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Wilson, E.O. 2003 Pheidole in the New World: A Dominant, Hyperdiverse Ant Genus.

Wilson, E.O. 2003 Pheidole in the New World: A Dominant, Hyperdiverse Ant Genus Harvard University Press, Cambridge, 794 p + CD ROM. ISBN 0-674-00293. Price US$110.00. Review with a special emphasis on the Brazilian Pheidole ant fauna and megadiversity distribution in American countries

Completing his academic career the way he began, one of the most outstanding scientists of this century, Edward O. Wilson, published his formidable monographic revision of the New World Pheidole. Though formally retired since 1997, Wilson continues to labor as Emeritus Pellegrino University Research Professor of the Museum of Comparative Zoology of Harvard University. Dr. Wilson is well known to biologists and ecologists as the author of Sociobiology, Biophilia, Island Biogeography, among others. Wilson, in spite of this breadth of vision and synthesis, is the foremost living myrmecologist. To those who know him, the sheer passion that he shares for ants he extends to all other living beings, and it is this commitment which has highlighted his Biodiversity advocacy, and he is now one of its principle spokesmen. His monumental revisionary monograph of Pheidole follows last year's publication of The Future of Life (Wilson 2002), the most accessible and impassioned of his appeals to humans to take heed of the millions of other species that make the planet habitable.

Pheidole in the New World is Wilson's masterful revision of the species rich ant genus that he has scrutinized for 16 years, as known to all friends and myrmecologists. To deal with the enormous number of species, Wilson defines the term "Hyperdiversity" to describe the species-richness of Pheidole. Hyperdiversity is employed by Wilson to discuss "when a genus or family ... contains exceptionally large number of species with reference to plant and animal diversity as a whole." The large number of species is to be expected as Pheidole is the dominant taxon of New World ant ground ant assemblages (Fowler 1994), but some species are also arboreal and others have symbiotic relationships with myrmecophytes, Indeed, Pheidole is now the most species-diverse group of New World organisms after beetles (Chrysomelidae and Curculionidae). Placed in another light, Levi (2002), Wilson's colleague at the Museum of Comparative Zoology, estimates 1500 species of araenid spiders divided into 65 genera, 47 in Brazil, as occurring in the New World and these data show how hyperdiverse is the genus Pheidole.

The origin of these omnipresent faunal elements is still unclear. Pheidole has a world-wide distribution, although it is apparently dominant in the New World (Wilson 1976, Fowler 1994). The oldest fossil is from Oligocene shales of North America (Carpenter 1930), and is not common in the Dominician ambers (Baroni Urbani 1995). 2N chromosome numbers of the studied species range in the neighborhood of 20, although an Indian species has a diploid number of 42, not allowing for a clear interpretation. Old World diversification is however much lower than that found in the New World. Sister groups are presently interpreted as being Aphaenogaster, or the Indonesian Chimaeridris. Thus, much more research is needed to clarify the origin of this highly successful genus.

Wilson recognizes 624 Pheidole species in the Western Hemisphere. However, he notes that "undescribed species ... were still pouring in to the Harvard Collection as (this) monograph went to press" suggesting that many more species undoubtedly exist. This monumental binary key revision was in part supported by the world's taxonomists, which provided type specimens, the oldest dating from 1826. Wilson's research depended upon local collaborators who shipped him samples from collections throughout the United States, Latin America, and Europe, resulting in 337 new species. But, how does one organize such a monumental venture? Wilson began by dividing the taxa into 19 species groups, 2 of which are of Old World origin (megacephala and teneriffana both from Africa) but represented in the New World. One species group includes social parasitic forms, most of which are from Argentina and should grow as more myrmecological research continues in the New World.

Pheidole is spread through 35 New World countries, but the number of species, as well as the relative percentage of all species, varies greatly (Table 1). How does Brazil fit into the distribution of New World Pheidole? Is the mega-diversity of Brazil (Mittermeier and Werner 1990) also found in this highly speciose group? Indeed, the documented Brazilian Pheidole fauna represents 24.3% of its total diversity (152 species), followed by Mexico 18.6% (116 species), Costa Rica 18.1% (113 species), Colombia 15.2% (95 species), Pena 12.5% (78 species) and the USA 12.2% (76 species) (Fig. 1). These values, however, only consider the absolute number of species for each country. Furthermore, the superficial area and the collecting effort varies greatly among countries. In order to minimize differences in the number of species per area of any country, we created an index to estimate the relative mega-diversity of species of New World Pheidole, by dividing the total number of recorded species per country by the number of 10[grados] latitudinal bands of each country from 50[grados] N to 50[grados] S. The highest relative megadiversity of species of Pheidole was found in Costa Rica, followed by Panama, Pena, Mexico, Colombia, Brazil, Ecuador, EUA, Trinidad, and Guatemala independently of their surface area (Fig. 2, Table 1). These data clearly show the high collecting effort in Costa Rica and Panama, two countries well studied by Wilson. Brazil's sixth position indicates that this country may exhibit a large Pheidole diversity, depending on increased collecting effort.

