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A monograph of the genus Pheidole in Florida (Hymenoptera : Formicidae)

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Title:
A monograph of the genus Pheidole in Florida (Hymenoptera : Formicidae)
Creator:
Naves, Marcio Antonio, 1943-
Publication Date:
Language:
English
Physical Description:
xiii, 105 leaves : ill. ; 28 cm.

Subjects

Subjects / Keywords:
Ants ( jstor )
Hair ( jstor )
Head ( jstor )
Occipital lobe ( jstor )
Paratypes ( jstor )
Pedicels ( jstor )
Petioles ( jstor )
Species ( jstor )
Thorax ( jstor )
Weather fronts ( jstor )
Ants -- Florida ( lcsh )
Dissertations, Academic -- Entomology and Nematology -- UF
Entomology and Nematology thesis Ph. D
Pheidole ( lcsh )
City of Gainesville ( local )
Genre:
bibliography ( marcgt )
non-fiction ( marcgt )

Notes

Thesis:
Thesis (Ph. D.)--University of Florida, 1976.
Bibliography:
Bibliography: leaves 103-104.
General Note:
Typescript.
General Note:
Vita.
Statement of Responsibility:
by Marcio Antonio Naves.

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University of Florida
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A MONOGRAPH OF THE GENUS PHEIDOLE IN FLORIDA (HYMENOPTERA:FORMICIDAE)

















By

Marcio A. Naves


















A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF THE
UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 1976





































To my parents, wife and son and in memory of

Father Walter W. Kempf, O.F.M.















ACKNOWLEDGEMENTS

I wish to express my gratitude to Dr. William F. Buren, Advisory

Committee Chairman, who patiently guided me and inspired me during every phase of this research. I also wish to acknowledge my sincere indebteness to Dr. George E. Allen, Dr. Francis W. Zettler and Dr. Clifford S. Lofgren, members of my Advisory Committee, for their constructive criticisms and guidance.

I sincerely thank Mrs. T. Carlysle for providing me with scanning electron micrographs.

I also wish to thank Dr. John F. Carroll, Dr. J. C. E. Nickerson, and Dr. D. P. Wojcik, for continuing assistance, and to Dr. Edward 0. Wilson and Dr. David R. Smith for their permission to study the collections at the Museum of Comparative Zoology and U.S. National Museum, respectively.

This research was also aided in part by Tall Timbers Research Station, Leon Co., Florida, through the kindness of Dr. E. V. Komarek, Director of the Station, and the sponsorship of Dr. Willard H. Whitcomb. The support of Empresa Brasileira de Pesquisa Agropecuaria is also acknowledged with gratitude.

To my wife, Lucilia, my utmost thanks and appreciation for her patience and support.













iii












TABLE OF CONTENTS

Page

ACKNOWLEDGMENTS....................... ......................... iii

LIST OF FIGURES ........ .......................................... viii

ABSTRACT..............................................................xii

INTRODUCTION..................................................... 1

REVIEW OF LITERATURE............................................. 3

MATERIALS AND METHODS ............................................. 5

KEY TO SPECIES OF PHEIDOLE OF FLORIDA (MAJORS ONLY).............. 8

KEY TO PHEIDOLE SPECIES OF FLORIDA (WORKERS ONLY) ................ 13

1. Pheidole adrianoi n. sp. ............................... 17
Diagnosis............................................. 17
Description........................................... 17
Major............................................... 17
Measurements...................................... 17
Structural characters ............................. 17
Sculpture......................................... 18
Pilosity.......................................... 19
Color............................................. 19
Worker.............................................. 19
Measurements...................................... 19
Structural characters ............................. 19
Sculpture......................................... 20
Pilosity ........ ................................. 20
Color............................................. 20
Female.............................................. 20
Measurements...................................... 20
Structural characters .............................. 20
Sculpture......................................... 21
Pilosity.......................................... 21
Color............................................. 22
Male................................................ 22
Holotype.......................................... 22
Paratypes......................................... 22
Discussion............................................ 22
2. Pheidole anastasii Emery ................................. 24
Diagnosis............................................. 24
Discussion............................................ 24







iv











TABLE OF CONTENTS Continued

Page

3. Pheidole bureni n. sp. ................................... 26
Diagnosis .............................................. 26
Description............................................ 26
Major ................................. .............. 26
Measurements............... .......................... 26
Structural characters............................... 26
Sculpture........................................... 27
Pilosity...............................................28
Color .............................................. 28
Worker.................................................. 28
Measurements ................ ...................... 28
Structural characters................................. 28
Sculpture.......................................... 29
Pilosity........................................... 29
Color.............................................. 30
Female............................................... 30
Measurements .......................................... 30
Structural characters.............................. 30
Sculpture.......................................... 30
Pilosity ........................................... 31
Color.............................................. 31
Male ......... ...................................... 31
Holotype........................................... 31
Paratypes.......................................... 31
Discussion.................................................32
4. Pheidole carrolli n. sp. ............................... 33
Diagnosis.............................................. 33
Description............................................ 33
Major ....... ....................................... 33
Measurements.. ...................................... 33
Structural characters.............................. 33
Sculpture .......... .............................. 34
Pilosity ............................................ 35
Color...................................................35
Worker............................................... 35
Measurements....................................... 35
Structural character ............................... 35
Sculpture ........... .............................. 36
Pilosity .... ...................................... 36
Color ............................................... 36
Holotype............................................. 36
Paratypes.......................................... 36
Discussion ...... ..................................... 37









V











TABLE OF CONTENTS Continued

Page

5.. Pheidole crassicornis Emery.............................. 39
Diagnosis .............................................. 39
Discussion.................................................39
6. Pheidole dentata Mayr..................................... 40
Diagnosis.............................................. 41
Discussion ............................................... 41
7. Pheidole dentigula M. R. Smith ........................... 43
Diagnosis.............................................. 43
Discussion............................................. 43
8. Pheidole diversipilosa Wheeler ............................ 44
Diagnosis.............................................. 44
Discussion............................................. 44
9. Pheidole fallax obscurithorax Santschi................... 46
Diagnosis.............................................. 46
Discussion............................................. 46
10. Pheidole floridana Emery ................................. 48
Diagnosis ................................................ 48
Discussion............................... .. 48
11. Pheidole greggi n. sp..................................... 49
Diagnosis.............................................. 49
Description ....... ................................... 49
Major................................................ 49
Measurements....................................... 49
Structural characters .............................. 49
Sculpture.......................................... 50
Pilosity........................................... 51
Color .............................................. 51
Worker............................................... 51
Measurements....................................... 51
Structural characters .............................. 51
Sculpture.......................................... 52
Pilosity ..................................... ....... 52
Color ..... ....................................... 52
Female and Male...................................... 52
Holotype ...... ................................... 52
Paratypes.......................................... 52
Discussion ....... .................................... 52
12. Pheidole lamia Wheeler................................... '55
Diagnosis.............................................. 55
Discussion............................................. 55
13. Pheidole littoralis Cole.................................. 57
Diagnosis............... ............................... 57
Discussion............................................. 57
14. Pheidole megacephala (Fabricius) ......................... 58
Diagnosis.............................................. 58
Discussion............................................. 59





vi











TABLE OF CONTENTS Continued

Page

15. Pheidole metallescens Emery ............................. 61
Diagnosis............................................ 61
Discussion ........................................... 61
16. Pheidole moerens Wheeler ............................... 62
Diagnosis............................................ 62
Discussion........................................... 62
17. Pheidole morrisi Forel.................................. 64
Diagnosis ........... .......... ..................... 64
Discussion........................................... 65
18. Pheidole auiescens n. sp................................ 66
Diagnosis............................................ 66
Description.......................................... 66
Major.............................................. 66
Worker........................................ 66
Measurements..................................... 66
Structural characters ............................ 66
Sculpture........................................ 67
Pilosity......................................... 67
Color............................................ 68
Female and male.................................... 68
Holotype......................................... 68
Paratypes ........................................ 68
Discussion....................................... 68
19. Pheidole tysoni Forel.................................. 71
Diagnosis............................................ 71
Discussion........................................... 71
20. Pheidole vinelandica Forel ............................. 72
Diagnosis ...... .................................... 73
Discussion........................................... 73

SPECIES NOT PROPERLY INCLUDED IN THE FLORIDA PHEIDOLE FAUNA .... 75

1. Pheidole pilifera (Roger)............................... 75
Diagnosis............................................ 75
Discussion........................................... 75

LITERATURE CITED ................................................ 103

BIOGRAPHICAL SKETCH............................................. 105








vii














LIST OF FIGURES

Figure Page

1 Head of major of P. lamia Wheeler (33X)................. 78

2 Scape of major of P. bureni n. sp. a member of the
crassicornis complex (81X) .............................. 78

3 Head of major of P. diversipilosa Wheeler (45X).......... 78

4 Gaster of major of P. diversipilosa (47X)................ 78

5 Head of major of P. crassicornis Emery (43X) ............ 78

6 Gaster of major of P. crassicornis (58X)................. 78

7 Scape of major of P. fallax obscurithorax Santschi
(81x) .................................................... 80

8 Head of major of P. fallax obscurithorax (40X).......... 80

9 Head of major of P. megacephala (Fabricius) (40X)....... 80 10 Pedicel of major of P. floridana Emery (128X) ........... 80

11 Propodeum, pedicel and gaster of major of P.
metallescens Emery (74X) ..................................... 80

12 Postpetiole of major of P. littoralis Cole (119X)....... 80 13 Head of major of P. dentigula M. R. Smith (64X)......... 82

14 Head of major of P. carrolli n. sp. (29X) .............. 82

15 Pedicel of major of P. metallescens Emery (163X)......... 82 16 Head of major of P. metallescens (62X) .................. 82

17 Head of major of P. anastasii Emery (68X) ............... 82

18 Postpetiole and gaster of P. anastasii (62X)............ 82

19 Head of major of P. floridana Emery (72X) ............... 84

20 Head of major of P. littoralis Cole (37X) ............... 84





viii











LIST OF FIGURES Continued

Figure Page 21 Head of major of P. greggi n. sp. (68X).................. 84

22 Head of major of P. moerens Wheeler (58X) ............... 84

23 Clypeus and frontal area of major of P. vinelandica Forel (186X) ............................................ 84

24 Head of major of P. bicarinata Mayr (53X) ............... 84

25 Head of major of P. adrianoi n. sp. (72X) ............... 86

26 Thorax, pedicel and gaster of major of P. adrianoi n. sp. (50X); white legs are artifacts due to electron
overcharge ........ ..................................... 86

27 Head of major of P. morrisi Forel (37X) ...... ........... 86

28 Thorax of major of P. morrisi (56X) ..................... 86

29 Head of major of P. dentata Mayr (48X) ............... 86

30 Teeth on the anterior ventral head margin of P.
anastasii (Emery (123X).................................. 86

31 Head of worker of P. lamia (97X) ........................ 88

32 Dorsal view of worker of P. vinelandica (62X) ........... 88

33 Dorsal view of worker of P. adrianoi n. sp. (95X)....... 88 34 Worker of P. meace hala (36X) .......................... 88

35 Thorax and pedicel of the worker of P. fallax
obscurithorax (41X) ....................................... 88

36 Head of worker of P. fallax obscurithorax (86X)......... 88 37 Thorax of the worker of P. morrisi (72X) ................ 90

38 Head thorax and pedicel of the worker of P. dentata
(72X)............. ...................... ............... 90

39 Thorax of the worker of P. dentata (79X) ................ 90






ix










LIST OF FIGURES Continued

Figure Page 40 Head of the worker of P. crassicornis (54X)............. 90

41 Thorax of the worker of P. metallescens (196X) ......... 90 42 Head of the worker of P. littoralis (113X) ............. 90

43 Worker of P. carrolii n. sp. (40X) ..................... 92

44 Head of the worker of P. moerens (102X) ................ 92

45 Head of worker of P. greggi n. sp. (102X)............... 92

46 Worker of P. quiescens n. sp. (45X) .................... 92

47 Dorsal view of worker of P. quiescens n. sp. (103X).... 92 48 Propodeum, pedicel, and gaster of worker of P.
dentigula (102X) ........................................... 92

49 Dorsal view of thorax of P. dentigul-a worker (115X).... 94 50 Head of worker of P. anastasii (109X) .................. 94

51 Head of worker of P. floridana (115X) .................. 94

52 Pedicel of worker of P. floridana (193X) ............... 94

53 Dorsal thoracic hairs of worker of P. adrianoi n. sp.
(1,401X and 5,174X, respectively) ...................... 94

54 Dorsal thoracic hairs of worker of P. adrianoi n. sp.
(1,401X and 5,174X, respectively) ...................... 94

55 Head of major of P. flavens (95X)........................ 96

56 Head of major of P. flavens (87X) ...................... 96

57 Occipital lobes of P. greggi n. sp. (250X) ............ 96

58 Postpetiole of the major of P. flavens sculptior (158X). 96 59 Head and thorax of the minor of P. flavens (83X)........ 96 60 Head of major of P. bureni n. sp. (39X)................. 98







X










LIST OF FIGURES Continued

Figure Page 61 Gaster and pedicel of P. bureni n. sp. (40X)............ 98

62 Head and pronotum of worker of P. bureni n. sp.
(52x).................................................. 96

63 Dorsum of propodeum of major of P. vinelandica (272X).................................................. 100

64 Thoracic dorsum of P. bicarinata (71X) .................. 100

65 Postpetiole of P. bicarinata (186X) ..................... 100

66 Head of P. vinelandica (66X)................... ........ 100

67 Map of Florida and the United States .................... 102









































xi












Abstract of Dissertation Presented to the Graduate Council
of the University of Florida in Partial Fulfillment of the Requirements
for the Degree of Doctor of Philosophy


A MONOGRAPH OF THE GENUS PHEIDOLE IN FLORIDA (HY.MENOPTERA:FORMICIDAE)

By

Marcio A. Naves

December, 1976

Chairman: William F. Buren
Major Department: Entomology and Nematology

Until recently only twelve species of Pheidole were reported from

Florida, P. anastasii Emery, P. bicarinata vinelandica Forel, P. dentigula M. R. Smith, P. flavens Roger, P. megacephala (Fabricius), P. moerens Wheeler, P. morrisi Forel, P. pilifera (Roger), P. dentata IMayr, P. floridana Emery, P. metallescens Emery and P. sitarches littoralis Cole. The last four have Florida as the type locality.

P. pilifera (Roger) is believed to have been errounbously reported from Florida and has been excluded from the Florida fauna. P. flavens (Gregg, 1958, nec Roger) is considered a misidentification and to be an undescribed species, P. greggi n. sp.

In addition to P. greggi n. sp. three additional new species of

Pheidole are described from Florida: P. adrianoi n. sp., P. carrolli n. sp., and P. 9.liescens n. sp. An exotic species, P. fallax obscurithorax (= P. fallax arenicola var. obscurithorax Santschi) probably introduced from South Aerica, is also added to the North American ant fauna.

PheidoIe cr-_ssicornis Emery ard P. diversip-ilosa Wheeler are recorded for the first time in Floridal. The latter species is resurrected from synomy1y .id is elevated to full species.


-4 t











Two additional taxa are elevated to full species: P. vinelandica (= P. bicarinata vinelandica Forel) and P. littoralis (= P. sitarches littoralis Cole). The P. crassicornis complex is recognized with the description of P. bureni n. sp. collected in Alice, Texas even though this species does not occur in Florida.

