NEO; January, July, August, September, October, No

vember)

Doeringiella {Triepeolus) sarothrinus (Cockerell 1929)**

Doeringiella {Triepeolus) sp. 1






41. Epeolus Latreille (HOL + NEO; August) Epeolus sp. 1

Ammobatoidini

42. Holcopasites Ashmead (NEA; September)

Holcopasites {Holcopasites) sp. 1

Neolarrini

43. Neolarra Ashmead (NEA; not collected during 1996-1997) Neolarra {Neolarra) aff. californica Michener 1939

Apinae

Exomalopsini

44. Anthophorula Cockerell (AMP; July)

Anthophorula {Anthophorula) interrupta (Timberlake 1980) 45. Exomalopsis Spinola (NW; January, February, July, August, September, October, No

vember)

Exomalopsis {Exomalopsis) similis modesta Timberlake 1980

Exomalopsis {Megomalopsis) frederici Cockerell 1914

Exomalopsis {Phanomalopsis) pueblana Timberlake 1980

Tetrapediini

46. Tetrapedia Klug (NEO; September)

Tetrapedia {Tetrapedia) new species 1

Emphorini

47. Ancyloscelis Latreille (NEO; October, November, December)

Ancyloscelis sp. 1


48. Diadasia Patton (AMP; February, March, April, May, July, August, September, Octo

ber, November, December)

Diadasia australis (Cresson 1878) Diadasia diminuta (Cresson 1878)* Diadasia friesei Cockerell 1898**

Diadasia rinconis Cockerell 1897

Diadasia tropicalis (Cockerell 1918)* Diadasia sp. 1

Diadasia sp. 2

49. Melitoma Lepeletier and Serville (NW; September, October, November, December) Melitoma marginella (Cresson 1872)* Melitoma monozonula (Cockerell 1949) Melitoma nudicauda (Cockerell 1949)

Eucerini

50. Eucera Scopoli (HOL; October) Eucera (Synhalonia) new species 1

51. Gaesischia Michener, LaBerge and Moure (AMP; February, September, November) Gaesischia (Gaesischiana) exul Michener, LaBerge and Moure 1955*

52. Melissodes Latreille (NW; February, April, May, June, July, August, September, Oc

tober, November, December)

Melissodes (Eumelissodes) confusa Cresson 1878

Melissodes (Eumelissodes) interrupta LaBerge 1961*

Melissodes (Eumelissodes) montana Cresson 1878

Melissodes (Eumelissodes) tristis Cockerell 1894

Melissodes (Melissodes) colliciata Cockerell 1910

Melissodes (Melissodes) gilensis crenata LaBerge 1956

Melissodes (Melissodes) tepaneca Cresson 1878

53. Peponapis Robertson (NW; July, August, September) Peponapis crassidentata (Cockerell 1949)

Peponapis new species 1

54. Svastra Holmberg (AMP; July, September) Svastra (Epimelissodes) albocollaris (Cockerell 1918) Svastra sp. 1

55. Syntrichalonia LaBerge (NEA; September) Syntrichalonia exquisita (Cresson 1878)

56. Tetraloniella Ashmead (HOL + African; August, September, October, November, De

cember) Tetraloniella (Loxoptilus) brevifellator (LaBerge 1957) Tetraloniella (Pectinapis) auricauda LaBerge 1970

Tetraloniella (Pectinapis) sp. 1

Tetraloniella (Tetraloniella) eriocarpi (Cockerell 1898) Tetraloniella (Tetraloniella) flagellicomis (Smith 1879)* Tetraloniella (Tetraloniella) perconcinna (Cockerell ?) Tetraloniella (Tetraloniella) new species 1


Tetraloniella {Tetraloniella) new species 2

Tetraloniella {Tetraloniella) new species 3

57. Thygater Holmberg (NEO; August)

Thy g?ter {Thygater) cockerelli (Crawford 1906)*

Thygater {Thygater) micheneri Urban 1967*

Anthophorini

58. Anthophora Latreille (HOL; April, May, July, August, September, October, November,

December)

