You are on page 1of 16

Diversity, Phenology and Biogeography of the Bees (Hymenoptera: Apoidea) of Zapotitln de las Salinas, Puebla, Mexico Author(s): Carlos

H. Vergara and Ricardo Ayala Source: Journal of the Kansas Entomological Society, Vol. 75, No. 1 (Jan., 2002), pp. 16-30 Published by: Allen Press on behalf of Kansas (Central States) Entomological Society Stable URL: http://www.jstor.org/stable/25086035 . Accessed: 21/02/2011 13:18
Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at . http://www.jstor.org/action/showPublisher?publisherCode=kes. . Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.

Kansas (Central States) Entomological Society and Allen Press are collaborating with JSTOR to digitize, preserve and extend access to Journal of the Kansas Entomological Society.

http://www.jstor.org

JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY


75(1), 2002, pp. 16-30

and Biogeography Phenology Diversity, of Zapotitl?n Apoidea) (Hymenoptera:


Puebla,
Carlos H. Vergara1

of the Bees de las Salinas,

Mexico
and Ricardo Ayala2

abstract: canic Belt

A faunistic

of central Mexico

The great and, of these, 3 are also new records for Mexico. level, was found for Apidae, with 33 genera and 92 species and, at the Curtis with 29 species, and Perdita Smith, with 20 species. The level, for Lasioglossum generic greatest monthly diversity was observed during September, when 57 genera were active. The main elements found, at the generic level, are of cosmopolitan affinity (22%), neotropical biogeographic that the Tehuac?n affinity (20%) and of neartic affinity (13%). Our results agree with the prediction est diversity, at the familial desert would mesic present a number of species areas of the Mexican Plateau. Also, intermediate the number between the xeric regions of the US and the to the deserts of Sonora of species common on the other, is low, reflecting differences in the Apoidea, diversity

total of 259 species, are new records for the state of Puebla

in the Transverse Neovol study of the bees (Apoidea) of a semi-desert was carried out by systematic monthly during one year. A collecting, 69 genera and seven families were recorded; 44 are undescribed species, 46

on the one hand, and Tehuac?n, and Chihuahua, floristic composition of the three regions. key words: bees, Hymenoptera, Biogeography,

Introduction Mexico
144 genera

has a rich fauna of bees, consisting


and seven families. This number

of 1800 species and subspecies


could rise to over 2000, once

grouped
several

in

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, recorded extratropical in deserts regions and of the world. The greatest regional number but of species from has the been corre shrublands of California and Arizona, records

sponding adjacent regions of Mexico


ico such as the Tehuac?n-Cuicatl?n

are lacking (Michener, 2000). Other deserts of Mex


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). inMexico have been carried out in Chamela, Faunistic studies on bee communities 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
Study area

and Materials

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 de las Am?ricas-Puebla, de Qu?mica y Biolog?a, Universidad Departamento 72820 Mexico. cvergara@mail.udlap.mx 2 Nacional Instituto de Biolog?a, Universidad Estaci?n de Biolog?a Chamela, Postal 21, San Patricio, Jalisco, 48980 M?xico. barajas@servidor.unam.mx

Santa Catarina M?rtir, Aut?noma de M?xico,

Puebla,

Apartado

VOLUME 75, ISSUE 1 97?34'W. The valley


system.

17

is drained by the Salado River, which

is part of the Papaloapan

station of Zapotitl?n Salinas, is dry with The climate, measured at the meteorological 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: tetetzo (A. Weber ex. K. Schum) Backeb (Cac thorny shrub, tetechera [Neobuxbaumia 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 de las Am?ricas-Puebla, Collection, Universidad
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; 1988). A graph representing the accumulated number of species per unit of col Magurran, was constructed. effort (hours of collecting/person) lecting Bee phenology Information on the presence of each genus is presented for every month between 1996 and June 1997, except for Neolarra Ashmead, since this genus was not collected ing that period. Biogeographic affinities July dur

affinities were determined at the generic level, based on existing infor Biogeographic 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 inMarch 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 SON = sonoran; AMP = amphitropical. Also, information on ical; MES = mesoamerican; 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 The for Mexico. 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.