The four new species, P. adrianoi n. sp., P. bureni n. sp., P.

carrolii n. sp., and P. quiescens n. sp., plus the introduced species P. fallax obscurithorax Santschi and the recognized species P. diversipilosa Wheeler increase the total Pheidole fauna of North America north of Mexico to seventy-seven. The Florida Pheidole fauna is increased to nineteen species and Pheidole is now the largest ant genus in the state.

Keys for the identification of both majors and workers are provided. Scanning electron micrographs are also provided as identification aides. The biology of each species is discussed.



























xiii















INTRODUCTION

The genus Pheidole is one of the more important ant genera in

the world, Approximately 1000 taxa are known. Over 400 taxa are known from the Neotropical region (Kempf, 1972), and about 77 taxa are now known from North America north of Mexico. Pheidole spp. are abundant in many areas, and live in varying habitats ranging from the humid tropics to deserts. They are able to survive in some areas by their habits of collecting and storing seeds as food resources. They are also scavengers of dead insects and other animals, and can be predaceous. Some species tend aphids and other homopterans, but this food source, so important to many ants genera, is probauby of relatively minor importance to most

species of Pheidole.

Pheidole apparently is a genus of rather recent evolutionary development. No fossil Pheidole are known previous to the Miocene (Bromn, 1973). There are no Pheidole in the Baltic armber (lower Oiigocene) although these ambers contain large numbers of specimens of other ant genera which have persisted almost unchanged since that epoch (Formica, Myrmica, Iridomyrmex, Caamoopotus, and others). It has been said that Pheidole has undergone a world wide dispersal since this time and has undergone a large proliferetion of taxai as well as adaptation to many ecological nitches (Brown, 1973).

A study_ of the Pheidole of any area is one of the prerequisites to understanrdi.ng the fornicid interactions of that area and this in turn could be importaJnt in the de.voopmrent of pest management strategies, since




1










ants are probably the most numerous and most important animals within their size range.

One species, Pheidole megacephala, has become a tropicopolitan tramp species, and is a serious pest in various areas (Wheeler. 1910, Flucker and Beardsley, 1970 and Lieburg, 1975). Some species sting severely and are comparable with the fire ants in aggressiveness (Buren, personal communication). Most of the species, however, appear innocuous and are usually unnoticed by man. Their importance in the ecosystem appears largely unappreciated.

Due to its size and distribution, Pheidole is a very difficult genus and has been a challenge to myrmecologists taxonomically as well as biologically.

Until recently only eleven species were known from Florida, which suggested that despite its subtropical warmth Florida was deficient in Pheidole when compared to Texas (36 species) and Arizona (26 species).

The Florida Pheidole fauna has long been known to be more extensive than that of northern states such as Iowa and New Jersey (3 species each, Buren, 1944 and Creighton, 1950).

This study was undertaken to increase our knowledge of Pheidole

of Florida. Scanning electron micrographs are provided along with keys for the identification of both majors and workers.















REVIEW OF LITERATURE

Four out of twelve species reported from Florida have that state as

the type locality: P. dentata Mayr (1886), P. floridana Emery (1895),

P. metallescens Emery (1895), and P. sitarches litorallis, Cole (1952).

Smith (1930) reported P. morrisi; Wheeler (1932) reported P. anastasii;

Smith (I 33) reported P. megacephaia; Smith (1944) reported P. dentigula;

Smith (1951) reported P. flavens sculptior (misidentification); Van Pelt

(1956) reported P. pilifera; Smith (1958) reported P. bicarinata vinelandica,

and Wojcik et al. (1975) reported P. moerens. Additional information is

discussed under each species by the respective authors.

Creighton (1950) was the first to try to give a extensive key to the

North America Pheidole; 63 species were listed and he used morphological

characters pertinent to both majors and workers. Creighton's statements

about the North American Pheidole are as follows:

Most of our species of Pheidole posses a dimorphic worker caste
with major and minor workers not connected by intermediates.
In a few species, however, the worker caste is polymorphic (kingi,
instabilis and torpescens, vasliti arizonica, etc.). Most of
the species garner seeds and it is believed that the large-headed major workers function as seed-huskers. The enlarged head of the major is mainly filled with mandibular muscles. This enables the
jaws to exert much pressure, which should be useful in cracking off the husks of seeds. It may be added that sometimes the head
of the m joar is so large in proportion to its body that if the insect is turned over on the back of its head it cannot regain
a normal. posture without help from other workers. Despite their preference for a graminivourous diet many species of
Phe.dole will accept other food as well. They seem less attracted to honey-dew than do many ants but will often feed voraciously
on animal tissue when the opportunity offers. The majority of our species form small colonies. In many cases there are only







3






4


about two or three hundred individuals in a fully developed
nest. Even in the case of the species which produce comparatively
large nests (morrisi, hyatti, desertorum, etc.) a colony of more
than two or three thousand individuals would be exceptionally large. By far the majority of our species nest in soil. The
nest may be built under a stone or in open soil without a covering object. In the latter case there is often a mound or craters
of excavated soil surrounding the nest entrance. The eastern
species dentata will nest in rotten logs as well as soil but
such flexibility in nesting habit is exceptional.

(p. 162)

Gregg (1958), published a new revision in which he included the

description of ten new taxa. He also placed ten taxa in synomymy and the status of eight taxa was revised. Later, Gregg (1969) described another new species, P. clementensis.

Smith (1967) reported P. moerens from Alabama as yet another exotic

species. This was the 71st Pheidole taxon reported for North America north of Mexico. Wojcik et al. (1975) reported that this species was also present in Florida.















MATERIALS AND METHODS

Four species of Pheidole (P. dentigula, P. fallax obscuri.thorax, P. bureni, n. sp. and P. gregi n. sp.) included in this research were received from other collectors. All other specimens were collected by the author during several trips in the state (fig. 67) or while at Tall Timbers Research Station in Leon Co., Florida, during the summers of 1974 and 1975.

Colonies of Pheidole spp. were located in several ways:

a. Examinging cavities under the bark of trees or examining grass

behind the leaf bases of palms.

b. Searching for nest craters, mounds, or any other modification

on the soil surface likely to have been produced by ants.

c. Searching for foraging majors or workers and following them to

the nest location.

d. Baiting with honey or dead arthropods and following trails of

workers and majors to the nest location.

3. Overturning stones, boards, or pieces of wood.

The two most successful methods were examining cavities under the bark of pine trees and looking for craters on the soil surface.

Colonies were collected by aspirating the ants once the chambers were located. Nests in the soil were excavated by removal of large blocks of soil, breaking it carefully and aspirating the ants found. The depth explored was variable, ranging from 30 to 50 cm for P. metallescens to over 1.5 in for P. lamia.




5






6


The ants were placed in snap-cap vials with 70% alcohol for

preservation or kept alive in snap-cap vials with a piece of moistened cotton to ensure adequate survival. The snap-cap vials were placed in a styrofoam box to protect them against excessive changes in temperature during transport back to the laboratory.

The colonies were kept in chambers made from petri dishes modified

to assure proper high moisture levels and ease of feeding and observation without disturbing the colony.

Plastic petri dishes 9 cms in diameter were used. A 0.5 cm hole was melted by a soldering iron toward theside of the bottom of one dish. A piece of artificial cotton matting was pushed half way through the hole and was coated inside with a mixture of plaster of paris (98%) and commercial cement (2%). The matting below the hole was kept continuously wet with distilled water and was retained in another petri dish bottom. It did not need to be replenished with water frequently, The nest chamber was formed by applying the petri dish top and sealing with tape. A small feeding chamber which remained dry vwas formed by affixing another petri dish bottom to the perri dish top of the nest chamber with an interconnecting exit hole. This construction allowed colonies to be maintained in the laboratory for long periods with ouly infrequent attention. The dry feeding area separated from the nest chamber inhibited the growth of molds. Large numbers of colonies were thus 'ept in a small laboratory space.

Colonies were fed with fleshly killed house-flies, fresh peanuts, and honey.

Whenever possible field observations were made concerning the ecology and biology of each species.






7



A trip was taken to the Museum of Comparative Zoology, Harvard University, Cambridge, Massachussetts, and to the National Museum of Natural History in Washington, D.C., in order to study types of Nearctic and Neotropical Pheidole.

All specimens were point mounted and studied by using a stereo microscope with magnification up to 160X. The pictures were taken using the scanning electron microscope of the insects Attractants Laboratory, Agricultural Research Service, U.S.D.A., Gainesville, Florida.

Several measurements were taken for the newly described species including:

a. Body length: the sum of the head length, thoracic length, pedicel

and gastric lengths.

b. Head length: lengths of the head in full face view (mandibles

excluded).

c. Head width: greatest width of the head in full face view.

d. Thoracic length: greatest length of thorax in lateral view.

e. Scape length: middle of antennal socket to the scape tip.

f. Scape index: found by computing the formula: Scape length X 100 head length

The following abreviations were used throughout this work:

M.C.Z. Museum of Comparative Zoology, Harvard University, Cambridge,

Massachussetts.

U.S. Nat. Mus. United States National Museum, Washington, D.C.

A.M.N.. ;kierican Mascum of Natural History, New York, N.Y.














KEY TO SPECIES OF PHEIDOLE OF FLORIDA (MAJORS ONLY)

la) Head cylindrical in cross section and obliquely truncate

anteriorly, the truncation involving the clypeus, frontal

area, anterior portions of genae, and mandibles.

(fig. 1)................................................ lamia Wheeler

b) Head not cylindrical in cross section and not truncate anteriorly......................................................... 2

2a) Antennal scape strongly bent, flattened and smooth at the

base, the base nearly as or as broad as the distal portions

of the scape (fig. 2)............................................... 3

b) Not as above, scape not flattened at base and usually distinctly narrower than distal portion; if thickened

and strongly bent at base, obviously not flattened ................. 4

3a) First gastric tergite covered with very numerous long and

short erect hairs; sometimes sparse,coarse, appressed,

pubescence is present. (fig. 3, 4)............ diversipilosa Wheeler

b) First gastric tergite covered with appressed pubescence and sometimes with sparse long hairs.

(fig. 5, 6).........................................crassicornis Emery

4a) Large species, total length over 6 mm; head heavily

sculptured; scapes very thick and bent at base. The

scapes seen in full face view are evenly curved measally,

but more angular laterally (fig. 7, 8)..fallax obscurithorax Santschi



8











b) Species usually measuring 5 mm or less; if larger the head never heavily sculptured; the scapes never thick or

strongly bent at base.............................................. 5

5a) Head cordate, gradually but distinctly narrowed toward

the mandibular insertions; broadest at the occipital

lobes; mesonotum not transversely impressed.

(fig. 9)...................................... mpgacePala (Fabricius)

b) Head not cordate and never strongly narrowed towards the mandibular insertions; mesonotum may or may not be

transversely impressed ............................................. 6

6a) Postpetiole, seen from above, bearing distinct lateral

connules; head covered with reticulate punctation or

longitudinally striate. (fig. 10, 11).............................. 7

b) Postpetiole seen from above, without distinct lateral connules; if indistinct lateral projections present

the posterior half of the head smooth and shining.

(fig. 12).......................................................... 11

7a) Occipital lobes with reticulate sculpture, with no trace

of transverse or longitudinal striation, the surface

opaque or feebly shining; post-petiolar width two times

length. (fig. 13) ............................. dentigula M. R. Smith

b) Occipital lobes smooth and shining or with sparse

punctures or striae; postpetiole width greater than length,

but never two times length. (fig. 10, 11).......................... 8

8a) Sides of head sub-parallel, narrowing slightly anteriorly;

head length 1.82 to 2.0 mm; scape index 41 to 42; pronotum






10



and mesonotum strongly sculptured with transverse and

sometimes with longitudinal rugae; dorsal crest of petiole

notched. (fig. 14).................................... carrolli n. sp.

b) Without all the above characters.................................. 9

9a) Erect gastric hairs sparse, tapered and almost all of the same size; posterior part of the head covered with sculpture;

anterior ventral margin of the head without teeth; body

sometimes with distinct violaceous or bluish metallic

reflections. (fig. 15, 16)........................ metallescens Emery

b) Gaster ocvered with long and short hairs; posterior part

of the head smooth and shining (fig. 17); anterior ventral

margin of the head with two or three teeth, the mesal

tooth usually short and blunt. (fig. 30).......................... 10

10a) Dorsal surface of first gastric tergite finely and densely

granulose, opalescent and opaque; dorsal areas of pronotum

punctate and sometimes with several short and fine transverse

striae; dorsal area of post-petiole punctate.

(fig. 17, 18)....................................... anastasii Emery

b) Dorsal surface of first gastric tergite smooth and shining;

dorsum of pronotum partially weakly punctate but also

interdispersed with smooth and shining areas; dorsal area

of post-petiole weakly punctate and shining.

(fig. 10, 19)........................................ floridana Emery

lla) Head covered with longitudinal striae; the dorsal area of

the pronotum and usually the mesothorax reticulo-punctate

and with transverse striae or sculpture. (fig. 20)................ 12










b) Posterior areas of the head smooth and shining; the pronotum,

seen from above, smooth and very shining........................... 14

12a) Head usually measuring over 1.4 mm in length and 1.1 mm

in width; sides of the head sub-parallel; scape length

shorter than half the head length (Scape Index around 40);

occipital lobes with distinct circular piligerous punctures.

(fig. 12, 20) ...................................... littoralis Cole

b) Head measuring less than .9 mm in length and width; occipital

lobes smooth and shining............................................. 13

13a) Frontal area concave and bearing 1 or 2 longitudinal

striae. Scrobes strongly punctate and opaque; occipital lobes smooth and weakly shining; on occiput interstitial

spaces between striae reticulo-punctate. (fig. 21).... greggi n. sp.

b) Frontal area concave, without longitudinal striae; posterior

half of the scrobe weakly punctate and shining; occipital

lobes and most of the head shining; on front, interstitial

spaces between striae smooth and shining. (fig. 22). moerens Wheeler 14a) Anterior clypeal margin bluntly bidentate, with deep

median notch; anterior ventral margin of head without

teeth. (fig. 23) ................................. vinelandica Forel

b) Anterior clypeal margin not bidentate and without notch;

anterior ventral margin of the head usually with two or

three teeth; if teeth are absent, shining black species............ 15

15a) Mesonotum without transverse impression, forming in profile

a continuous smooth curve with the pronotum. (fig. 26) ........... 16






12




b) Mesonotum, with a well developed transverse impression............. 17

16a) A yellow species; frontal area and middle of front smooth

and shining without striae; anterior ventral margin with

3 teeth; thorax smooth and shining ...................... tysoni Forel

b) A black species; anterior half of head with longitudinal striae; dorsum of pronotum smooth and shining; pleurae

of pronotum, mesonotum, and propodeum.heavily sculptured.

(fig. 25, 26)......................................... adrianoi n. sp.

17a) Propodeum angular at the junction of the basal and declivous

faces, but the angles not produced into distinct teeth or

spines. (fig. 27, 28)................................ morrisi Forel

b) Propodeum armed with distinct teeth or spines.