Anthophora {Anthophoroides) californica Cresson 1869

Anthophora {Heliophila) squammulosa Dours 1864

Anthophora {Mystacanthophora) capistrata Cresson 1878

59. Deltoptila LaBerge and Michener (MES; September)

Deltoptila aurulentocaudata Dours 1869*

Deltoptila new species 1

60. Habropoda Smith (HOL; September, October)

Habropoda salviarum Cockerell 1898

Centridini

61. Centris Fabricius (NEO; January, February, March, April, May, June, July, August,

September, October, November, December) Centris {Acritocentris) albiceps Friese 1899*

Centris {Centris) adani Cockerell 1949

Centris {Centris) eisenii Fox 1899

Centris {Hemisiella) transversa P?rez 1905

Centris {Hemisiella) trigonoides Lepeletier 1841

Centris {Paracentris) atripes Mocsary 1899

Centris {Paracentris) mexicana Smith 1854*

Centris {Paracentris) nigrocaerulea Smith 1874*

Centris {Paracentris) zacateca Snelling 1966*

Ericrocidini

62. Ericrocis Cresson (SON; May, July, August, September) Ericrocis pintada Snelling and Zavortink 1985

63. Mesocheira Lepeletier and Serville (NEO; September) Mesocheira bicolor (Fabricius 1804)

64. Mesoplia Lepeletier (NEO; May)

Mesoplia {Mesoplia) rufipes (Perty 1833)*

Euglossini

65. Euglossa Latreille (NEO; January, March, May, December)

Euglossa {Euglossa) viridissima Friese 1899

66. Eulaema Lepeletier (NEO; November, December) Eulaema {Apeulema) poly chroma (Mocsary 1899)

Bombini

67. Bombus Latreille (HOL + NEO; May, June, September, October, November, December) Bombus {Fervidobombus) pennsylvanicus sonorus Say 1837


Bombus (Pyrobombus) ephippiatus Say 1837

Bombus (Psithyrus) intrudens (Smith 1861)

Meliponini

68. Plebeia Schwarz (NEO; January, February, March, May, June, July, August, Septem ber, October, December)

Plebeia (Plebeia) frontalis Friese 1911

Plebeia (Plebeia) mexica Ayala 1999

69. Trigonisca Moure (NEO; September) Trigonisca azteca Ayala 1999

Distribution of species abundance

The data set on species abundance distribution was fitted to a truncated log-normal model

(Magurran, 1988), X2 = 3.07, P = 0.93; d.f. = 8 and S* (expected number of species) = 305.2

Species accumulation curve

A collecting effort of 776 hours/person was used to collect the 259 species found in the

study. A species accumulation curve is presented in Fig. 1.

Discussion

The main objective of the study was to know the diversity of Apoidea found in the Val

ley of Zapotitl?n de las Salinas. The Valley contains a higher number of species than any other locality studied in Mexico (Ayala, 1988; Godinez, 1991; Roubik et al., 1991; Estrada,

1992; Fierros, 1998). This finding agrees with the prediction (Ayala et al., 1996) that the

Tehuac?n region would present a number of species intermediate between the xeric regions of the United States and the mesic areas of the Mexican Plateau. Also, the number of

species which are common to the deserts of Sonora and Chihuahua, on the one hand, and

Tehuac?n, on the other, will possibly be found to be low, indicating differences in the floris tic composition of the three regions, which has evolved in recent times, as a result of in

creasing local aridity (Axelrod, 1979). Of the 126 taxa recognized as species in this study, 46 are new records for the state of

Puebla, and 3 of these [Diadasia friesei Cockerell 1898, Megachile (Xanthosarus) augus tini Cockerell 1905, and Doeringiella (Triepeolus) sarothrinus (Cockerell 1929)] are new

records for Mexico, according to the list presented by Ayala et al. (1996). This new infor mation would put Puebla as the fifth ranking Mexican state in bee species richness, with 302 species, after Chihuahua, Sonora, Jalisco, and Baja California.

Due to the current state of taxonomic knowledge of the group, only 65% of the collected

specimens were determined to species. The main problems are in taxa for which there are

no revisions for Mexico, as it is the case for Hylaeus, Protandrena (Pterosarus), La

sioglossum and Doeringiella (Triepeolus) which, together, make up 18% of the species found in this study.