each subgenus, species names were ordered in the following manner: 1. valid names followed by author and description in alphabetic order; 2. new year, species species; and, 3. species that were not assigned specific epithets (see previous paragraph.) Within

COLLETIDAE
COLLETINAE

1. Collet?s

(COS except Indo-Australia; tember, October) Collet?s algarobiae Cockerell 1900 Collet?s macconelli Metz 1910 1954 Collet?s ?ff. panamensis Michener 1897* Collet?s wootoni Cockerell Collet?s new species 1 Collet?s new species 2
Collet?s sp. DlPHAGLOSSINAE Caupolicanini

Latreille

January, February, March, August,

Sep

2. Caupolicana Caupolicana 3. Crawfordapis Crawfordapis

Spinola (AMP; September) {Caupolicana) new species Moure (MES; September) new species 1

Hylaeinae

4. Hylaeus tober)
Hylaeus Hylaeus Hylaeus Hylaeus Hylaeus

Fabricius
sp. sp. sp. 1 2 3

(COS; January, February, March, April,

July, August,

September, Oc

sp. 4 sp. 5

Hylaeus
Hylaeus

sp. 6
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

VOLUME 75, ISSUE 1 Andrena Andrena Andrena Andrena Andrena new new new new new 2 3 4 5 6
Panurginae Protandrenini

19

(Callandrena) (Callandrena) (Callandrena) (Callandrena) (Callandrena)

species species species species species

6. Protandrena Protandrena Protandrena Protandrena


Protandrena Protandrena

Cockerell

(NW; August, September) (Heterosarus) fuscitarsis Timberlake 1975* 1975* (Heterosarus) gracilis Timberlake (Heterosarus) parvulus (Friese 1916)
(Heterosarus) (Heterosarus) new new species species 1 2

Protandrena
Protandrena Protandrena Protandrena Protandrena

(Protandrena)
(Protandrena) (Protandrena) (Protandrena) (Protandrena)

?ff. punctulata
new new new sp. 1 species species species 1 2 3

Timberlake

1976

Protandrena Protandrena Protandrena


Protandrena Protandrena Protandrena Protandrena Protandrena

(Protandrena) sp. 2 (Pterosarus) bidentis (Cockerell 1986)* 1893 (Pterosarus) ?ff. solidaginis Roberson
(Pterosarus) (Pterosarus) (Pterosarus) (Pterosarus) (Pterosarus) sp. sp. sp. 1 2 3

sp. 4 sp. 5

7. Pseudopanurgus Pseudopanurgus
Pseudopanurgus

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


new species 1

Perditini (NEA; April, May, June, July, August, September, October, Decem ber) Macrotera 1954 (Cockerellula) azteca Timberlake new species 1 Macrotera (Cockerellula) Macrotera (Macrotera) bicolor (Smith 1853) Macrotera 1980 (Macroteropsis) parkeri Timberlake 9. Perdita Smith (SON; February, March, April, May, June, July, August, September, Oc
tober, November, December)

8. Macrotera

Smith

Perdita Perdita Perdita Perdita Perdita Perdita Perdita Perdita

(Epimacrotera) new species 1 (Perdita) aculeata Timberlake 1968 1968* (Perdita) annectens Timberlake 1964 (Perdita) aff. caerulescens Timberlake (Perdita) decemnotata Timberlake 1962 1899 (Perdita) aff. luciae Cockerell 1964* (Perdita) maerens Timberlake (Perdita) maesta Timberlake 1964

20

JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

Perdita {Perdita) marginata Timberlake 1964* Perdita {Perdita) aff. mexicanorum Cockerell 1896 1964 Perdita {Perdita) oaxacana Timberlake Perdita {Perdita) obscurifacies Timberlake 1968* Perdita {Perdita) aff. pernitens Timberlake 1980 Perdita {Perdita) pueblana Timberlake 1964 1964 Perdita {Perdita) spheralceae balteata Timberlake Perdita {Perdita) new species 1 Perdita {Perdita) new species 2 Perdita {Perdita) sp. 1 Perdita {Perdita) sp. 2 Perdita {Perdita) sp. 3 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) sp. 1 Agapostemon Agapostemon {Agapostemon) sp. 2 Cameron (NEO; February, July, September, November, December) 13. Caenohalictus
Caenohalictus Caenohalictus Caenohalictus sp. sp. sp. 1 2 3