In Brazil, of the 19 species groups of Pheidole recognized by Wilson, only four arc not present. The species groups crassicornis, granulata and pilifera are almost exclusively Nearctic. Additionally, the species group teneriffana, which is native to North Africa, has turned up in disturbed habitats in Cuba, Pena and California. Of the 15 species groups of Pheidole found in Brazil, tristis, diligens, fallax And flavens are, respectively, the most speciose and encompass 84% of the fauna. These species groups also represent the highest number of New World species, with flavens being the most species rich, followed by tristis, fallax and diligens (Fig. 3).

Of the 152 Brazilian species of Pheidole, 45.4% are apparently endemic to Brazil. Of these, 71% are known only from type localities, suggesting a broader geographic distribution, due to lack of systematic collections, or conversely restricted distributions which could lead to their rapid extinction without ever knowing their natural history. Of the total documented Brazilian fauna, 42.1% have unknown biologies, or in another words, 65.2% of the Brazilian species are endemic and with unknown biologies (Table 2), suggesting a formidable amount of natural history still ahead, as well as the probable discovery of more species. In spite of this fact, no Pheidole appear in the Red List of Threatened Species for Brazil (MMA 2003), which surprisingly has only three ant species listed.

Wilson graciously recognized many prominent myrmecologists who have worked largely solitarily in Latin America, by naming species after them, including the masters W.L. Brown and N. Kusnezov, and the present generation of myrmecologists J. Trager, W. MacKay, J. Lattke, C.R.F. Brandao, P. Oliveira, and H.G. Fowler. Wilson recognized the importance of the contributions in support of tropical conservation by Harrison Ford, Peter A. Seligmann and Sal Roush. Indeed, Wilson dedicated the book to his deceased mentor, W.L. Brown, who began the revision of this genus with Wilson more than 20 years ago (Brown 1981).

Wilson meticulously illustrated by hand all species, and showed another innovation (Fig. 4). Wilson introduced anatomical markers which highlight keyed taxonomic characters and should greatly assist new myrmecologists, as well as more experienced ones by making explicit identifying traits.

A CD accompanies the printed book. The CD micrographically illustrates the species and should facilitate production of local and regional faunal keys. Combined with work underway at the Museum of Comparative Zoology and elsewhere, this additional innovation assures that such painstaking research will henceforth be refined and accelerated using digital imaging and genomics techniques, and then promptly disseminated to users worldwide by the Internet. In scholarly terms, it bridges the established science of biological systematics and its imminent future. However, the CD contains a mistake concerning the total number of Pheidole species in Brazil. Instead of 150 species presented in the CD list, we have detected that the book shows 152 species for Brazil, including P laevinota (diligens group) and P. nitidicollis (flavens group).


The authors are grateful to Cynthia P.A. Prado for the valuable suggestions on the manuscript and Thiago S. de Azevedo for the help with the figures.


Baroni Urbani, C. 1995. Invasion and extinction in the West Indian ant fauna revisited: the example of Pheidole (Amber Collection Stutgart Hymenoptera: Formicidae, VII: Myrmicinae, partim). Stutg. Beitr. Naturkd. Ser. B. (Geol. Palaontol.) 62: 1-10.

Brown, W.L. Jr. 1981. Preliminary contributions toward a revision of the ant genus Pheidole (Hymenoptera: Formicidae), Part I. J. Kansas Entomol. Soc. 54: 523-530.

Carpenter, F.M. 1930. The fossil ants of North America. Bull. Mus. Comp. Zool. 70: 1-66.

Fowler, H.G. 1994. Relative representation of Pheidole (Hymenoptera: Formicidae) in local ground ant assemblages of the Americas. An. Biol. 19: 29-37.

Levi, H.W. 2002. Keys to the genera of araneid orbweavers (Araneae, Araneidae) of the Americas. J. Arachnol. 30: 527-562.

Ministerio do Meio Ambiente, IBAMA, Brazil. 2003. Lista de especies ameacadas de extincao.

Mittermeier, R.A. & T.B. Werner. 1990. Wealth of plants and animals unites 'megadiversity' countries. Tropicus 4: 4-5.

Pauwels, G. J. 1997. Atlas geografico melhoramentos. Melhoramentos, Sao Paulo. 80 p.

Wilson, E.O. 1976. Which are the most prevalent ant genera? Stud. Entomol. 19:187-200.

Wilson, E.O. 2002. The Future of Life. Harvard University, Cambridge.

F.J. Zara (1) & H.G. Fowler (2)

(1) UNESP, Campus do Litoral Paulista--Unidade de Sao Vicente, CEP 11330-900, Sao Vicente, SP, Brazil; fjzara@csv.