We found 44 species new to science (17% of the total). This percentage is higher than

expected, considering the prediction that the number of species of bees found in Mexico could increase by 11% (Ayala et al., 1996).

According to the truncated log-normal model of species abundance distribution, the to tal expected number of species is 305. This would mean that 85% of the total number of

species present in the study area have been collected. This situation is typical of one-year


300

250

2 200

s c 150

% 100

3

200 300 400 500 600 700

Collecting effort (hours/person) Figure 1. Curve of species accumulation after 776 hour/person of collecting effort.

800

collecting studies (Magurran, 1988). On the other hand, the structure of the species accu

mulation curve indicates that the phase of fastest species accumulation has been already

completed and that the curve is reaching the asymptote parallel to the x-axis. The method

ology used in the study did not permit collecting of matinal, crepuscular or nocturnal bees.

Also, parsivoltine species were not collected.

We recorded activity of bees all year round. There is a peak of activity between Sep tember and October, which may be related to the rain fall regime in the Valley of Zapoti tl?n. Many annual and perennial Asteraceae as well as several species of Salvia (Lami

aceae) bloom right after the main rain period starts. Asteraceae is the most diverse and

abundant plant family visited by bees in the Valley of Zapotitl?n (G?mez, 1999); so, it is

expectable that their blooming period will determine the greatest abundance of bees in the area. Four bee genera {Agapostemon, Megachile, Centris, and Xylocopa) are active all year

round. Agapostemon is a New World genus and the other three are speciose cosmopolitan

or neotropical genera. These genera may present both oligolectic and polylectic species which can alternate to make use of the different plants, blooming in succession along the

year. At the other end of the spectrum, we found 16 pollen collecting genera and 6 clep

toparasitic genera, in several families, which were taken only during one month of the year.

Thirteen of the pollen-collecting and three of the cleptoparasitic genera are active in Sep tember, reflecting the greatest availability of floral resources in the region.

Almost 22% of the genera collected are of cosmopolitan affinity. Of the rest, 20% are

of neotropical affinity, 13% are of neartic affinity. These proportions reflect the main bio

geographic elements that determine most of the biological diversity of Mexico, and more

specifically, of the Transverse Volcanic Belt, where the study area is located. Other geo

graphical affinities well represented in the bee fauna of Zapotitl?n are holartic (15%), New

World (13%), and amphitropical elements. The scarce representation of genera (3,

4.4%)and the small number of species of sonoran affinity may possibly indicate that these


faunistic elements have recently penetrated into the Tehuac?n area, even though am

phitropical elements, which, according to Michener (2000) are primarily of Sonoran and

Chichuahuan origin, are not rare (10%) in the bee fauna of the area studied. Only two (3%) mesoamerican genera are present.

Acknowledgements

We want to thank Terry W. Griswold (Bee Biology and Systematics Laboratory,

USDA/ARS-University of Utah, Logan, Utah), Wallace LaBerge (Illinois Natural History Survey), and Michael S. Engel and Robert W. Brooks (Snow Entomological Division, Nat ural History Museum, University of Kansas, Lawrence, Kansas) for the identification of

specimens sent to them. Rhonda and Terry Griswold and Robert W. Brooks participated in the collection of specimens in the field. Mariana Cuautle helped with field work for the

entire collecting season of 1996-1997 and Martha Lopezaraiza helped solving some lo

gistic problems during the late stage of preparation of the manuscript. Two anonymous re

viewers gave useful suggestions to improve the manuscript.

Funding was provided by the Comisi?n Nacional para el Conocimiento y Uso de la

Biodiversidad (CONABIO) to project H278 'Apoidea (Hymenoptera) del Valle de Za

potitl?n de las Salinas, Puebla', and by the Consejo Nacional de Ciencia y Tecnolog?a (CONACyT), to project 2635-PN 'Diversidad, fenolog?a y flora visitada por apoideos del

Valle de Zapotitl?n de las Salinas, (Puebla)'. The Dean office of the Instituto de Investi

gaci?n y Posgrado (INIP) at U. de las Am?ricas-Puebla, provided funding and logistic support.

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