14. Halictus
tember,

Latreille
October,

(HOL + African;
November,

January, February, April,

June, July, August,

Sep

December)

Halictus {Odontalictus) ligatus Say 1837* Halictus {Seladonia) lutescens Friese 1921 Halictus {Seladonia) sp. 1 15. Lasioglossum Curtis (COS; January, February, March, April, May, September, October, December) {Dialictus) sp. 1 Lasioglossum Lasioglossum Lasioglossum Lasioglossum Lasioglossum Lasioglossum Lasioglossum Lasioglossum Lasioglossum {Dialictus) {Dialictus) {Dialictus) {Dialictus) {Dialictus) {Dialictus) {Dialictus) {Dialictus) sp. sp. sp. sp. sp. sp. sp. sp. 2 3 4 5 6 7 8 9

June, July, August,

VOLUME

75,

ISSUE

21

(Dialictus) sp. 10 Lasioglossum (Dialictus) sp. 11 Lasioglossum (Dialictus) sp. 12 Lasioglossum (Dialictus) sp. 13 Lasioglossum (Dialictus) sp. 14 Lasioglossum (Dialictus) sp. 15 Lasioglossum (Dialictus) sp. 16 Lasioglossum (Dialictus) sp. 17 Lasioglossum (Dialictus) sp. 18 Lasioglossum (Dialictus) sp. 19 Lasioglossum (Dialictus) sp. 20 Lasioglossum (Evylaeus) sp. 1 Lasioglossum Lasioglossum (Evylaeus) sp. 2 (Evylaeus) sp. 3 Lasioglossum sp. 1 Lasioglossum (Lasioglossum) sp. 2 Lasioglossum (Lasioglossum) sp. 3 Lasioglossum (Lasioglossum) sp. 4 Lasioglossum (Lasioglossum) sp. 5 Lasioglossum (Lasioglossum) sp. 6 Lasioglossum (Lasioglossum) 16. Sphecodes Latreille (COS; September) Sp he code s sp. 1 Sphecodes sp. 2
AUGOCHLORINI

17. Augochlora
tember,

Smith

(NW; January, February, March, Cockerell

April,

June, July, August,

Sep

December)

Augochlora Augochlora Augochlora Augochlora


18. Augochlorella

(Augochlora) nigrocyanea (Augochlora) sp. 1 (Augochlora) sp. 2 sp. 1 (Oxystoglossella)


Sandhouse (NW; January,

1897*

February,

April,

July, August,

September,

Oc

tober, December) Augochlorella neglectula (Cockerell 1897) 19. Augochloropsis Cockerell (NW; April, June, July, August, September, December) Augochloropsis ignita (Smith 1861)* (Augochloropsis) metallica (Fabricius 1793) (Paraugochloropsis) Augochloropsis vesta (Smith 1853) (Paraugochloropsis) Augochloropsis 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 Hesperapis

(NEA + African; (Disparapis) sp. 1 Cockerell

September)

22

JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

MEGACHILIDAE
Megachilinae

LlTHURGINI 23. Lithurgus Berthold (COS; March, April, May) Lithurgus {Lithurgopsis) planifrons (Friese 1908) Lithurgus {Lithurgopsis) new species 1
Osmiini

Cockerell (NEA; February, March, April, May, June, July, August, tember, October) Ashmeadiella {Ashmeadiella) new species 1 Ashmeadiella {Ashmeadiella) cactorum (Cockerell 1897)* 25. Atoposmia Cockerell (SON; December) {Eremosmia) tequila (Peters 1972) Atoposmia 26. Heriades Spinola (COS except South America and Australia; April, May, June) Heriades {Neotrypetes) new species 1 Heriades {Neotrypetes) new species 2 Heriades {Neotrypetes) new species 3 27. Osmia Panzer (HOL; September, October) Osmia {Diceratosmia) azteca Cresson 1878 Osmia {Diceratosmia) new species 1
Anthidiini