(fig. 29).............................................. dentata Mayr
















KEY TO PHEIDOLE SPECIES OF FLORIDA (WORKERS ONLY)

la) Entire head smooth and shining. (fig. 31, 36)...................... 2

b) Head reticulo-punctate or partially reticulate with some

smooth areas. (fig. 40, 41, 44)................................... 9

2a) Scapes failing to reach the posterior border of the head,

or surpassing it by an amount no greater than the first

funicular joint. (fig. 31)........................................ 3

b) Scapes surpassing the posterior border of the head by an amount greater than the first funicular joint. (fig. 34).......... 6

3a) Thorax smooth, nearly free of sculpture and strongly

shining; yellow.................................................... 4

b) Pronotum smooth and shining; mesonotum and propodeum punctate or sculptured; color various. (fig. 32, 33) .............. 5

4a) Propodeum armed with angular teeth which are broad at the

base and do not resemble spines; mesonotum and propodeum

in profile separated by a deep, wide impression.

(fig. a ) .............................................. lamia Wheeler

b) Propodeum armed with spines; mesonotum and propodeum in profile separated by a normal impression ............... tysoni Forel

5a) Yellowish to brownish species, gaster covered with numerous

long fine hairs.(fig. 32) ......................... vinelandica Forel

b) Ebony black species, gaster with plumose or subplumose hairs which are spaced at distances more or less equal to

the hair's length. (fig. 33) ...................... adrianoi n. sp.



13






14



6a) Mesonotum without a distinct tranverse impression; in

profile postpetiole longer than petiole and dorsal and

ventral outlines both smoothly convex.

(fig. 34) ................................... megacephala (Fabricius)

b) The mesonotum with a distinct transverse impression.

(fig. 35, 37, 39).................................................. 7

7a) Posteriorly on head a distinct collar visible in full

face view (fig. 36); the mesonotum and base of propodeum

heavily punctuate. (fig. 35) ......... fallax obscurithorax Santschi

b) Posterior collar of head not visible in full face view; the mesonotal dorsum and propodeal base weakly sculptured

and shining ........................................................ 8

8a) Usually yellowish or pale species, the propodeal spines

absent. (fig. 37) ..................................... morrisi Forel

b) Dark brown to ferruginous species; propodeal spines well formed. (fig. 38, 39)................................. dentata Mayr

9a) Scapes surpassing the posterior margin of the head by a

distance greater than the first funicular joint. (fig. 40) ....... 10 b) Scapes either not surpassing posterior margin of the head, or surpassing less than the length of the first funicular

joint. (fig. 43)................................................. 11

10a) First gastric tergite with appressed pubescence and only a

few scattered long hairs ......................... crassicornis Emery

b) First gastric tergite covered with numerous long and short hairs; pubescence usually erect or semi-erect, rarely

appressed...................................... diversipilosa Wheeler





15



lla) Head not completely punctate, the front of the head with

irregular smooth and shining areas. (fig. 42).................... 12

b) In full face view, head entirely covered with reticulate punctation or with longitudinal striae. (fig. 43)................ 13

12a) Pleurae of pronotum heavily reticulate; head, thorax and

gaster usually with distinct violaceous or bluish metallic

reflections; eyes with less than 30 facets.

(fig. 41)......................................... metallescens Emery

b) Pleurae of pronotum smooth and shining; body without

violaceous or bluish reflections; eyes with more than

45 facets. (fig. 42)................................ littoralis Cole

13a) Eyes large, with more than 40 facets. (fig. 43)..... carrolli n. sp.

b) Eyes small, with less than 25 facets. (fig. 50, 51) .............. 14

14a) Postpetiole small, little wider than the petiole.

(fig. 44) ..... .................................... moerens Wheeler

or (fig. 4 ).. ......................................... greggi n. sp.

b) Postpetiole nearly or more than twice as wide as the

petiole. (fig. 47)............................................... 15

15a) Body and mainly the gaster covered with numerous fine

long hairs; gaster enlarged, larger than the thorax;

the propodeal spines very thick and obliquely curved and

pointed laterad; first gastric tergite opalescent and

subopaque. (fig. 46, 47) .......................... quiescens n. sp.

b) Not with this combination of characters ........................... 16






16




16a) Postpetiole without lateral angles, hairs slender and

pointed; postpetiole and gaster smooth and shining.

(fig. 48)...................................... dentigula M. R. Smith

b) Postpetiole with angular sides, seen from above; hairs short and thick................................................... 17

17a) Dorsal surface of first gastric tergite finely and densily

granulose, opalescent and subopaque; dorsum of pronotum

and mesonotum punctate; dorsum of postpetiole punctate

and opaque. (fig. 50)............................... anastasii Emery

b) First gastric tergite smooth and shining; dorsal area of

postpetiole weakly smooth and shining; dorsum of pronotum

usually with partial smooth shining areas.

(fig. 51). .......................................... floridana Emery











1. Pheidole adrianoi n. sp.

Diagnosis:

A small ebony black species differing from P. metallescens by the

absence of the violaceous or bluish reflections and without the reticulate sculptured occipital lobes. It also differs from P. littoralis by the small shining head, the head length usually less than 1.0 mm in the major as opposed to over 1.4 numm in littoralis; workers with the erect hairs plumose or semi-plumose at apex.

Description:

Ma or:

Measurements: Body length 2.70 to 3.10 mm. Head length 0.82

to 0.97 mm; head width 0.79 to 0.95 mm. Thorax length 0.79 to 0.90 mm;

scape length 0.45 to 0.51 mm. Scape Index 51 to 59.

Structural characters: Head shape as in fig. 25. Head, excluding mandibles, usually slightly longer than wide, broadest behind

the eyes from where the sides slightly curve and narrow posteriorly,

the occipital excision gently concave. Frontal area impressed; anterior

border of clypeus with a wide, shallow median notch. A weak, scarcely

depressed antennal semi-scrobe present; a distinct frontal furrow

arising behind the frontal area, widening posteriorly into the

occipital excision. Scapes slender at base, thickening apically and longer than the half the head length; mandibles stout, weakly curved laterally, with two apical blunt teeth and two basal denticles. Eyes

small with approximately 35 facets.

Pronotum and mesonotum strongly convex in profile, posteriorly

the mesonotum more abruptly descending into the mesopropodeal impression.



17






18



Mesonotum without transverse impression; spines thick and long but blunt apically and pointed obliquely dorsad. Petiole stout, anterior and posterior faces sloping, the crest weakly angular in profile.

Seen from above, the sides of post petiole semi-angular, the lateral projections blunt and not well developed; gaster truncate at base and smaller than head.

Sculpture: Strong cephalic longitudinal striae diverging

posteriorly and disappearing in the area posterior to the scape when in repose; interstitial areas between striae with weak, fine reticulate punctation; frontal carinae diverging and surpassing the scapes when in repose; vertex, frontal area as well as the clypeus smooth and shining; the semi-scrobes not distinct and marked only by lack of sculpture; the genae with strong longitudinal striae, the interstitial areas weakly reticulo-punctate; occipital lobes smooth and shining with sparse small circular piligerous punctures, ventral surface of head smooth and shining, its anterior margin without teeth; frontal furrow with its posterior half bearing transverse rugules.

Seen from above, dorsum of the pronotum and anterior mesonotum

nearly smooth and weakly shining, mesopropodeal impression and adjacent posterior area of mesonotum reticulo-punctate; basal area of propodeum weakly reticulate and shining; declivous face usually shining.

In profile, pronotum, mesonotum and propodeum heavily reticulate, the inner areas finely reticulo-punctate and shining; petiole finely reticulo-punctate. Seen from above, dorsum of pronotum and anterior mesonotum nearly smooth and weakly shining; mesopropodeal impression and adjacent posterior area of mesonotum reticulo-punctate; basal





19




area of propodeum reticulate and shining; declivous face with transverse striae and usually the interstitial areas finely reticulopunctate and shining; several fine interspinal striae present; dorsum of postpetiole weakly sculptured or almost smooth and shining.

Pilosity: Hairs yellow, fine, numerous and of various lengths, measuring 0.07 to 0.18 mm; dorsal and ventral surface of head with semi-erect short and long hairs; dorsum of pronotum and mesonotum with numerous predominantly long erect hairs; propodeal base with few scattered short hairs; pedicel with numerous semi-erect long hairs; gaster with numerous predominantly long semi-erect hairs, most of them blunt apically. Pubescence on antennae, sparse on legs and absent on gaster.

Color: Head and gaster ebony black; thorax and legs a little lighter.

Worker:

Measurements: Body length 1.50 to 1.60 mm. Head length 0.44 to

0.51; head width 0.38 to 0.41 mm. Thoracic length 0.46 to 0.51 mm. Scape length 0.40 to 0.46 mm. Scape Index 97 to 106.

Structural characters: Head, excluding mandibles, slightly longer than broad; occipital border slightly concave medianly; frontal area shallowly depressed; frontal carinae straight, short and slightly divergent; scapes surpassing the occipital lobes by an amount less than the first funicular joint; mandibles slender, apically with two sharp teeth, the remaining masticatory border with several smaller denticles.






20



Pronotum and mesonotum, in profile, convex; propodeal spines long but blunt, projecting obliquely dorsad; petiole stout, the apex of the node angular; postpetiole strongly convex dorsally and almost flat ventrally; gaster truncate at the base.

Sculpture: Head smooth and strongly shining; pleural surfaces of pronotum shining; pleurae of mesonotum, propodeum and petiole strongly reticulo-punctate, postpetiole weakly punctate.

Seen from above dorsum of pronotum and anterior mesonotum smooth and strongly shining, posterior area of mesonotum, propodeal base and propodeal declivity reticulo-punctate; dorsum of postpetiole and gaster smooth and shining.

Pilosity: Hairs white, measuring less than 0.08 mm; sparse all over the body, semi-erect and of mixed sizes on the head, long and erect on the dorsum of thorax and long and slender on gaster; under the stereo microscope the hairs seemed spatulate, but scanning micrographs later revealed that such hairs are branched at the tips (fig. 53, 54), therefore, plumose or sub-plumose.

Color: Ebony black, the appendages lighter, mandibles yellowish to dark brown.

Females:

Measurements: Body length 4.60 mm. Head length 0.90 mm; head width 0.95 mm. Thoracic length 1.46 mm. Scape length 0.51 mm. Scape Index 58.

Structural characters: Head broader than long, broadest posteriorly; occipital border slightly concave; sides slightly convex and






21



narrowing anteriorly toward the mandibular insertions; anterior border of clypeus weakly notched in the middle.

Frontal area depressed; antennae like those on the major;

scapes longer than half the head length; eyes measuring about 0.24 mm in diameter and located on anterior half of head; ocelli measuring about 0.12 mm in diameter.

Thorax flat dorsally, narrower than the head measured through

the wing insertions. Mesoscutum and scutellum well separated; propodeal base and declivous face not forming a distinct angle. Propodeal spines stout, blunt and directed posteriorly; petiole stout, the node seen from above with parallel sides, the crest slightly depressed in the middle; postpetiole twice as wide as petiole, with rather angular lateral projections; gaster truncate at the base.

Sculpture: Cephalic rugae strong, covering most of the anterior

part of the head, but disappearing before reaching the occipital border, the interstitial areas between rugae with widely spaced weak reticulate punctation; frontal area smooth and shining with two mesal longitudinal striae; frontal carinae long and diverging posteriorly.

Sides of pronotum and propodeum reticulate; episternum smooth and shining; propodeal base and declivous face smooth and shining; petiole and postpetiole punctate and opaque; gaster, seen from above, with fine longitudinal striae near the base, the interstitial areas between striae shining but with widely spaced reticulate punctation, the rest shining and with circular piligerous punctures.

Pilosity: Hairs yellow, pointed; dorsum of thorax, pedicel and gaster with numerous short and long hairs; gaster usually more pilose than the remaining parts; pubescence present on the antennae and legs.






22




Color: Reddish brown; the gaster darker.

Male:

Unknown.

This species is named in honor of my beloved first born son,

Adriano de Resende Naves.

Holotype: Major from Gainesville, Florida, collected by M. A.

Naves.

Paratypes: The type material comprises 30 majors and 60 workers

collected in Gainesville, Florida from several colonies on August 13, 1975 and 25 majors and 33 workers from several colonies from Archbold Biological Research Station at Lake Placid, Florida, in August, 1975.

All were collected by M. A. Naves.

The holotype and several paratypes will be deposited at the Museum

of Comparative Zoology, Harvard University, Cambridge, Massachussetts.

Paratypes will also be deposited in the National Museum of Natural

History, Washington, D.C., Florida State Museum, Department of Plant

Industry, Gainesville, Florida and in the W. F. Buren and M. A. Naves

collections.

Discussion:

This species is fairly common in Florida. Its range seems to be from central to northern Florida. It may also occur in southern Georgia., Its preferable habitat seems to be sandy soils, in clear areas among trees, the same habitat as in P. metallescens, with which it is very similar to the naked eye.

The colonies have about 60 majors and more than 300 workers. The main chamber is located 30 to 40 cm deep in the soil and is directly connected to a vertical tunnel of about 1 to 2 mm in diameter, which usually is the





23



center of a small crater on the soil surface. The main flights of sexuals occur in the summer in July and August. The species is entirely diurnal. The main sources of food are small seeds, although it also scavenges small dead arthropods. The majors as well as the workers usually forage for food and help carry it back to the colony. It is not an agressive species. The majors when disturbed try to escape to hiding places.

P. adrianoi differs from P. metallescens by its ebony black color,

absence of reticulate punctation on the head and by its smooth and strongly shining occipital lobes and lack of metallic reflections. P. adrianoi also differs from P. littoralis by its small head, and the occipital lobes not strongly rounded, sides of head not parallel and mandibles not stout and strongly curved. P. metallescens and P. littoralis usually have a reddish brown head and thorax, the gaster always darker or blackish as opposed to P. adrianoi which has a shining concolorously ebony black body.






24




2. Pheidole anastasii Emery

P. anastasii Emery, 1896. Soc. Ent. Ital., Bol. 28:76. Forel,

1901. Naturhist. Mus. Hamburg, Mitt. 18:78. Wheeler, 1932. N.Y. Ent. Soc., Journ. 40:6. Smith, 1933. Fla. Ent. 17:23. Creighton, 1950. Bull. Mus. Comp. Zoll. 104:169. Gregg, 1958. N.Y. Ent. Soc. 66:16.

P,. floridana Van Pelt, 1956. Amer. Midland Nat., 56:376, 384.

Carroll, 1970. Unpublished M.S. thesis, University of Florida, p. 32-34. Whitcomb, Denmark, Bhatkar, and Greene, 197. Fla. Ent., 55:133. (nec Emery).

Type locality: Jimenes, Costa Rica

Types : None in the U.S.

Range : Florida, southern Alabama and Georgia Diagnosis:

A small yellowish species differing from P. floridana in having opalescent surfaces dorsally on the first gastric tergite; differing from P. moerens and P. greggi n. sp. by having the postpetiole twice as broad as the petiole. Distinct lateral connules are present. It differs from P. dentigula in having smooth and shining occipital lobes. Discussion:

Wheeler (1932) reported P. anastasii (fig. 17) from Florida. It seems hard to believe that it is an introduced species as Wheeler suggested. I have found this species throughout Florida from the keys to the Panhandle. Its constant nesting at the base of pines and the occurrence of two parasites, P. guiescens n. sp., an inquiline ant, and a hymenopteran parasite in the genus Orasema, seem to contradict Wheeler's suggestion that P. anastasii is an exotic species.






25



Due to the close resemblance of this species to P. floridana Emery, P. anastasii probably has been misdetermined in several records from Florida.

The difficulty of having access to Emery's types of P. anastasii

makes it difficult to determine if the Florida species is conspecific with the Costa Rican types. Wheeler, however, did have knowledge of the types of P. anastasii when he reported it from Florida.