(2) Depto de Ecologia UNESP, Rio Claro--C. P. 199, CEP 13506-900, Rio Claro, SP, Brazil;

Statistics of species richness and country characteristics used in
our analysis

                                  Number of                 Relative
                                  latitude                    mega-
                                  bands of                  diversity
                                 10[grados]    Country       Index
                                  covering       area       Species/
                      Species       each       ([Km.sup.    latitude
      Country         richness     country       2]) *     10[grados]

Argentina                59           4        2.766.889      14.75
Brazil                  152           5        8.511.965      30.6
Uruguay                   4           1         186.925        4
Chile                     1           5         751.625        0.2
Paraguay                  9           2         406.750        4.5
Bolivia                  34           2        1.098.575      17
Peru                     78           2        1.285.215      39
Ecuador                  53           2         461.475       26.5
Colombia                 95           3        1.138.915      31.7
Venezuela                31           2         912.045       15.5
French Guiana             7           1         91.000         7
Suriname                  8           1         163.820        8
Guyana                   13           1         214.970       13
Trinidad                 23           1          5.130        23
Curacao (Dutch            1           1           993          1
  West Indies)
Panama                   46           1         78.515        46
Costa Rica              113           2         50.900        56.5
Nicaragua                11           1         148.000       11
Honduras                 19           1         112.085       19
El Salvador              15           1         21.395        15
Guatemala                20           1         108.890       20
Belize                    4           1         22.965         4
Bermudas                  1           1           54           1
Cuba                     12           2         114.525        6
Jamaica                   7           1         11.425         7
Haiti                     4           1         27.750         4
Domenican Republic        7           1         48.440         7
Dominica                 10           1           751         10
Puerto Rico               8           1          8.960         8
St. Kitts and Nevis       1           1           262          1
Bahamas                   4           1         13.865         4
St. Vincent and
The Grenadines            7           1           389          7
Grenada                   4           1           345          4
USA                      76           3        9.363.130      25.3
Mexico                  116           3        1.972.545      38.7

(*) from Pauwels (1997)


Synoptic listing of what is known of the biologies and distributions of
recorded Brazilian species of Pheidole