24. Ashmeadiella

Sep

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) 1988 Dianthidium {Adanthidium) anophrys Griswold and Michener new species 1 Dianthidium {Adanthidium) Dianthidium {Dianthidium) new species 1 Dianthidium {Mechanthidium) macrurum (Cockerell 1913)* Cockerell and Cockerell (NEA; September) Paranthidium {Rapanthidium) new species 1 33. Stelis Panzer (COS except Australia; May, October) costaricencis Stelis {Dolichostelis) (Friese 1921) 34. Trachusa Panzer (COS except Australia; September, October) 1948 Trachusa {Ulanthidium) mitchelli Michener 32. Paranthidium
Megachilini

35. Coelioxys Latreille (COS; February, September, December) {Acrocoelioxys) azteca Cresson 1878* Coelioxys

VOLUME 75, ISSUE 1 Coelioxys (Boreocoelioxys) pratti Crawford 1914 1897 Coelioxys (Neocoelioxys) menthae Cockerell new species 1 Coelioxys (Synocoelioxys) 36. Megachile Latreille(COS; January, February, March, April, May, September, October, November, December) \e (Argyropile) aff. rossi Mitchell 1943 Megachi \e (Argyropile) aff. paralella Smith 1853 Megachi 1900* Megachii ]e (Chelostomoides) chilopsidis Cockerell ]e (Chelostomoides) otomita Cresson 1878* Megachi ]e (Chelostomoides) aff. subexilis Cockerell 1903 Megachii texensis Mitchell ]e (Chelostomoides) 1976 Megachi new species 1 \e (Chelostomoides) Megachi ]e (Cressoniella) zapoteca Cresson 1878 Megachi ]e (Leptorachis) petulans Cresson 1878* Megachi 'e (Leptorachis) new species 1 Megachi 1900* Megachh ]e (Lit?megachile) lippiae Cockerell aff. inscita Mitchell ]e (Pseudocentron) 1930 Megachi \e (Sayapis) aff. frugalis Cresson 1872 Megachi ]e (Say apis) frugalis p s eudofrugalis Mitchell 1937* Megachi ]e (Sayapis) inimica sayi Cresson 1878 Megachi Megachii ]e (Sayapis) zaptlana Cresson 1878 }e (Ty lome gachile) toluca Cresson 1878 Megachi e (Xanthosarus) augustini Cockerell 1905** Megachn

23

June, July, August,

APIDAE
Xylocopinae Xylocopini

37. Xylocopa

(COS except Australia; January, February, March, April, May, July, August, September, October, November, December) Xylocopa (Neoxylocopa) varipuncta Patton 1879 Xylocopa (Notoxylocopa) tabaniformis melanosoma O'Brien and Hurd 1965 (Stenoxylocopa) micheneri Hurd 1978 Xylocopa Xylocopa (Xylocopoides) cyanea Smith 1874
Ceratinini

Latreille

June,

38. Ceratina

Latreille (COS; January, February, March, April, December) Ceratina (Zadontomerus) eximia Smith 1862* Ceratina Ceratina Ceratina Ceratina Ceratina Ceratina Ceratina (Zadontomerus) (Zadontomerus) (Zadontomerus) (Zadontomerus) (Zadontomerus) (Zadontomerus) (Zadontomerus) ignara Cresson 1878* nautlana Cockerell 1897* new species 1 new species 2 new species 3 new species 4 new species 5

July, August,

September,

24

JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

NOMADINAE NOMADINI

39. Nomada
Nomada Nomada Nomada

Scopoli
sp. sp. sp. 1 2 3

(COS; August,

September)

Epeolini

40. Doeringiella vember)

Holmberg

(HOL + NEO;