P. anastasii was usually found nesting under the bark at the base of pines or along the roots and only rarely in the soil. The colonies are monogynous and have more than 100 majors and over 500 workers. Several chambers are constructed interconnected by a string of galleries under the pine bark. The workers forage over 4 m from the colony and once food is located majors are recruited to help transport it back to the colony. It feeds on seeds, fruits, and scavenges on small dead arthropods and is predaceous on small live arthropods.

Laboratory colonies are very easy to keep and usually accept members of other colonies. Colonies that have lost their queens will also accept any other queens of anastasii. Several times I collected two or three nest founding females found together in one chamber. In the lab one female always killed the others before the first workers were reared.





26




3. Pheidole bureni n. sp.

Diagnosis:

A medium sized brownish red species with darker gaster, difering from P. diversipilosa and P. crassicornis tetra by the absence of long erect hairs on the dorsum of the first gastric tergite. It differs from P. crassicornis by having the occipital excision deeper and much more angularly V shaped, and the occipital lobes sharper. Description:

Major:

Measurements: Body length 5.30 to 5.70 mmn. Head length 1.54

to 1.67 mm; head width 1.43 to 1.54. Thoracic length, 1.44 to 1.54 mm. Scape length, 0.85 to 0.92 mm; Scape Index, 54 to 55. Depth of

occipital excision usually over 0.12 mm.

Structural characters: Head shape as in fig. 60. Head excluding

mandibles longer than wide, broadest above the eyes, from where the sides curve and narrow posteriorly; the occipital excision deep and wide and angularly V shaped; occipital lobes much more angular than

in crassicornis or diversipilosa; frontal area impressed; anterior

border of clypeus with shallow median notch; frons strongly pronounced and the vertex inclined posteriorly; genae flat and depressed

laterally. Frontal furrow distinct and extending from frontal area

to the occipital excision; scapes flattened and curved at base, a

little longer than half head length; mandibles curved laterally with

two strong, sharp teeth apically; the rest of the border without

teeth or with several small denticles; eyes measuring about 0.26 mm

in diameter and with over 100 small facets.





27



In profile, pronotum and anterior mesonotum convex, mesonotum with a wide impression in the middle; posteriorly the mesonotum abruptly descending to the mesopropodeal impression; propodeal base longer than declivity; propodeal spines short and pointed dorsally; petiole stout, the anterior face and posterior face sloping, the node quite large and its crest in profile varying from weakly angular to strongly convex; the ventral border slightly convex anteriorly; postpetiole strongly convex dorsally and flat ventrally.

Seen from above, pronotum with pronounced but rounded shoulders; petiole with parallel sides, the crest of node with a shallow concave impression in the middle; postpetiole almost two times wider than long; slightly angular to blunt laterally; gaster truncate at the base.

Sculpture: Fine longitudinal striae present on the frons, interstitial areas finely and weakly reticulo-punctate; vertex and occipital lobes smooth and shining; genae with longitudinal rugae, the interstitial areas weakly punctate; frontalicarinae short and slightly diverging posteriorly; frontal area smooth and shining; clypeus smooth and shining, usually with a weak median carinae; frontal furrow with transverse short rugulae posteriorly; ventral area of head smooth and shining, its anterior margin with two well developed sharp teeth.

Pronotal pleurae weakly reticulate; mesonotum, propodeum and

petiole finely reticulo-punctate, the floor of each punctation strongly shining.

Seen from above, pronotum and adjacent area of mesonotum with weak transverse striae and weakly reticulated; mesonotal dorsum smooth and strongly shining; mesonotal impression not easily seen;





28




mesopropodeal impression depressed and shining; prododeal base weakly reticulo-punctate and shining; propodeal declivity smooth and strongly shining; petiole and postpetiole finely reticulopunctate and opaque; gaster smooth and shining.

Pilosity: Hairs yellow and pointed, measuring up to 0.26 mm;

vertex with two long hairs on each side and several along the frontal carinae; anterior ventral face of head with semi-erect short hair; posterior of head and mandibles and scapes with fine appressed pubescence; a few erect hairs on the scapes; dorsum of pronotum and mesonotum with a few mixed short and long erect hairs; scattered appressed pubescence on pronotum; each side of the petiolar node with long semi-erect hairs; postpetiole with several semi-erect long hairs; first gastric tergite with numerous fine appressed pubescent hairs; ventrally the posterior half of gaster with mixed short and long erect and semi-erect hairs; appendages with sparse appressed pubescence; funiculus with very fine pubescence.

Color: Reddish brown, the gaster usually darker. Worker:

Measurements: Body length 3.10 to 3.20 mm. Head length 0.77

to 0.79 mm; head width 0.70 to 0.77mm. Thoracic length 1.05 to 1.13. Scape length 0.82 to 0.90. Scape Index 106 to 114.

Structural characters: Head, excluding mandibles, a little

longer than broad; occipital border flat to slightly concave; sides strongly convex; frontal area broader than long and shallow; frontal carinae short and parallel; scapes surpassing the occipital border






29




by an amount greater than the first funicular joint; mandibles slender, curved apically and with two sharp teeth, the rest of the border with several small denticles; anterior clypeal border convex; eyes measuring 0.18 mm in diameter and with more than 65 facets.

In profile, pronotum and anterior mesonotum strongly convex; mesonotum with a wide transverse impression; propodeal base longer than the declivity; propodeal spines short and obliquely pointed dorsad; petiole stout, the node strongly convex in profile; postpetiole strongly convex dorsally in profile and flat ventrally.

Seen from above, postpetiole 1.5 times wider than long and the sides strongly convex; gaster truncate at the base.

Sculpture: Head finely reticulo-punctate except the frons and vertex which have smooth and shining areas; frontal area smooth and shining; clypeus with a median carina and usually smooth or weakly reticulo-punctate; genae with several short longitudinal striae emerging from anterior reticulation.

Pleurae of throax reticulo-punctate; petiole and postpetiole finely reticulo-punctate; gaster smooth and shining.

Dorsum of mesonotum with small smooth and shining areas; basal face of propodeun strongly reticulo-punctate and the declivity weakly reticulate and shining.

Pilosity: Hairs yellow, numerous, measuring up to 0.20 mm;

mixed short and long semi-erect hairs present on head; erect, short and long hairs on the dorsum of thorax; two long semi-erect hairs on the pediole; numerous short and long, erect and semi-erect hairs on postpetiole and gaster.





30




Color: Concolorously dark brown. Female:

Measurements: Body length 7.50 mm. Head length 1.20 mm; head

width 1.38 mm. Thoracic length 2.41 mm. Scape length 1.03 mm. Scape Index 86.

Structural characters: Head broader than long, broadest posteriorly; occipital border straight; sides slightly convex and narrowing anteriorly; middle of anterior border of clypeus with an impression; frontal area broad and depressed; scapes stout, longer than 3/4 the head length, the basal portions strongly curved and flattened, the flattening more latero-mesal rather than dorso-ventral as in a major. Eyes measuring about 0.40 mm in diameter and with over 200 small facets; ocelli measuring about 0.15 mm in diameter; frontal carina expanded and raised. Thorax flat dorsally, measured through the Wing insertions slightly wider than the head; mesoscutum and scutelum separated by a distinct suture.

Propodeal base and declivity not forming a distinct angle;

propodeal spines very short, thick and pointed dorsad; petiole stout; postpetiole twice as broad as long, the sides produced into distinct, blunt lateral projections; these slightly convex anteriorly and posteriorly weakly concave; gaster truncate at the base.

Sculpture: Frontal area and clypeus smooth and shining as well as a narrow area of the frons between the frontal area and the median ocellus; frons and lower vertex longitudinally striate laterally; posterior areas of head smooth and shining; genae with pronounced





31



rugae; frontal carinae short and diverging posteriorly; ventral

surface of the head smooth and shining, its anterior margin with

three distinct teeth, the median tooth short and lobe-like and the lateral ones long and sharp; sides of pronotum opaque; esisternum

smooth and shining; propodeum with transverse striae, the interstitial areas weakly punctate and shining; propodeal base and

declivity smooth and shining; sides of pedicel finely reticulopunctate; gaster smooth and shining.

Pilosity: Hairs yellow and pointed and measuring up to 0.40 mm

in length; head with numerous short and long erect and semi-erect hairs; dorsum of thorax and petiole with numerous short and long

pointed hairs; postpetiole and gaster with numerous, predominantly

short semi-erect hairs; short sparse fine and pointed semi-erect

hairs on scapes and apendages.

Color: Light reddish brown

Male: Unknown

Holotype: Major, collected at Alice, Texas

Paratypes: 6 majors, 7 workers and 1 female, from the same

colony as the Holotype.

The type material comprises part of a colony collected by William F. Buren on October 23, 1942 at Alice, Texas.

The holotype, the female paratype and one worker paratype will be deposited in the Museum of Comparative oology, Harvard University, Cambridge, Massachussetts. Paratypes will also be deposited in the NationalMuseum of Natural History, Washington, D.C., the Florida State





32



Museum, Department of Plant Industry, Gainesville, Florida, and in the W. F. Buren and M. A. Naves collections. Discussion:

This species is named in honor of Dr. William F. Buren, Professor of Entomology, University of Florida.






33





4. Pheidole carrolli n. sp.

Diagnosis:

Areddishbrown species related to the P. pilifera group. It

differs from P. pilifera and P. pilifera artemisia by the absence of coarse reticulate occipital rugae and blunt postpetiolar connules. It differs from P. pilifera coloradensis by its larger head, strongly convex pronotum and anterior mesonotum, and longer and finer thoracic and gastric hairs. It differs from P. pilifera pacifica by its larger size and cephalic rugae extending to the occipital lobes and absence of transverse occipital rugae.

Description:

Major:

Measurements: Body length 3.20 to 3.60 mm. Head length

1.80 to 1.90 mm; head width 1.69 to 1.80 mm. Thoracic length 1.36

to 1.44 mm. Scape length0.77 to 0.82 mm. Scape Index 42 to 44.

Structural characters: Head shape as in fig. 14. Head, excluding mandibles, longer than broad, occipital lobes prominent and well

rounded, separated by a broad and deep excision; a furrow present,

starting from the frontal area to the occipital excision. Frontal area depressed and bearing a median longitudinal striation. Clypeus with a weak median carinae which is continuous with the frontal'area

striation; anterior border of clypeus with a shallow weak emargination; frontal carinae divergent; anterior ventral margin of the head

without teeth; scapes slender at base, thickening apically and measuring less than half the head length. Mandibles stout, strongly curved laterally and with two apical large blunt teeth, the remaining border





34



edentate, except for a small basal tooth-like lobe. Eyes small, measuring about 0.25 mm in diametLer with over 60 facets and located on the anterior third of the head. Seen from the side, thorax smaller than head; pronotum and anterior mesonotum strongly convex; posterior of mesonotum descending to the mesopropodeal impression, forming an obtuse angle with the propodeal base; base of propodeum slightly shorter than declivity; propodeal spines short and projecting dorsad; petiole, in profile, stout and with a short anterior peduncle, anterior face gently curved and posterior face short; apex of node weakly angular to moderately convex; seen from above the sides of the petiole parallel and the node notched in the middle; postpetiole in profile strongly convex dorsally, slightly concave and much shorter ventrally; seen from above, postpetiole two times or more wider than petiole and with lateral connules; gaster oval and much smaller than the head.

Sculpture: Cephalic longitudinal rugae extending posteriorly and somewhat divergently; the lateral rugae usually do not reach the occipital lobes; occipital lobes smooth and shining laterally and dorsally; rugae present on the genae, extending longitudinally; posterior to the eyes the surfaces smooth and shining; head posteriorly with coarse circular piligerous punctures. Dorsum of pronotum and mesonotum with longitudinal or transverse rugae or both, the interstitial areas with fine reticulate punctation. The mesopropodeal impression wide and with longitudinal rugules, the interstitial areas smooth and weakly shining; the propodeal base and declivity






35




finely reticulo-punctate and weakly shining. Several transverse striae present between bases of spines; petiole and postpetiole opaque and weakly reticulo-punctate; gaster smooth and shining except in the basalmost area where it is opaque and weakly punctate.

Pilosity: Hairs yellow, pointed, short and semi-erect and sparse on the anterior face of the head, short and long and semi-erect ventrally; dorsum of thorax with numerous short and long erect hairs measuring up to 0.26 mm; pedicel with numerous semi-erect short and long hair; sparse pubescence confined to antenae and appendages.

Color: Reddish brown, appendages lighter; dorsum of gaster slightly darker; anterior margin of clypeus and the border of mandibles dark.

Worker:

Measurements: Body length 2.30 to 2.40 mm. Head length 0.56

to 0.64 mm; head width 0.56 to 0.58 mm. Thoracic length 0.74 to 0.77 mm. Scape length 0.53 to 0.59 mm. Scape Index 92 to 94 mm.

Structural characters: Head usually a little longer than broad, occipital borders weakly convex; frontal area impressed and with a longitudinal striation in the middle; frontal carinae slightly divergent; clypeus with a weak median carina and 2 longitudinal striae laterally; anterior border curved and without any notch. Mandibles slender with

2 large and sharp apical teeth and 4 or 5 smaller teeth on the remaining masticatory border; scapes usually surpassing the occipital border by an amount no greater than the first funicular joint; eyes large with about 45 facets and measuring about 0.15 mm in diameter.






36




In profile, pro-mesonotum !moderately convex; propodeal spines small, sharp, and projecting obliquely dorsad; petiole as in the major, except the crest is proportionately much broader; postpetiole convex dorsally and flat ventrally, seen from above the sides are strongly convex and almost 2 times wider than petiole; gaster oval.

Sculpture: Head reticulo-punctate and opaque; a few rugules on the genae, between the eyes and on the mandibles; thorax, in profile, reticulo-punctate; dorsum of pronotum and mesonotum usually with a few longitudinal or transverse striae; petiole reticulopunctate; sides of postpetiole weakly punctate, dorsum weakly shining; gaster smooth and shining.

Pilosity: Erect hair yellow, short on the dorsal surface of head, longer on the ventral surface and anterior clypeal margin; long and sparse on thorax and pedicel; gaster with rather sparse hair of mixed lengths; pubescence confined to the antennae and legs.

Color: Reddish brown, sometimes the mandibles and legs are lighter.

Holotype: Major, collected near Gainesville, Florida by M. A. Naves.

Paratype: 3 majors and 20 workers probably from the same nest as the holotype, collected as follows:

Dr. John F. Carroll collected one major and several workers on August 2, 1973, near Gainesville, Florida. M. A. Naves collected three majors and several workers in August, 1975, at the same locality.





37




The holotype major and 3 worker paratypes will be deposited at

the Museum of Comparative Zoology, Harvard University, Cambridge, Massachussetts. Paratypes will also be deposited in the National Museum of Natural History, Washington, D.C., Florida State Museum, Department of Plant Industry, Gainesville, Florida and in the M. A.

Naves collections.