                                           Known      only in
      Group               Species         biology      Brazil

aberrans             aberrans               Yes          No
                     cavifrons              Yes          No
                     fracticeps             Yes          No
                     minensis               No          Yes
biconstricta         biconstricta           Yes          No
                     simplex                Yes          No
                     socrates               No           No
crassiconnis         ausency                in         Brazil
diligens             angusta                No          Yes
                     blumenauensis          No          Yes
                     bruensi                Yes          No
                     comptostela            No          Yes
                     cataractae             Yes          No
                     chrysops               Yes          No
                     coffeicola             No           No
                     cyrtostela             Yes         Yes
                     diligens               Yes          No
                     embolopyx              Yes          No
                     fowleri                Yes         Yes
                     geraesensis            No          Yes
                     laevifrons             No          Yes
                     laevinota              Yes          No
                     laidlowi               No           No
                     lancifer               Yes          No
                     lemur                  No          Yes
                     longiscapa             Yes          No
                     longiseta              Yes          No
                     oxyops                 Yes          No
                     peregrina              Yes         Yes
                     perryorum              Yes         Yes
                     pubiventris            Yes          No
                     radoszkowskii          Yes          No
                     reichenspergeri        No          Yes
                     rochai                 No          Yes
                     rufipilis              No          Yes
                     sensitiva              Yes          No
                     strigosa               Yes         Yes
                     triconstricta          Yes          No
                     vafra                  Yes          No
                     veletis                Yes          No
distorta             bufo                   Yes          No
                     deima                  No          Yes
                     dolon                  Yes          No
                     monstrosa              No          Yes
fallax               aenescens              No          Yes
                     alienata               No          Yes
                     araenoides             Yes         Yes
                     arcifera               No          Yes
                     cardinalis             Yes         Yes
                     claviscapa             Yes          No
                     cuevasi                Yes          No
fallax               cursor                 Yes          No
                     eidmanni               No          Yes
                     gigas                  Yes         Yes
                     impressa               No          Yes
                     jacutifera             No          Yes
flavens              flavens                Yes          No
                     flavida                No          Yes
                     goeldii                No          Yes
                     guajirana              Yes          No
                     ligniocola             No           No
                     lucaris                Yes         Yes
                     minutula               Yes          No
                     nana                   No          Yes
                     nitella                Yes          No
                     nitidicollis           No           No
                     obtusopilosa           Yes          No
                     peltastes              No          Yes
                     pholeops               Yes          No
                     rudigenis              Yes          No
                     schmalzi               No           No
                     sospes                 Yes          No
                     subreticulata          No          Yes
                     tambopatae             Yes          No
                     termitobia             Yes         Yes
                     tetrica                No          Yes
                     victma                 No          Yes
gertrudae            capillata              Yes          No
                     gertrudae              No           No
                     jeannei                Yes         Yes
granulata            ausency                in         Brazil
lamia                colbopsis              Yes          No
megacephala          megacephala            Yes          No
perpusilla           perpusilla             Yes          No
pilifera             crusency               in         Brazil
pucntatissima        meinerti               Yes          No
                     rugiceps               Yes          No
scrobifera           mamore                 Yes          No
tachigaliae          mendicula              Yes          No
                     pedana                 No           No
teneriffana          ausency                in         Brazil
transversostriata    scolioceps             Yes          No
                     transversostriata      Yes          No
tristis              alexeter               Yes         Yes
                     allarmata              Yes          No
                     alpinensis             No          Yes
                     ambigua                No          Yes
                     aper                   No          Yes
                     auropilosa             No           No
                     avia                   No          Yes
                     bambusarum             Yes          No
                     brevicona              No          Yes
                     brunnescens            No          Yes
                     bucculenta             Yes         Yes
                     carapuna               Yes          No
                     cephalica              Yes          No
                     cramptoni              Yes          No
Tristis              descolei               No           No
                     dyctiota               Yes         Yes
                     fabricator             Yes          No
                     fera                   No          Yes
                     fimbriata              Yes          No
                     jelskii                Yes          No
                     jujuvensis             No           No
                     lovejoyi               Yes         Yes
                     lucretii               No          Yes
                     midas                  Yes          No
                     nesiota                No          Yes
                     nitidula               No           No
                     obscurior              Yes          No
                     paraensis              No          Yes
                     punctithorax           No          Yes
                     puttemansi             No           No
                     tijucana               No          Yes
                     trageri                No          Yes
                     valens                 Yes         Yes
                     wallacei               No          Yes
                     wolfringi              No          Yes
flavens              arhuaca                Yes          No
                     asperithorax           Yes          No
                     bidens                 No          Yes
                     borgmeieri             Yes         Yes
                     dinophica              Yes         Yes
                     exigua                 Yes          No
                     germaini               No          Yes
                     gibba                  No          Yes
                     grandinodus            No          Yes
                     guilelmimuelleri       No           No
                     hetschkoi              Yes         Yes
                     lutzi                  Yes         Yes
                     moseni                 Yes         Yes
                     polita                 Yes          No
                     praeses                Yes         Yes
                     risii                  No           No
                     rosae                  Yes          No
                     rutilana               Yes         Yes
                     sarcina                No           No
                     schwarzmaieri          No           No
                     senilis                Yes         Yes
                     sigillata              Yes         Yes
                     spininodis             Yes          No
                     stulta                 Yes          No
                     subarmata              No           No
                     synarmata              Yes          No
                     tristicula             Yes          No
                     tristis                No           No

Fig. 1. The 15 New World countries with the largest number of species of
Pheidole. Numbers above bars indicate the relative percentage of each
country to the total number of species recognized by Wilson (2003).

Guyana        2
Honduras      3
Guatemala     3.3
Trinidad      3.7
Venezuela     5
Bolivia       5.5
Panama        7.4
Ecuador       8.5
Argentina     9.5
USA          12.2
Peru         12.5
Colombia     15.2
C. Rica      18.1
Mexico       18.6
Brazil       24.5

Nota: Tabla derivada de grafico de barra.

Fig. 2. Relative Pheidole megadiversity (number of species/number of
10[grados] latitudinal bands from 50[grados]N to 50[grados]N to
50[grados]S) for the 15 most speciose countries of the New World.

Guyana       13
Argentina    14.7
Venezuela    15.5
Bolivia      17
Honduras     19
Guatemala    20
Trinidad     23
USA          25.3
Ecuador      26.5
Brazil       30.6
Colombia     31.7
Mexico       38.7
Peru         39
Panama       46
C. Rica      56.5

Nota: Tabla derivada de grafico de barra.

Fig. 3. Species richness of each 15 species groups of Pheidole found in
Brazil (white bars) compared to the total New World species group
(black bars).

                     Pheidole found
                       in Brazil       Total New World species group

tristis                   132                           41
transversostriata           5                            2
tachigalie                  5                            2
scrobifera                 12                            1
punctatissima              12                            2
perpusila                   4                            1
megacephala                 1                            1
lamia                       4                            1
gertrudae                   3                            3
flavens                   165                           27
fallax                    103                           28
distorta                    7                            4
diligens                   86                           32
bicontricta                 6                            3
aberrans                   12                            4

Nota: Tabla derivada de grafico de barra.
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Author:Zara, F.J.; Fowler, H.G.
Publication:Revista de Biologia Tropical
Article Type:Resena de libro
Date:Mar 1, 2005
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