January, July, August, 1929)**

September, October, No

Doeringiella {Triepeolus) Doeringiella {Triepeolus) Doeringiella {Triepeolus) Doeringiella {Triepeolus) Doeringiella {Triepeolus) Doeringiella {Triepeolus) Doeringiella {Triepeolus) 41. Epeolus Latreille (HOL
Epeolus sp. 1

sarothrinus (Cockerell sp. 1 sp. 2 sp. 3 sp. 4 sp. 5 sp. 6 + NEO; August)

Ammobatoidini

42. Holcopasites Holcopasites

Ashmead

{Holcopasites)

(NEA; September) sp. 1


Neolarrini

43. Neolarra Ashmead Neolarra {Neolarra)

(NEA; not collected during 1996-1997) aff. californica Michener 1939


Apinae Exomalopsini

44. Anthophorula Cockerell (AMP; July) {Anthophorula) interrupta (Timberlake 1980) Anthophorula 45. Exomalopsis Spinola (NW; January, February, July, August, September, October, No vember) 1980 {Exomalopsis) similis modesta Timberlake Exomalopsis Exomalopsis Exomalopsis 1914 {Megomalopsis) frederici Cockerell 1980 {Phanomalopsis) pueblana Timberlake
Tetrapediini

46. Tetrapedia Klug (NEO; September) Tetrapedia {Tetrapedia) new species

1
Emphorini

47. Ancyloscelis Ancyloscelis

Latreille sp. 1

(NEO; October, November,

December)

VOLUME

75,

ISSUE

25

48. Diadasia Diadasia Diadasia Diadasia Diadasia Diadasia


Diadasia Diadasia

Patton (AMP; February, March, April, May,


December)

July, August,

September, Octo

ber, November,

australis (Cresson 1878) diminuta (Cresson 1878)* 1898** friesei Cockerell rinconis Cockerell 1897 tropicalis
sp. 1 sp. 2

(Cockerell

1918)*

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

December)

Eucerini

50. Eucera Scopoli (HOL; October) Eucera (Synhalonia) new species 1 51. Gaesischia Michener, LaBerge and Moure (AMP; February, September, November) Gaesischia exul Michener, LaBerge and Moure 1955* (Gaesischiana) 52. Melissodes Latreille (NW; February, April, May, June, July, August, September, Oc
tober, November, December)

Melissodes Melissodes Melissodes Melissodes Melissodes Melissodes Melissodes 53. Peponapis Peponapis
Peponapis

(Eumelissodes) confusa Cresson 1878 (Eumelissodes) interrupta LaBerge 1961* montana Cresson 1878 (Eumelissodes) tristis Cockerell 1894 (Eumelissodes) colliciata Cockerell 1910 (Melissodes) crenata LaBerge 1956 (Melissodes) gilensis (Melissodes) tepaneca Cresson 1878 Robertson (NW; July, August, September) crassidentata (Cockerell 1949)
new species 1

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

1918)

55. Syntrichalonia LaBerge (NEA; September) Syntrichalonia exquisita (Cresson 1878) 56. Tetraloniella Ashmead (HOL + African; August, cember) Tetraloniella Tetraloniella Tetraloniella Tetraloniella Tetraloniella Tetraloniella Tetraloniella

September, October, November,

De

(Loxoptilus) brevifellator (LaBerge 1957) (Pectinapis) auricauda LaBerge 1970 (Pectinapis) sp. 1 (Tetraloniella) eriocarpi (Cockerell 1898) (Tetraloniella) flagellicomis (Smith 1879)* (Tetraloniella) perconcinna (Cockerell ?) (Tetraloniella) new species 1

26

JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

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 December)

Latreille

(HOL; April, May,

July, August,

September, October, November,

Anthophora {Anthophoroides) californica Cresson 1869 Anthophora {Heliophila) squammulosa Dours 1864 capistrata Cresson 1878 Anthophora {Mystacanthophora) 59. Deltoptila LaBerge and Michener (MES; September) aurulentocaudata Dours 1869* Deltoptila new species 1 Deltoptila 60. Habropoda Smith (HOL; September, October) salviarum Cockerell 1898 Habropoda
Centridini