This species was also collected by me at Tall Timbers Research

Station, but the specimens, unfortunately, were inadvertantly lost. Discussion:

This species has unusual behavior which is not common to other Pheidole known to me. It was always found in shaded areas in sandy soils among tall trees such as pines and oaks. It forages when the temperatures are not high and avoids direct sunlight. It is easily found at dusk or in the morning. The workers usually go out at random. Usually there are no more than 15 workers foraging at one time, and the foraging territory usually is within 5 meters from the nest entrance. The nest entrance is small and usually very hard to spot. The worker is sluggish compared with most other species. The majors are very rare. For a year two colonies were periodically checked and yielded only one callow major in 1975 and two majors in March, 1976. Dr. John F. Carroll also collected one major in the same locality in August, 1973. Periodic excavation usually also yielded a chamber approximately 4X2X2 cms located 30 cm deep and full of various seeds, some apparently with a black fungal mycelium. Although the excavation for one of these colonies at one time was 1.7 meters deep 1.5 meters in diameter, I was unable to locate the main chamber or any majors





38




or any sexual forms. Probably the main chamber or chambers are located very deep in the ground. It seems quite obvious that the colonies are not large. The disproportional size of the major and its rarity also seems to indicate that it has limited importance in sustaining the colony, which is not true for most species of the genus.

Another feature of this species when disturbed is the ability of the workers to flign death. The minors try to flex themselves around particles of soil which make them very difficult to see. The majors hide themselves and also seem to avoid light, but their large size and more shining surfaces compared with the workers make them easier to spot.

Workers of this species, although not many, have been captured by baiting near Gainesville.

One colony was located at Tall Timbers Research Station in 1974, which failed to yield any major or sexual form.

Perhaps P. carrolli is a species which may be losing its major in the evolutionary process. The fewer majors available show rather more morphological variation than is normal in most Pheidole species.

This species is named after Dr. John F. Carroll, my friend and colleague during several years of graduate study at the University of Florida.





39



5. Pheidole crassicornis Emery

P. crassicornis Emery, 1985. Zool. Jahrb., Abt. F. System. 8:289, 296. Forel, 1901. Soc. Ent. Belg., Ann. 45:350. Wheeler 1904. Amer.

Mus. Nat. Hist., Bul. 20:302. Creighton, 1950. Bull. Mus. Comp. Zool.

104:175-176. Gregg, 1958. N.Y. Ent. Soc., 66:20.

Type locality: Charlotte, North Carolina

Types : Cotypes, M. C. Z. and A. M. N. H.

Range : Southeastern States Diagnosis:

P. crassicornis (fig. 5) differs from P. diversipilosa and P. crassicornis tetra by the absence of erect and semi-erect hairs on the gaster. It also differs from P. bureni n. sp. by the convex occipital lobes and the occipital impression which is not sharply V shapped. P. crassicornis can be easily distinguished from all Florida Pheidole other than diversipilosa by the basally strongly bent and flattened scapes. Discussion:

The finding of P. crassicornis at Gainesville and at Tall Timbers Research Station will help the understanding of this group because it is sympatric with P. diversipilosa at Tall Timbers. P. crassicornis is apparently confined to the northern part of Florida where it occurs in open areas, usually among trees. Its nests seem to be in deep soil. After digging as deep as 60 cm, I was unable to find any chambers of three colonies. The opening at the soil surface is hard to see. The typical craters common to other species were never observed. The major and workers were seen carrying live termites and other small dead arthropods. The workers and a few majors were observed foraging approximately 4 meters or more from the nest opening.






40




6. Pheidole dentata Mayr

P. morrisi var. dentata Mayr 1886. Vehr. Zool Bot. Ges. Wein, 36:457.

P. dentata Forel, 1901. Ann. Soc. Ent. Belg., 45:351 Wheeler,

1904. Amer. Mus. Nat. Hist., Bull. 20:302. Mitchell and Pierce, 1912. Ent. Soc. Wash., Proc. 14:71. Smith, 1918. Ent. News, 29:22. Smith, 1924. Ent. News 35:77. Smith, 1930. Fla. Ent. 14:3. Wheeler, 1932. N.Y. Ent. Soc., 40:6. Dennis, 1938. Ent. Soc. Amer., Ann. 31:281,

304. Van Pelt, 1950. Ent. News 61:161-163. Creighton, 1950. Bull. Mus. Comp. Zool. 104:177. Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc. 55:71. Gregg, 1953. N.Y. Ent. Soc. 66:21-22. Smith, 1965.

U.S. Dept. Agr., Tech. Bul. 1326, pp. 27-28. Wilson, 1975. Science

190:798-800.

P. dentata var. faisonica Forel., 1901. Ann. Ent. Soc. Belg., 45:352.

P. commutata Mayr 1886. Verb. Zool. Bot. Ges. Wein. 360459. Emery, 1895. Zool Jahrb. Syst., 8:289.

Leptothorax tennesseensis Cole, 1938. Amer. Midland Nat. 19:238.

P. dentata var. commutata Wheeler, 1907. Amer. Mus. Nat. Hist.,

Bull. 7:18. Wheeler, 1908. Amer. Mus. Nat. Hist., Bull. 24:460-461.

Smith, 1924 Ent. News 35:77. Cole, 1940. Amer. Midland Nat. 24:29, 44. Smith, 1951. Cat. Hym. Amer. No. of Mex.,U.SD.A. Mon 2:802.

Type locality: Florida

Types : None in this country

Range : Southeastern States, Tennessee and Gulf states to Texas.





41




Diagnosis:

P. dentata (fig. 30) is a reddish yellow to dark brown species

differing from P. morrisi by the distinct and sharp propodeal spines. It differs from P. megacephala in not having a cordate head and differs from P. vinelandica by the larger size and absence of a longitudinal striation on the frontal area, and having the mesonotum transversely impressed in the middle.

Discussion:

P. dentata is one of the more common species in Florida. It is

easy to understand why this species caused so many misunderstandings in the past. There is much variation, not only in size and color, but also in habitat preference. I have found colonies of small dark colored specimens in wooded areas around Gainesville and large specimens that nest in sandy soil on beaches in south Florida and the Florida Keys. I also found a yellowish variant that inhabits the marshlands of the keys and another variant with quite large majors that vary in color from reddish to very dark brown nesting in open areas around Gainesville. This morphological variation seems to be without taxonomic significan.

Three mated young queens collected at Gainesville were easily reared in the laboratory. All three had majors after 50 days. The species is highly carnivorous and the colonies have a tremendous growth when fed with living arthropods. One of the colonies was inadvertantly left without food and moisture for three weeks. The result was that the adults formed from the stressed broods were intermediate in size and structure between workers and majors. Some of the intermediates were not as active






42





as the normal caste. This is a very interesting species for ecological and laboratory studies. Recent studies by Wilson (1975) have shown that workers of P. dentata use odor trails to recruit nestmates to food discoveries and new nest sites. The same pheromone is used also to recruit major workers to the vicinity of intruders. This type of alarm recruitment proved rather narrowly specific and works best against some of the potentially more important enemies of P. dentata in the genus Solenopsis.





43




7. Pheidole dentigula M. R. Smith P. dentigula Smith, 1927. Ent. News 38:310. Smith, 1928. Ent. News 39:245-246. Cole, 1940. Amer. Midland Nat. 24:42, 45. Smith, 1944. Fla. Ent. 27:14. Creighton, 1950. Bull. Mus. Comp. Zool, 104: 178. Gregg, 1958. N.Y. Ent. Soc. 66:21. Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:14.

Type locality: Mississippi State University, Starkville, Mississippi.

Types : Cotypes, National Museum

Range : Tennessee, southward through Alabama and Mississippi, and north Florida

Diagnosis:

P. dentigula (fig. 13) is a small yellowish species that differs from P. greggi, P. anastasii, P. floridana, P. tysoni, P. bicarinata and P. moerens by the reticulated, opaque occipital lobes. It differs from P. metallescens by having a smooth and shining propodeal base and lacks metallic reflections. The teeth on the anterior ventral border of the head are prominent.

Discussion:

Smith (1944) reported P. dentigula from Wakulla County (Florida).

Dr. John F. Carroll also collected this species in Marion County (Florida) on April 3, 1973. According to Smith (1944) this species nests in the soil and in well rotted stumps in wooded areas, especially where the soil conatins considerable hummus.






44




8. Pheidole diversipilosa Wheeler P. crassicornis var. diversipilosa Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:467.

P. crassicornis Creighton, 1950. Bull. Mus. Comp. Zool., 104:175176.

Type locality: Fort Davis, Texas.

Types : M.C.Z.

Range : Northwestern Florida to western Texas Diagnosis:

P. diversipilosa (fig. 3) is a reddish yellow species that differs from P. crassicornis and P. bureni n. sp. by the numerous erect hairs on the gaster. In P. crassicornis tetra the gastric hairs are nearly all of the same short length rather than obviously of mixed lengths as in diversipilosa.

Discussion:

P. diversipilosa was collected at Tall Timbers Research Station several times where P. crassicornis is also present. Morphological distinction is constant between the two species. There is no real overlap in the pilosity characters. Therefore, I have treated P. diversipilosa as a full species on the basis that it can be sympatric with P. crassicornis but still retains constant morphological distinction. Creighton (1950) treated diversipilosa as an intergrade between crassicornis and tetra. The finding of both crassicornis and diversipilosa in the same locality in Florida invalidates Creighton's viewpoint.

Specimens of P. crassicornis collected in Gainesville were within the variation shown by the Tall Timbers specimens. I was unable to find






45





specimens of P. diversipilosa in Gainesville. Comparison with the types and cotypes for the two taxa in the M. C. Z. and further material of diversipilosa collected at Columbus and Mission, Texas by W. F. Buren as well as the finding of another related species, P. bureni n. sp. (fig. 60, 61, 62) emphasize that we are dealing with a species group that is widely spread and contains several discrete species. P. crassicornis is predominantly an eastern species. P. diversiiilosa, P. bureni and P. crassicornis tetra are western species. Of these apparently only P. diversipilosa reaches east into Florida. The lack of better records for the distribution of these species probably led Creighton (1950) to synomymizing crassicornis and diversipilosa. P. crassicornis tetra appears to be a very distinct form and may be a discrete species also, but I have not attempted taxonomic treatment in the present paper, since it does not occur in Florida.






46




9. Pheidole fallax obscurithorax Santschi

P. fallax arenicola var. obscurithorax Santschi, 1923. A~m Soc. Ent. Belg. 63:58. Kempf, 1972. Studia Ent. 15:191.

Type locality: Cordoba, Argentina

Types : None in this country

Range : Probably westernmost Florida to at least Mobile, Alabama in U.S. Probably widely ranging in South America.

Diagnosis:

P. fallax obscurithorax (fig. 7, 8), is a large, very dark species over 6 mm in body length. Its characteristics, such as the heavily sculptured head and thick scape which is curved mesally and more angular laterally are unique among the Pheidole of North America. Discussion:

P. fallax obscurithorax (fig. 8) is a member of the difficult large neotropical fallax group which has 6 subspecies and 6 varieties. The specimens collected near the Florida border matched specimens in the M.C.Z. identified by W. W. Kempf.

This species was collected close to the Florida and Alabama border in Balbwin County, Alabama by Dr. John F. Carroll. E. O. Wilson also (personal communication) collected this species in Mobile, Alabama in 1950 within 1/2 mile of the ship docking area. It is apparent that this introduced species has been in the U.S. for at least 26 years but, unlike the imported fire ant, has not spread widely.

It is not a surprise to find another exotic ant species from South America in the U.S. Brachymyrmex patagonicus Mayr (not published),





47




Iridomyrmex humilis (Mayr), Solenopsis invicta Buren, S. richteri Forel, and Pheidole fallax obscurithorax appear to be a group of South American ants inadvertently imported to the ports of New Orleans, and/or Mobile, and/or possibly Pensacola.

All five species have ranges along or near the Paraguay and La Plata Rivers in Western Brazil, Paraguay and Argentina. The Paraguay River is navigable by ocean going vessels up river to Asuncion, Paraguay.





48




10. Pheidole floridana Emery

P. flavens floridana Emery, 1895. Zool. Jahrb., Abt. f. System. 8:293. Smith, 1930. Fla. Ent. 14:3.

P. floridana Emery, 1895. Bull. Soc. Ent. Ital. 28:77. Wheeler, 1932. N.Y. Ent. Soc., 40:6. Creighton, 1950. Bull. Mus. Comp. Zool.

104:179. Smith, 1951. Cat. Hym. Amer. No. of Mex., U.S.D.A. Mon.

2:802. Gregg, 1958. N.Y. Ent. Soc. 66:21-22, 35. Kempf, 1972.

Studia Ent. 15:192.

Type locality: Coconut Grove, Florida

Types : M.C.Z.

Range : Southwest Florida Diagnosis:

P. floridana (fig. 19) is a small yellowish species differing from P. greggi n. sp. and P. moerens by the distinct lateral connules on the postpetiole. It differs from P. anastasii by the absence of opalescence on the dorsum of the first gastric tergite. Discussion:

P. floridana seems to be confined to southeast Florida in the Miami area. This is the only place where I was able to locate this species. Due to its close relationship to P. anastasii the latter has been misidentified as P. floridana many times, thus, mistakenly extending the supposed range of P. floridana. P. anastasii is actually the species widely distributed in Florida, while floridana is absent or at least must be rare in most of the state.





49





11. Pheidole greggi n. sp.

P. flavens sculptior M. R. Smith, 1951. Cat. Hym. Amer. No. of Mex. U.S.D.A. No. 2, p. 802 (nec Forel).

P. flavens Gregg, R. E. 1958. N.Y. Entomol. Soc. 66:21 (nec Roger).

Diagnosis:

A small yellowish species closely related to P. flavens Roger, P. flavens sculptior Forel, P. floridana Emery, P. anastasii Emery, and P. moerens Wheeler. Antennal scrobe weakly depressed but punctate and opaque. Frontal carinae not surpassing apices of scapes in repose. Postpetiole without lateral projections. P. greggi has been previously confounded with P. flavens sculptior and P. flavens. Description:

Major:

Measurement: Body length 2.60 to 2.90 mm. Head length 0.82

to 0.89 mm; head width 0.79 to 0.82. Thoracic length 0.72 to 0.77 mm.

Scape length 0.41 to 0.46 mm. Scape Index 52 to 56.

Structural characters: Head shape as in fig. 21; head, excluding mandibles, usually slightly longer than broad, broadest behind

the eyes. The occipital lobes strongly curved and convex laterally

but less curved or even angular mesally; median occipital sulcus

shallow. Frontal area small and depressed; antennal scrobes weakly

or not at all depressed. Clypeus with weak median carina, its

anterior margin with weak median emargination. Frontal carinae

short and divergent. Scapes slender at base, widening apically and usually half or more the head length; mandibles stout, each curved






50



apically and bearing two teeth, the remaining border edentate, except for a small basal tooth. Eyes small and with approximately 23 facets.

In profile, pronotum and mesonotum strongly convex and descending through an abrupt declivity to the mesopropodeal impression which is rather weakly impressed; propodeal spines small and sharp, pointed obliquely dorsad; petiole with a short anterior peduncle; anterior face of node gently sloping, the posterior face sharply declivous; crest of node convex and blunt in profile but not thickened and sub-truncate as in anastasii and floridana, ventrally the petiole straight rather than convex as in anastasii and floridana; postpetiole strongly convex dorsally, flat to slightly concave ventrally.

Seen from above shoulders of pronotum well rounded; sides of

petiole parallel; postpetiole small; the sides convex and the shape sub-trapezoidal, without lateral connules, little wider than petiole; gaster truncate at base and slightly smaller than the head.

Sculpture: Head opaque, cephalic rugae sub-parallel mesally, slightly diverging laterally and reaching close to occipital lobes before disappearing; the interstitial areas between rugae with weak reticulate punctation; the occipital lobes smooth and shining; the scrobes finely reticulo-punctate; genae and areas posterior to the eyes, as well as the area anterior to the eyes longitudinally rugose, the rugae disappearing before reaching the occipital lobes, occipital lobes with fine circular piligerous punctures.






51



Sides of thorax, petiole and postpetiole reticulo-punctate, opaque to subopaque. Pronotal dorsum with transverse irregular striae; the interstitial areas weakly reticulo-punctate to smooth and weakly shining; mesopropodeal impression wide and with longitudinal rugulae; the interstitial areas smooth and shining; propodeal base reticulo-punctate and opaque; postpetiolar dorsum smooth and shining; gaster smooth and shining.

Pilosit : Erect hair yellow, pointed, numerous; both short and long hairs on head, thorax, pedicel and gaster; pubescence sparse on antennae and legs, absent on gaster.

Color: Concolorously yellow

Worker:

Measurements: Body length 1.50 to 1.80 mm. Head length 0.46

to 0.52 mm; head width 0.43 to 0.46 mm. Thoracic length 0.41 to 0.59 mm. Scape length 0.41 to 0.43 mm. Scape Index 82 to 89.

Structural characters: Head as in fig. 45, slightly longer than broad, broadest through the eyes; posterior border of head slightly notched medially; clypeal border curved and even; clypeus with a median and 2 lateral striae; frontal area depressed and with a median longitudinal striation; frontal carinae sub-parallel; antennal scapes slender, more robust apically, reaching or surpassing the occipital border by an amount less than first funicular joint; eyes with less than 20 facets.

In profile, pronotum and mesonotum strongly convex and descending sharply to the mesopropodeal impression, the latter with longitudinal rugulae. Propodeal spines sharp and pointed obliquely





52



dorsad; petiole slender anteriorly. Seen from above, postpetiole

sub-globular and slightly wider than the petiole.

Sculpture: Head reticulo-punctate; frontal area and clypeus

weakly reticulo-punctate and opaque; mandibles with piligerous

punctures; thorax, basal and declivious face of propodeum as well

as the petiole reticulo-punctate; dorsum of postpetiole and gaster

smooth and shining.

Pilosity: Hairs yellow, dorsal surface of head as well as the

ventral surface with numerous mixed short and long erect and semierect hairs; dorsum of thorax with perdominantly long erect hairs;

pedicel and gaster with long semi-erect numerous hairs.

Color: Concolorously yellow

Female and Male:

Unknown

Holotype: Major (collected on December 19, 1945 in Miami,

Florida by W. F. Buren).

Paratypes: 39 majors, 26 workers Discussion:

The type material comprises23 majors and 16 workers, collected on

December 19, 1945 in Miami, Florida, by W. F. Buren; 6 majors and 6 workers collected in 1975 in Baldwin County, Florida by D. P. Wojcik; 10 majors and 8 workers collected on November 25, 1975, in Naples, Florida by M. A. Naves.

The holotype and several paratypes will be deposited in the Museum of Comparative Zoology, Harvard University, Cambridge, Massachussetts. Paratypes will be deposited in the Nation Museum of Natural History, Washington, D.C., Florida State Museum, Department of Plant Industry, Gainesville, Florida; and in the W. F. Buren and M. A. Naves collections.





53



The species is named in honor of Dr. Robert E. Gregg, Professor

of Entomology, University of Colorado.

Gregg (1958) expended a great deal of effort trying to identify

a sample of this species that he and Mrs. Gregg collected in Brickel

Hammock, Miami, Florida. In his own words:

The specimens of sculptior from Miami, Florida (Buren det.) and the sample obtained from Brickel Hammock are
indistinguishable. Despite the similarities of the
scrobes and their sculpture, these ants are not identical,
however, with the examples of sculptior from Puerto Rico
and Martinique, nor with the types from St. Vincent. This
is evident especially from the smooth (almost sculptureless) and shining occipital lobes of the Florida ants. In
sculptior, the cephalic rugae and punctures completely
cover the head, making it opaque, except at the extreme posterior margin around the foramen which is smooth and
shining. In addition, the epinotal spines of the Brickel
Hammock ants are longer and sharper than those of sculptior.
These two samples of the flavens group in Florida cannot
represent P. flavens sculptior (from the West Indies), and
unless this subspecies is known from Florida by other
specimens, or is there but not yet collected, this discovery
requires a revision of the North American list. Pheidole
flavens sculptior must be dropped, and that is the plan
followed in this paper.

(p. 45)

Fig. 21 shows the head of P. greggi from the original specimens

collected at Miami, Florida, by W. F. Buren. This confirms Gregg's

conclusion that they are not conspecific with sculptior. The occipital

lobes of P. flavens sculptior from St. Thomas, Virgin Islands, also

collected by Buren on January 18, 1952, are shown in fig. 56.

The specimens collected at Naples were from a colony at the base

of a telephone pole on sandy soil close to the beach. The colony was

quite sizeable as in P. anastasii, P. floridana and P. moerens.




54




Although Gregg showed that Buren's original identification of sculptior from Florida specimens was in error, and that apparently sculptior does not occur in the U.S., he left unresolved the question of what name should be applied to the Florida species. The present studies indicate that it is a new, undescribed species distinct from P. flavens Roger (fig. 55), P. flavens sculptior Forel, P. floridana Emery, P. anastasii Emery and P. moerens Wheeler.

Much more study is needed in order to fully understand this

difficult group which has several related species, 8 subspecies and 13 varieties in the new world.

This new species differs from P. anastasii and P. floridana by

the shape of the postpetiole which is slightly larger than the petiole and does not have the distinct lateral connules. It differs from P. flavens by having the scrobe heavily reticulo-punctate, and the vertex not smooth and shining. It differs from P. flavens sculptior by having smooth and shining occipital lobes. It also differs from P. moerens by having a reticulo-punctate scrobe, which is opaque as well as the entire head.





55




12. Pheidole lamia Wheeler

P. lamia Wheeler, 1901. Amer. Nat. 35:534. Wheeler, 1908. Amer.

Mus. Nat. Hist., Bul. 24:477-478. Wheeler, 1910. Ants, pp. 212, 248.

Wheeler, 1926. Ants, pp. 212, 248. Smith, 1931. Ent. News 42:21-22. Creighton, 1950. Bull. Mus. Comp. Zool. 104:182. Smith, 1951. Cat. Hym. Amer. No. of Mex. U.S.D.A. Mon 2:803. Gregg, 1956. Ent. News

67:37-39. Gregg, 1958. N.Y. Ent. Soc. 66:23.

Type locality: Austin, Texas

Types : M.C.Z., A.M.N.H.

Range : North Florida to Texas Diagnosis:

P. lamia is a small, yellowish, shining species with a subcylindrical head in the major which is truncate anteriorly. The soldier of P. lamia has the most striking head shape in the genus. Discussion:

The head shape of P. lamia led Wheeler (1908) to suggest that it was a phragmotic ant. He also believed it to be a very rare species. Buren et al. (In Press), showed by laboratory and field observations that phragmosis does not occur in this species.

At Tall Timbers Research Station where this species is common, I had the opportunity to observe the biology of this interesting species. Colonies are monogynous and sometimes have up to about 200 majors and 1,000 workers. The nests are in the ground, usually in grassy areas as well as among taller vegetation. The nesting sites, at least at Tall Timbers, can be easily spotted in the morning due to the reddish clay





56




soil cast up in very small honey-combed hummocks. These clumps consist of fine soil particles and can have several openings connected to a vertical tunnel of about 2 mm in diameter. From the vertical tunnel branch single secondary galleries each directly connected to an oval chamber of about 0.5 to 1.5 cm in height and about .4 to 6 cm in length. Each colony has several chambers. The first is found usually over 40 cm deep. I have dug over 1.5 m deep following the main tunnel without reaching the end of it in four colonies. In the summer of 1973, I found the first female after digging seven colonies. In July, 1974, I found about thirty males in tunnels close to the soil surface. In July, 1975, I found 18 females inside a large chamber of about 50 cm deep. One mating pair that fell on the ground was collected in the late afternoon on July 15, 1974. The soil where this species nests usually has the first 30 cm predominantly sandy. Deeper there is a reddish clay where this species has its chambers.

This species usually forages in the morning or when there is no sunlight. Majors sometimes are found among debris with the workers. The pale yellow color of this species also could indicate that this species is nocturnal. The workers were observed carrying small seeds, small insects and mites to the nest. Some majors in the chambers might be repletes since they have quite extended gasters full of liquid. Buren et al. (In Press) show that this species has stylized defenses against Solenopsis (Diplorhoptrum) spp.





57




13. Pheidole littoralis Cole

P. sitarches littoralis Cole, 1952. Ann. Ent. Soc. Amer. 45:443. Gregg, 1958. N.Y. Ent. Soc. 66:30-39.

Type locality: Lido Beach, Sarasota, Florida

Type : A. C. Cole collection; M.C.Z.: Nat. Mus., A.M.N.H.

and W. S. Creighton collection. Both the A. C. Cole collection and W. S. Creighton collection are now at the Los Angeles County Museum of Natural History. Range : Central Florida

Diagnosis:

P. littoralis (fig. 20) has a reddish brown head and thorax, the gaster black and shining. It differs from P. sitarches, P. sitarches compestris and P. sitarches soritis by the presence of deep, large piligerous foveolae on the occipital lobes which also have no distinct transverse striae. Discussion:

P. littoralis is a distinct species related to the sitarches complex as Cole (1952) suggested. This species was collected nesting in white sand at the Archbold Biological Research Station, where it is very common. It was found nesting side by side with P. metallescens and P. adrianoi n. sp. The colonies seem to be quite small. Digging to 50 cm deep failed to produce any brood chamber or any sexual forms. At about 10 to 20 cm deep there was always a chamber full with small grass seeds and never more than four majors. The major has a large head which is disproportional considering the small size of the rest of the body. Majors were never observed outside the nest foraging. The small black, shining workers can easily be confounded with P. metallescens and P. adrianoi n. sp. workers.






58




14. Pheidole megacephala (Fabricius)

Formica megacephala Fabricius, 1793. Ent. Syst. 2:361.

P. gacep hala Emery 1921. Gen. Insect. Fasc. 174, p. 85. Phillips

1934. (Hawaii Univ.) Expt. Sta. Pineapple Prod. Coop. Assoc. Bul. 15:5-12.

Smith, 1936. Puerto Rico Univ., Jour. Agr. 20:843-844. Broekhuysen, 1948. Union So. Africa Dept. Agr. Bul. 266:1-40. Smith, 1951. Cat. Hym. Am.

No. of Mex. U.S.D.A. Mon 2:803. Wheeler and Wheeler, 1953. Ent. Soc.

Wash., Proc. 55:75. Brown, 1958. Act. Hym. 1:47. Gregg, 1958. N.Y.

Ent. Soc. 66:23. Brown, 1959. Bul. Ent. Res. 50:523. Weber, 1960. Ent.

Soc. Wash., Proc. 62:232. Taylor and Wilson, 1961. Psyche 68:143. Kempf, 1962.-Studia Ent. 5:18-19. Haskins and Haskins, 1965. Ecology 46:737. Wilson and Taylor, 1967. Pacific. Insects Monog. 14:46-48. Fluker, Huddleston, and Beardsley, 1968. Jour. Econ. Ent. 61:474. Fluker and Beardsley, 1970. Ent. Soc. Amer., Ann. 63:1290-1296. Kempf, 1972. Studia Ent. 15:196.

Oecophthora pusilla Heer, 1852. Stueck 66:15. Wheeler, 1922. Bull. Amer. Mus. Nat. Hist. 45:812.

P. laevigata Mayr, 1862. Verli. Zool. Bot. Ges. Wien 12:747. Mayr, 1870. Vehr. Zool. Bot. Ges. Wien 20:981.

Type locality: Isle de France (Mauritius)

Types : None in this country

Range : This species has become a world tramp; in Florida it apparently is largely confined to South Florida. Diagnosis:

P. megacephala (fig. 9), is a dark yellow to brownish-ferruginous

species that differs from P. morrisi by having distinct and sharp propodeal spines. It also differs from P. morrisi and P. dentata by having a cordate






59



head narrowing toward the mandibular insertion and the mesonotum is not distinctly transversely impressed.

Discussion:

P. megacephala is a well known world tramp (Gregg, 1958). It has been present in Florida for 43 years, at least, being reported by Smith (1933) from Everglades, Key West and St. Augustine. I collected this species in downtown Homestead, Florida on November 25, 1975 nesting under a paved sidewalk and at Archbold Biological Research Station. Buren observed large colonies of this species trailing in large numbers from the colony to nearby trees (Personal Communication). Its ecological impact has been studied in relation to another world tramp, Iridomyrmex humilis. Flucker et al. (1970) reported P. megacephala was losing territory to I. humilis in Hawaii, although it did not appear to be

evenly remotely approaching extermination. They also found that P. megacephala did not seem to occupy territory above 915 m, above which I. humilis was usually found.
In Florida the two species seem to occupy two distinct areas. I. humilis, although not common, occurs in the northernmost and northwest






60




areas of the state. P. megacephala, also not common, occurs in the southern area of the state. In the central area of the state I was able to find only P. megacephala. At least in Florida the two species are not sympatric and the inability of either of them to increase must be due to some other cause, possibly competition with Solenopsis invicta Buren or Solenopsis geminata (Fabricius).






61




15. Pheidole metallescens Emery

P. metallescens Emery, 1895. Zool. Jahrb., Abst. of System. 8:289, 294. Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:476-477. Smith,

1930. Fla. Ent. 14:3. Wheeler, 1932. N.Y. Ent. Soc. 40:7. Creighton,
S
19/0. Bull. Mus. Comp. Zool. 104:183. Smith, 1951. Cat. Hym. Am. No. of

Mex., U.S.D.A. Mon 2:803. Gregg, 1958. N.Y. Ent. Soc. 66:23. Wheeler

and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:12.

Type locality: St. George, Florida

Types : None in this country

Range : Gulf States

Diagnosis:

P. metallescens (fig. 16) is a small species usually with reddish

brown head and thorax and black gaster, although completely dark to black specimens have been found. It differs from P. greggi n. sp. and P. moerens by the distinct lateral connules on the postpetiole. It also differs from P. floridana and P. anastasii by the strong reticulation posteriorly on the head and from P. dentigula by having a reticulo-punctate propodeal base. The workers display strong, metallic, iridescent bluish reflections over most of the body.

Discussion:

P. metallescens is very common in Florida. The preferable habitat seems to be under the shade of trees or any high vegetation. Usually it is easy to find several small craters connected to a central vertical tunnel to the main chamber, rarely over 40 cm deep in the ground. It is a monogynous species. The majors usually forage along with the workers. They feed on small grass seeds and also scavenge dead arthropods.






62




16. Pheidole moerens Wheeler

P. moerens Wheeler, 1908. Bull. Amer. Mus. Nat. Hist. 24:136. Smith, 1936. Puerto Rico Univ., Jour. Agr. 20:842-843. Smith, 1967. Cat. Hym. Amer. No. of Mex. U.S.D.A. Mon 2 (Suppl. 2):354. Wheeler

and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:244. Kempf, 1972. Studia Ent. 15:196. Wojcik, Banks, and Buren, 1975. Coop. Econ. Ins. Rpt. U.S.D.A. 25449-5,6:906.