61. Centris

(NEO; January, February, March, April, May, 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 1841 Centris {Hemisiella) trigonoides Lepeletier Centris {Paracentris) atripes Mocsary 1899 Centris {Paracentris) mexicana Smith 1854* Smith 1874* Centris {Paracentris) nigrocaerulea Centris {Paracentris) zacateca Snelling 1966*
Ericrocidini

Fabricius

June, July, August,

62. Ericrocis Cresson (SON; May, July, August, September) Ericrocis pintada Snelling and Zavortink 1985 63. Mesocheira Lepeletier and Serville (NEO; September) bicolor (Fabricius 1804) Mesocheira 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, sonorus Say 1837 Bombus {Fervidobombus) pennsylvanicus December)

VOLUME 75, ISSUE 1 Bombus Bombus (Pyrobombus) ephippiatus Say 1837 (Psithyrus) intrudens (Smith 1861)
Meliponini

27

68. Plebeia

(NEO; January, February, March, May, ber, October, December) Plebeia (Plebeia) frontalis Friese 1911 Plebeia (Plebeia) mexica Ayala 1999 69. Trigonisca Moure (NEO; September) Trigonisca azteca Ayala 1999 of species abundance

Schwarz

June, July, August,

Septem

Distribution

The data set on species abundance distribution was fitted to a truncated log-normal model = 3.07, P = 0.93; d.f. = 8 and S* = 305.2 (expected number of species) (Magurran, 1988), X2
Species accumulation curve

A collecting
study. A species

effort of 776 hours/person


accumulation curve

was used to collect


in Fig. 1.

the 259 species found in the

is presented

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


tic composition of the three regions, as which has evolved in recent times, as

in the floris
a result of in

creasing
Of the

local aridity (Axelrod,


126 taxa recognized

1979).
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

as it is the case for Hylaeus, Protandrena for Mexico, (Pterosarus), La and Doeringiella which, together, make up 18% of the species sioglossum (Triepeolus) 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 inMexico could increase by 11% (Ayala et al., 1996). to the truncated log-normal model of species abundance distribution, the to According 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

28

JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

300 250

2 200 s 150 c % 100 3


200
Figure 1. Curve of species

300
accumulation

400

500

600
of collecting

700
effort.

800

Collecting effort (hours/person)


after 776 hour/person

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 aceae) bloom annual right and after perennial the main Asteraceae rain period as well starts. as several species of Salvia diverse (Lami and Asteraceae is the most

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. round. Four bee genera Agapostemon {Agapostemon, is a New World Megachile, genus and Centris, the other and Xylocopa) three are are speciose active all year 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

and three of the cleptoparasitic genera are active in Sep of the pollen-collecting the greatest availability of floral resources in the region. tember, reflecting 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 and more geographic elements that determine most of the biological diversity of Mexico, of the Transverse Volcanic Belt, where the study area is located. Other geo specifically, graphical affinities well represented in the bee fauna of Zapotitl?n are holartic (15%), New of genera (3, elements. The scarce representation (13%), and amphitropical the small number of species of sonoran affinity may possibly indicate that these 4.4%)and

World

VOLUME 75, ISSUE 1

29

faunistic

elements

have

recently

penetrated

into

the

Tehuac?n

area,

even

though

am

(2000) are primarily of Sonoran and phitropical elements, which, according toMichener Chichuahuan origin, are not rare (10%) in the bee fauna of the area studied. Only two (3%)
mesoamerican genera are present.

Acknowledgements to thank Terry W. Griswold and Systematics (Bee Biology Laboratory, of Utah, Logan, Utah), Wallace LaBerge (Illinois Natural History USDA/ARS-University 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 We want
viewers gave useful suggestions to improve the manuscript.

y Uso de la Funding was provided by the Comisi?n Nacional para el Conocimiento to project H278 Biodiversidad (CONABIO) 'Apoidea (Hymenoptera) del Valle de Za potitl?n de las Salinas, Puebla', and by the Consejo Nacional de Ciencia y Tecnolog?a 'Diversidad, fenolog?a y flora visitada por apoideos del (CONACyT), to project 2635-PN 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.

Literature
D.