P. moerens dominicensis Wheeler, 1913. Bull. Amer. Mus. Nat. Hist. 32:241. Kempf, 1972. Studia Ent. 15:197. New Synonymy.

Type locality: Antilha Culebra

Types : M.C.Z.

Range : Southern Alabama, Florida, West Indies Diagnosis:

P. moerens (fig. 22) is a small reddish yellow to almost black species closely related to P. grggi n. sp., differing from it by the absence of the heavily punctate scrobe and in having a carinated shining clypeus. It differs from P. dentigula, P. anastasii, P. floridana and P. metallescens by the absence of distinct lateral connules on the postpetiole. Discussion:

P. moerens is a common species in north Florida. Its color was

observed in lab colonies to vary from reddish yellow to almost black due possibly to variation in the diet. Colonies fed with honey and seeds retained a lighter color than those fed with freshly killed house flies. Wheeler's subspecies dominicensis appears to represent only a color variant often found among the normal population. I have examined





63




the types of dominicensis and they appear identical with the types of moerens except in color. I do not consider dominicensis a valid taxon.

This species was found nesting in various places such as under boards, at base of oak trees and fence posts, along roots, under palm leaves, inside wall crevices, and rarely in the ground. Usually small chambers are constructed. It is a monogynous species which has a small blackish female. Its main flight is usually in July. Several queens may start founding a nest, but before the first brood emerges, the dominant female will have killed the others.

The chambers usually are built with small soil or debris particles and have small openings. A colony may have over 100 majors and over 300 workers. They feed on seeds and scavenge and prey on small dead or live arthropods. They forage very close to the nesting sites and sometimes a major is found foraging along with the workers.






64





17. Pheidole morrisi Forel

P. morrisii Forel, 1886. Soc. Ent. Belg., Bul. 30:46. Mayr, 1887. Verh. Zool. Bot. Ges. Wien 37:568. Forel, 1901. Soc. Ent. Belg.,

Ann. 45:350. Wheeler, 1904. Amer. Mus. Nat. Hist., Bul 20:302. Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:380. Wheeler, 1908. Amer. Mus.

Nat. Hist., Bul. 24:461. Smith, 1918. Ent. News 29:21. Davis and Bequaert, 1922. Brooklyn Ent. Soc., Bul. 17:8-9. Smith, 1924. Ent. News 35:53.

Smith, 1930. Fla. Ent. 14:3. Wheeler, 1932. N.Y. Ent. Soc. 40:7.

Cole, 1940. Amer. Midland Nat. 24:44-45. Creighton, 1950. Bull. Mus. Comp. Zool. 104:183-184. Smith, 1951. Cat. Hym. Amer. No. of Mex.,

U.S.D.A. Mon 2:803. Gregg, 1958. N.Y. Ent. Soc. 66:24. Wheeler and

Wheeler, 1960. Ent. Soc. Amer., Ann. 53:12.

P. morrisi var. vancea Forel, 1901. Soc. Ent. Belg., Ann. 45:351. Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20:302. Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:461. Smith, 1924. Ent. News 35:53. Smith, 1951. Cat. Hym. Amer. No. of Mex., U.S.D.A. Mon 2:803-804.

Type locality: Vineland, New Jersey Types : None in this country

Range : Southern New Jersey, south to Florida and eastern Gulf States.

Diagnosis:

P. morrisi (fig. 27) is a pale to darker yellowish species differing from P. dentata and P. vinelandica by the absence of distinct propodeal teeth or spines. It is larger than P. vinelandica and has a transverse mesonotal impression.





65




Discussion:

P. morrisi is another common species in Florida. It always nests in the ground and the colonies are quite large. It is a monogynous species and is primarily a scavenger.

Several nest founding females displayed singular habits in the

laboratory which may not have been noted previously in this genus. In constructing the claustral chamber each female used the soil dug from the bottom of the vertical passageway to form a continuous plug of the passageway above her rather than bringing the soil particles to the surface. The claustral chamber at the 20 to 30 cm level below ground thus is entirely sealed from the surface by a long plug. I interpret this habit as a defense mechanism against various predators during the claustral period. The first workers are reared in about 30 days and the first majors in about 50 days.

The colony increases in size rapidly. In 8 months it can have several hundred workers and majors. The workers forage alone, although the majors can help to bring food back to the colony once food is located. Workers can forage over 8 m from the nesting site. The species is mainly a scavenger but will also gather seeds.





66




18. Pheidole quiescens n. sp.

Diagnosis:

A small yellowish brown species (fig. 46, 47) parasitic in nests

of P. anastasii. Queens, soldiers and males are still unknown. Soldier caste probably lacking as in other species parasitic on Pheidole. Obviously closely related to its host and to other small species in this species group (P. floridana, P. moerens, and P. greggi n. sp.) but distinctive in its very long and delicate pilosity and its large and singularly shaped postpetiole which seems broadly and almost immovable articulated with the gaster. In behavior the species seems unique for its habit of standing motionless with open mandibles for long periods at a distance from the host workers and their activities. Description:

Major:

Unknown. Probably does not exist in this species.

Worker:

Measurements: Body length 1.90 to 2.08 mm. Head length 0.46

to 0.49 mm; head width 0.41 to 0.46 mm. Thoracic length 0.56 to

0.62 mm. Scape length 0.38 to 0.46 mm. Scape Index 89 to 96.

Structural characters: Head, excluding mandibles, slightly

longer than broad, posteriorly with a median sulcus and weak impression which forms weak occipital lobes. Frontal area small, slightly depressed, with two longitudinal striae. Clypeus with a fine median striation; anterior margin curved and slightly notched in the middle.

Frontal carinae short and divergent; antenae 12 segmented and with a

3 jointed club as in nonparasitic species; scapes long and slender,





67

the posterior third thicker, usually passing beyond the posterior margin of the head by an amount less than the length of first funicular segment. Mandibles slender, curved apically and with two large, sharp teeth, the remaining border with smaller sharp teeth; eyes convex, measuring less than 1/4 the head length and with approximately 22 facets. In profile,pronotum and anterior mesonotum convex; posterior of mesonotum more abruptly descending to the wide mesopropodeal impression which is slightly depressed below the anterior propodeal margin. Juncture of base and declivity of propodeum rounded; propodeal spines broad and heavy at base and pointed obliquely laterad. Petiole with gently sloping anterior dorsal face, the posterior face vertical, the node in profile very blunt apically.

Postpetiole, seen from above, slightly wider than long, two

times or more wider than petiole, the sides slightly angular but not forming distinct conules, broadest in the posterior half, gaster truncate at base and always noticeably larger than the head.

Sculpture: Head including frons and clypeus opaque and finely reticulo-punctate; mandibles smooth and shining. Frontal area small and finely reticulo-punctate. Sides of thorax and petiole, subopaque and reticulo-punctate. Postpetiole weakly reticulo-punctate and weakly shining. First gastric tergite and usually the postpetiolar dorsum with opalescent reflections and subopaque.

Pilosity: Hairs yellow, long, very slender and pointed, some

measuring up to 0.15 mm, numerous all over the body, even the ventral surface of the head; short and sparse hairs on the legs; pubescence limited to the antennae, absent on gaster and other regions of body.






68




Color: Light reddish brown, appendages lighter.

Female and Male:

Unknown

Holotype: 1 worker, collected at Archbold Research Station.

Paratypes: 3 workers collected in two colonies of the host

Pheidole anastasii at the Archbold Biological Research Station in

Lake Placid, Florida, by M. A. Naves in August, 1975.

The holotype will be deposited in the Museum of Comparative

Zoology, Harvard University, Cambridge, Massachussetts. Paratypes

will also be deposited in the National Museum of Natural History,

Washington, D.C., the Florida State Museum, Department of Plant

Industry, Gainesville, Florida, and M. A. Naves Collection. Discussion:

Wheeler (1904) described two parasitic species, Epipheidole inquilina and Sympheidole elecebra. E. inguilina was parasitising colonies of P. pilifera coloradensis and S. elecebra was parasitising colonies of P. ceres. Both parasitic species were collected in Colorado and up to now have been the only known ant parasites for Pheidole in the U.S.

E. inquilina was described from males and females collected in three colonies of its host. The lack of workers and absence of the host queen led Wheeler to the assumption that the host queen is dispatched by its own workers once the parasite had secured entry into the nest, and that this species deserved placement under a separate genus. Later Dr. M. R. Smith (1941) announced the discovery of a single worker of inquilina, possibly jeopardizing the status of Epipheidole as separate from Pheidole.






69




S. elecebra was described from one queen and 18 males. Later another queen was found. The worker caste, if any exists, is still unknown.

Brown (1973) listed both Epipheidole and Sympheidole as synonymous with Pheidole, but these listings were not annotated and the validity of these synonymizations is in doubt.

P. quiescens, in contrast with E. inquilina and S. elecebra, still produces its worker caste. Several attempts to collect sexual forms of P. quiescens have failed. In two cases parasitised colonies still retained the host queen.

One of these colonies was brought to the lab for observation. It

was noted that the parasite workers essentially stand in the colony doing nothing for long periods of time. They obtain their food by trophallaxis from the host workers. They always stand with the mandibles open, possibly in a defensive position, at a distance from the host workers. If the colony happens to be disturbed the parasite will run away, apparently seeking refuge. Twice they were observed helping to move brood from one place to another but never were observed doing any other useful work. The interspecific relationship seems to be inquilinism which means that the parasite spends its whole life cycle in the host colony.

Subsequent collections of host workers and parasite workers were added to the lab colony and no rejection of the newcomers was ever observed. Later on, the host queen died and another queen of P. anastasii collected in Gainesville, Florida was readily accepted by the colony. The host is very well distributed all over the state, but collections throughout the state failed to have parasites, although P. quiescens seems to be common in colonies of its host at Archbold Biological Research Station.






70



Permanent social parasitsm with a useless worker caste is already

known to occur among ants. Buren (1942), when describing Formica reflexa suggested that the consistant presence of workers of both F. subsericea, the host, and of small, timid workers of F. reflexa in every colony that he found, was possibly a case of F. reflexa being a permanent inquiline. King (1949) presented further evidence confirming Buren's suggestions. King and Sallee (1951) suggested that the absence of pure colonies of F. reflexa confirmed Buren's suggestions.

P. guiescens seems to fit Buren's and King's descriptions of a

permanent social parasite which has retained a useless worker caste. P. quiescens was collected several times and always the host-workers outnumered the parasite at least several to one. The new species was never found as a pure colony.





71




19. Pheidole tysoni Forel

P. tysoni Forel, 1901. Soc. Ent. Belg., Ann. 45:348. Wheeler,

1904. Amer. Mus. Nat. Hist. 20:302. Smith, 1918. Ent. News 29:22.

Smith, 1924. Ent. News 35:78. Dennis, 1938. Ent. Soc. Amer. 31:282, 304. Wesson and Wesson, 1940. Amer. Midland Nat. 24:92. Cole, 1940. Amer. Midland Nat. 24:42, 45. Creighton, 1950. Bull. MusComp. Zool. 104:191. Smith, 1951. Cat. Hym. Amer. No. of Mex., US.D.A. Mon 2:805.

Gregg, 1958. N.Y. Ent. Soc. 66:32.

Type locality: Mt. Mitchell, North Carolina

Types : A. M. N. H., M. C. Z.

Range : North Florida, Georgia, western North Carolina, southwestern Virginia and eastern Tennessee. Diagnosis:

P. tysoni is a small yellowish species closely related to P. vinelandica, differing from it by the teeth on the anterior ventral margin of head. The minor has the propodeal pleurae largely free from sculpture and strongly shining.

Discussion:

P. tysoni was collected once in a soybean field at Tall Timbers

Research Station in 1973. The colony was quite large and the main chamber was about 50 cm deep in the ground. Only minors were foraging. This is the only record known for the state and I am assuming that P. tysoni is rare in Florida.






72




20. Pheidole vinelandica Forel

P. bicarinata race vinelandica Forel. 1886. Ann. Soc. Ent. Belg. 30:45.

P. vinelandica Mayr, 1886. Verh. Zool. Bot. Ges. Wien 36:458. Emery, 1895. Zool. Jahrb. Syst. 8:292. Forel, 1901. Am. Soc. Ent. Belg. 45:348. Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 24:458.

Smith, 1918. Ent. News 29:22. Davis and Beguaert, 1922. Brook. Ent. Soc. 17:9. Smith, 1924. Ent. News, 35:54. Hayer, 1925. Ent. News 36:42 Cole, 1940. Amer. Midland Nat. 24:42. Wesson and Wesson, 1940. Amer. Midland Nat. 24:92-93.

P. (Allopheidole) vinelandica Forel, 1912 Mem. Soc. Ent. Belg. 19:237.

P. vinelandica laeviuscula Emery, 1895. Zool. Jahrb. Syst. 8:292. Smith, 1951. Cat. Hym. Amer. No. of Mex. U.S.D.A. Man. 2:805.

P. vinelandica subsp. buccalis Wheeler, 1908. Bull. Amer. Mus. Nat. Hist. 24:454. Smith, 1951. Cat. Hym. Amer. No. of Mex. U.S.D.A.

Mon 2:805.

P. vinelandica longula var. castanea. Wheeler, 1915. Amer. Mus. Nat. Hist. 34:405. Cole, 1937. Ent. News. 48:100.

P. bicarimata buccalis Creighton, 1950. Bull. Mus. Comp. Zool. 104:171. Cole, 1953. Tenn. Ac. Sc. 28:297.

P. bicarinata vinelandica Creighton. 1950. Bull. Mus. Comp. Zool. 104:17a. Cole, 1956. Tenn. Acad. Sc. 31:114. Gregg, 1958. N.Y. Ent.

Soc. 66:18, 36. Reid and Nugara, 1961. Jour. Parasitology 47:885-889. Smith, 1965. U.S. Dept. Agr., Tech. Bul. 1326:25-26.






73




P. vinelandica vinelandica. Smith, 1951. Cat. Hym. Amer. No. of Mex. U.S.D.A. Mon. 2:806.

Type locality: Vineland, New Jersey.

Types : Cotypes, A.M.N.H.

Range : Southeastern and Gulf States and along the east coast to New Jersey

Diagnosis:

P. vinelandica is a small yellowish species differing from P. bicarinata by having the basal face of propodeum in the major largely punctate (fig. ) and having a longitudianl striation on the frontal area (fig. 66). It differs from P. tysoni by the absence of teeth on the anterior ventral margin of the head.

Discussion:

I had the opportunity to study specimens of P. vinelandica from New Jersey, North Carolina, South Carolina, Georgia, Alabama, and Texas. I have also collected this species at Tall Timbers Research Station as well as in Putman Co., Florida. I have compared these with specimens of P. bicarinata (fig. 24, 64, 65) from Illinois in the M.C.Z. I also have studied specimens of bicarinata collected at Colorado Springs, Colorado; Ames and Lewis, Iowa; Devils Tower, Wyoming; Ortonville, Minnesota, and Corinne, Utah. It was evident that the two forms are not conspecific and have constant morphological differences, emphasizing that P. vinelandica deserves full species rank. Gregg (1958) included the var. nebrascensis Forel and the var. huachucana Smith as synonymus of vinelandica. I have not studied the types of these taxa but am doubtful about their inclusion under vinelandica rather than bicarinata. Records of vinelandica from





74




North Dakota and Colorado also seem doubtful, and seem much more likely to be misdetermined records of bicarinata.