Cited

I. 1979. Age and origin of Sonoran Desert vegetation. Occasional Axelrod, Papers of the California Acad 132:1-74. emy of Sciences silvestres de Chamela, Folia Entomol?gica Mexicana 77:395-493. Jalisco, M?xico. Ayala, R. 1988. Abejas and S. H. Bullock. 1993. The native bees of Mexico. In T. P. Ramamoorthy, R. Bye, Ayala, R., T. L. Griswold, and J. Fa (eds.). Biological Press, New York. University and D. Yanega. R., T. L. Griswold, A. Lot, Diversity of Mexico, Origins and Distribution, pp. 179-227. Oxford

Ayala,

1996. Apoidea In J. Llorente-Bousquets, A. N. Gar (Hymenoptera). and E. Gonz?lez-Soriano taxonom?a y biogeograf?a de artr?podos de cia-Aldrete, (eds.). Biodiversidad, M?xico: Hacia una s?ntesis de su conocimiento, de M?xico, pp. 423^164. Universidad Nacional Aut?noma M?xico D. F. de Licen Folia En

Estrada,

C. M. 1992. Abejas Silvestres de la Sierra del Tigre, Jalisco. Tesis (Hymenoptera: Apoidea) ciatura en Biolog?a. Facultad de Ciencias, Universidad de Guadalajara, Guadalajara. del Volc?n de Tequila, Jalisco, M?xico. Fierros, H. F. 1998. Abejas silvestres (Hymenoptera: Apoidea) 102:21-70. tomol?gica Mexicana

G?mez,

S. 1999. Grado de especificidad floral de abejas que visitan las Aster?ceas del Valle de Zapotitl?n de en Biolog?a. Departamento las Salinas, Puebla. Tesis de Licenciatura de Qu?mica y Biolog?a, Universi dad de las Am?ricas-Puebla, Cholula.

al Sistema de Clasificaci?n Clim?tica de Koppen. Instituto Garc?a, E. 1973. Modificaciones Segunda Edici?n. de Geograf?a, Universidad Nacional Aut?noma de M?xico, M?xico D. F 1991. Algunos Aspectos de la Fenolog?a de las Abejas Silvestres God?nez, G. L. M. (Hymenoptera: Apoidea) de San Gregorio, Guanajuato. Tesis para obtener el t?tulo de Bi?logo, Facultad de Ciencias, Universidad Nacional Aut?noma de M?xico, M?xico, D. F. and Its Measurement. Croom Helm, London. Magurrran, A. E. 1988. Ecological Diversity C. D. 1979. Biogeography of the bees. Annals of the Missouri Botanical Garden 66:277-347. Michener, C. D. 2000. The Bees of theWorld. Johns Hopkins University Michener, Press, Baltimore, Maryland. C. D., R. J.McGinley, and B. Danforth. 1994. The Bee Genera of North and Central America. Michener, sonian Pielou, E C. Institution Press, Washington, Diversity. D.C. New York, New York. 1975. Ecological Wiley,

Smith

30

JOURNAL OF THE KANSAS ENTOMOLOGICAL SOCIETY

and classification bees 1993. Studies of the phylogeny of long-tongued A., and C. D. Michener. of Kansas Science Bulletin 55:123-162. University (Hymenoptera: Apoidea). de la Reserva de la E. E Cabrera-Cano and W. Colli-Ucan. 1991. Abejas Nativas Roubik, D., R. Villanueva, de la Biosfera Biosfera de la Reserva de de Sian Ka'an. In L. D. Navarro y G. John (eds.). Diversidad Sian Ka'an, Quintana Roo, Mexico, pp 317-320. CIQRO, Chetumal, Quintana Roo. Roig-Alsina, de M?xico. Editorial Limusa, M?xico, D.E J. 1981. Vegetaci?n Rzedowski, en el valle semi?rido de Zapotitl?n, 1982. Estudios Zavala, J. A. ecol?gicos basada en atributos binarios de presencia o ausencia de la vegetaci?n, Puebla. I. Clasificaci?n Bi?tica num?rica 7:99-120.

de las especies.