P. vinelandica is found nesting in grassy areas at Tall Timbers

Research Station. The minors and majors are easily seen foraging close to the nesting site. The nest has several exits. They were observed carrying seeds and small dead arthropods.





75








SPECIES NOT PROPERLY INCLUDED IN THE FLORIDA PHEIDOLE FAUNA

1. Pheidole pilifera (Roger)

Leptothorax pilifera Roger, 1863. Berl. Ent. Zeitschr., 7:180.

P. pilifera var. simulans Wheeler, 1908. Bull. Amer. Mus. Nat. Hist., 24:436.

P. pilifera subsp. septentrionalis Wheeler, 1908. Bull. Amer. Mus. Nat. Hist. 24:436.

P. pennsylvanica Roger, 1863. Berl. Ent. Zeitschr., 7:199. Mayr, 1870. Verh. Zool-Bot. Ges. Wien, 20:981. Mayr, 1886. Verh. Zool-Bot.

Ges. Wien, 36:455. Mayr, 1877. Verh. Zool-Bot. Ges. Wien, 37:588.

P. pilifera Emery, 1895. Zool. Jahrb. Syst. 8:290. Creighton, 1950. Bull. Mus. Comp. Z-ol. 104:184-186. Van Pelt, 1956. Amer. Midland Nat.

56:377, 834. Smith, 1958. Cat. Hym. Amer. No. of Mex., U.S.D.A. Mon.

2 (Suppl. 1):123. Gregg, 1958. N.Y. Ent. Soc., 66:24-25.

Type locality: Pennsylvania

Types : None in this country

Range : Massachussetts to North Carolina and West to Iowa and Nebraska

Diagnosis:

P. pilifera differs from P. carrolli n. sp. and P. littoralis Cole by having the occipital rugae of the major notably reticulate and often coarse; its occipital lobes are not well rounded and pronounced. Discussion:

Attempts were made by the author to find P. pilifera in Florida as

reported by Van Pelt (1956). Dr. Van Pelt, however, expressed some reservations about the correctness of this identification. No voucher specimens





76




of P. pilifera are presently available from Dr. Van Pelt's work. Although it seems possible that P. pilifera occurs in Florida, I am unconvinced that there is any direct evidence. I believe all reported specimens were probably P. carrolli n. sp., P. littoralis Cole, or other species.













PLATE 1



Figure i. Head of major of P. lamia Wheeler (33X) Figure 2. Scape of major of P. bureni n. sp., a member
of the crassicornis complex (81X)

Figure 3. Head of major of P. diversipilosa Wheeler (45X) Figure 4. Gaster of major of P. diversipilosa (47X) Figure 5. Head of major of P. crassicornis Emery (43X) Figure 6. Gaster of major of P. crassicornis (58X)






78





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/ b IC1"4 i

C ;i I 1: :e

Y I!
i: .-i :r i;all t
,i Ir
"


*i ?. ,. :4

~ .

=:*~--,

~l;p~YTf+L~f~ 'I ~

,, r
t r
b I.il ~"i ~ \ r \ ~':mR>
~ r
~cr f
I
'i
Y ~II
1~, ( 7c, s ...
i ''
3.
r ";,
i r

*





~ 1






;Ct~t;, r ~r
I ~ ss-~e~~rui~~~u\~h:' Ct
I iS'

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3 ,
I-rs .\

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r;:
r '


5)jl y ~














PLATE 2

Figure 7. Scape of major of P. fallax obscurithorax
Santschi (81X)

Figure 8. Head of major of P. fallax obscurithorax (40X) Figure 9. Head of major of P. megacephala (Fabricius)
(40X)

Figure 10. Pedicel of major of P. floridana Emery (128X) Figure 11. Propodeum, pedicel and gaster of major of P.
metallescens Emery (74X)

Figure 12. Postpetiole of major of P. littoralis Cole (119X)



















__ /. I-Iowl 00,



14'C

Ir $rip~;














i'i .4f- (...........i















PLATE 3

Figure 13. Head of major of P. dentigula M. R. Smith
(64X)

Figure 14. Head of major of P. carrolli n. sp. (29X) Figure 15. Pedicel of major of P. metallescens Emery
(163X)

Figure 16. Head of major of P. metallescens (62X) Figure 17. Head of major of P. anastasii Emery (68X) Figure 18. Postpetiole and gaster of P. anastasii (62X)








82







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\g ( IBil r


ii

.

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,
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~:; ~* ~~r! :d:~ ~Ic 1, vz~s~ c ri c, "P1

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ji
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PLATE 4

Figure 19. Head of major of P. floridana Emery (72X) Figure 20. Head of major of P. littoralis Cole (37X) Figure 21. Head of major of P. greggi n. sp. (68X) Figure 22. Head of major of P. moerens Wheeler (58X) Figure 23. Clypeus and frontal area of major of P.
vinelandica Forel (186X)

Figure 24. Head of major of P. bicarinata Mayr (53X)





84












191















7 I














PLATE 5

Figure 25. Head of major of P. adrianoi n. sp. (72X) Figure 26. Thorax, pedicel and gaster of major of P.
adrianoi n. sp. (50X); white legs are artifacts due to electron overcharge.

Figure 27. Head of major of P. morrisi Forel (37X) Figure 28. Thorax of major of P. morrisi (56X) Figure 29. Head of major of P. dentata Mayr (48X) Figure 30. Teeth on the anterior ventral head margin of
P. anastasii Emery (123X)







86









tiff
















"al.
M I







































I~vr
-u i
'de












IY
I '~E E'








m














PLATE 6

Figure 31. Head of worker of P. lamia (97X) Figure 32. Dorsal view of worker of P. vinelandica
(62X)

Figure 33. Dorsal view of worker of P. adrianoi n. sp.
(95X)

Figure 34. Worker of P. megacephala (36X) Figure 35. Thorax and pedicel of the worker of P. fallax
obscurithorax (41X)

Figure 36. Head of worker of P. fallax obscurithorax
(86X)




Full Text
PLATE 13
Figure 67. Maps of Florida and the United States
showing:
Tall Timbers Research Station
O Gainesville
T Archbold Biological Research Station
Locations where collections of Pheidole
spp. were made in Florida
V -
Alice, Texas


LIST OF FIGURES Continued
Figure
61 Gaster and pedicel of P_. bureni n. sp. (40X) .
62 Head and pronotum of worker of P_. bureni n. sp.
(52X)
63 Dorsum of propodeum of maior of P. vinelandica
(272X) 7
64 Thoracic dorsum of P. bicarinata (71X)
65 Postpetiole of P_. bicarinata (186X)
66 Head of P. vinelandica (66X)
67 Map of Florida and the United States
Page
98
100
100
100
100
102
xi


80


34
edentate, except for a small basal tooth-like lobe. Eyes small,
measuring about 0.25 mm in diameter with over 60 facets and located
on the anterior third of the head. Seen from the side, thorax
smaller than head; pronotum and anterior mesonotum strongly convex;
posterior of mesonotum descending to the mesopropodeal impression,
forming an obtuse angle with the propodeal base; base of propodeum
slightly shorter than declivity; propodeal spines short and project
ing dorsad; petiole, in profile, stout and with a short anterior
peduncle, anterior face gently curved and posterior face short; apex
of node weakly angular to moderately convex; seen from above the
sides of the petiole parallel and the node notched in the middle;
postpetiole in profile strongly convex dorsally, slightly concave
and much shorter ventrally; seen from above, postpetiole two times
or more wider than petiole and with lateral connules; gaster oval
and much smaller than the head.
Sculpture: Cephalic longitudinal rugae extending posteriorly
and somewhat divergently; the lateral rugae usually do not reach
the occipital lobes; occipital lobes smooth and shining laterally
and dorsally; rugae present on the genae, extending longitudinally;
posterior to the eyes the surfaces smooth and shining; head posteriorly
with coarse circular piligerous punctures. Dorsum of pronotum and
mesonotum with longitudinal or transverse rugae or both, the inter
stitial areas with fine reticulate punctation. The mesopropodeal
impression wide and with longitudinal rugules, the interstitial
areas smooth and weakly shining; the propodeal base and declivity


10
and mesonotum strongly sculptured with transverse and
sometimes with longitudinal rugae; dorsal crest of petiole
notched. (fig. 14) carrolli n. sp.
b) Without all the above characters 9
9a) Erect gastric hairs sparse, tapered and almost all of the
same size; posterior part of the head covered with sculpture;
anterior ventral margin of the head without teeth; body
sometimes with distinct violaceous or bluish metallic
reflections. (fig. 15, 16) metallescens Emery
b) Gaster ocvered with long and short hairs; posterior part
of the head smooth and shining (fig. 17); anterior ventral
margin of the head with two or three teeth, the mesal
tooth usually short and blunt. (fig. 30) 10
10a) Dorsal surface of first gastric tergite finely and densely
granulse, opalescent and opaque; dorsal areas of pronotum
punctate and sometimes with several short and fine transverse
striae; dorsal area of post-petiole punctate.
(fig. 17, 18) anastasii Emery
b) Dorsal surface of first gastric tergite smooth and shining;
dorsum of pronotum partially weakly punctate but also
interdispersed with smooth and shining areas; dorsal area
of post-petiole weakly punctate and shining.
(fig. 10, 19) floridana Emery
11a) Head covered with longitudinal striae; the dorsal area of
the pronotum and usually the mesothorax reticulo-punctate
and with transverse striae or sculpture, (fig. 20)
12


19
area of propodeum reticulate and shining; declivous face with trans
verse striae and usually the interstitial areas finely reticulo-
punctate and shining; several fine interspinal striae present; dorsum
of postpetiole weakly sculptured or almost smooth and shining.
Pilosity: Hairs yellow, fine, numerous and of various lengths,
measuring 0.07 to 0.18 mm; dorsal and ventral surface of head with
semi-erect short and long hairs; dorsum of pronotum and mesonotum
with numerous predominantly long erect hairs; propodeal base with
few scattered short hairs; pedicel with numerous semi-erect long
hairs; gaster with numerous predominantly long semi-erect hairs, most
of them blunt apically. Pubescence on antennae, sparse on legs and
absent on gaster.
Color: Head and gaster ebony black; thorax and legs a little
lighter.
Worker:
Measurements: Body length 1.50 to 1.60 mm. Head length 0.44 to
0.51; head width 0.38 to 0.41 mm. Thoracic length 0.46 to 0.51 mm.
Scape length 0.40 to 0.46 mm. Scape Index 97 to 106.
Structural characters: Head, excluding mandibles, slightly longer
than broad; occipital border slightly concave medianly; frontal area
shallowly depressed; frontal carinae straight, short and slightly
divergent; scapes surpassing the occipital lobes by an amount less
than the first funicular joint; mandibles slender, apically with
two sharp teeth, the remaining masticatory border with several smaller
denticles.


28
mesopropodeal impression depressed and shining; prododeal base
weakly reticulo-punctate and shining; propodeal declivity smooth
and strongly shining; petiole and postpetiole finely reticulo-
punctate and opaque; gaster smooth and shining.
Pilosity: Hairs yellow and pointed, measuring up to 0.26 mm;
vertex with two long hairs on each side and several along the frontal
carinae; anterior ventral face of head with semi-erect short hair;
posterior of head and mandibles and scapes with fine appressed
pubescence; a few erect hairs on the scapes; dorsum of pronotum
and mesonotum with a few mixed short and long erect hairs; scattered
appressed pubescence on pronotum; each side of the petiolar node
with long semi-erect hairs; postpetiole with several semi-erect
long hairs; first gastric tergite with numerous fine appressed
pubescent hairs; ventrally the posterior half of gaster with mixed
short and long erect and semi-erect hairs; appendages with sparse
appressed pubescence; funiculus with very fine pubescence.
Color: Reddish brown, the gaster usually darker.
Worker:
Measurements: Body length 3.10 to 3.20 mm. Head length 0.77
to 0.79 mm; head width 0.70 to 0.77mm. Thoracic length 1.05 to 1.13.
Scape length 0.82 to 0.90. Scape Index 106 to 114.
Structural characters: Head, excluding mandibles, a little
longer than broad; occipital border flat to slightly concave; sides
strongly convex; frontal area broader than long and shallow; frontal
carinae short and parallel; scapes surpassing the occipital border


PLATE 3
Figure 13.
Figure 14.
Figure 15.
Figure 16.
Figure 17.
Head of major of P. dentigula M. R. Smith
(64X)
Head of major of P. carrolli n. sp. (29X)
Pedicel of major of P. metallescens Emery
(163X)
Head of major of _P. metallescens (62X)
Head of major of P_. anastasii Emery (68X)
Figure 18.
Postpetiole and gaster of P. anastasii (62X)


11
b) Posterior areas of the head smooth and shining; the pronotum,
seen from above, smooth and very shining 14
12a) Head usually measuring over 1.4 mm in length and 1.1 mm
in width; sides of the head sub-parallel; scape length
shorter than half the head length (Scape Index around 40);
occipital lobes with distinct circular piligerous punctures.
(fig. 12, 20) littoralis Cole
b) Head measuring less than .9 mm in length and width; occipital
lobes smooth and shining 13
13a) Frontal area concave and bearing 1 or 2 longitudinal
striae. Scrobes strongly punctate and opaque; occipital
lobes smooth and weakly shining; on occiput interstitial
spaces between striae reticulo-punctate. (fig. 21).... greggi n. sp.
b) Frontal area concave, without longitudinal striae; posterior
half of the scrobe weakly punctate and shining; occipital
lobes and most of the head shining; on front, interstitial
spaces between striae smooth and shining. (fig. 22). moerens Wheeler
14a) Anterior clypeal margin bluntly bidentate, with deep
median notch; anterior ventral margin of head without
teeth. (fig. 23) vinelandica Forel
b) Anterior clypeal margin not bidentate and without notch;
anterior ventral margin of the head usually with two or
three teeth; if teeth are absent, shining black species 15
15a) Mesonotum without transverse impression, forming in profile
a continuous smooth curve with the pronotum. (fig. 26) 16


This dissertation was submitted to the Graduate Faculty of the College
of Agriculture and to the Graduate Council, and was accepted as partial
fulfillment of the requirements for the degree of Doctor of Philosophy.
December, 1976
Dean, Graduate School


4
about two or three hundred individuals in a fully developed
nest. Even in the case of the species which produce comparatively
large nests (morrisi, hyatti, desertorum, etc.) a colony of more
than two or three thousand individuals would be exceptionally
large. By far the majority of our species nest in soil. The
nest may be built under a stone or in open soil without a cover
ing object. In the latter case there is often a mound or craters
of excavated soil surrounding the nest entrance. The eastern
species dentata will nest in rotten logs as well as soil but
such flexibility in nesting habit is exceptional.
(p. 162)
Gregg (1958), published a new revision in which he included the
description of ten new taxa. He also placed ten taxa in synomymy and the
status of eight taxa was revised. Later, Gregg (1969) described another
new species, P_. clementensis.
Smith (1967) reported P_. moerens from Alabama as yet another exotic
species. This was the 71st Pheidole taxon reported for North America north
of Mexico. Wojcik et al. (1975) reported that this species was also
present in Florida.


BIOGRAPHICAL SKETCH
Marcio Antonio Naves was born February 26, 1943, in Lavras, Minas
Gerais, Brazil. He is married to Lucilia de Rezende Naves, and has a
son, Adriano de Rezende Naves.
In December, 1970, he graduated from Escola Superior de Agricultura
de Lavras, Lavras, M. G., where he received a BS in Agronomy. In 1971
he was hired as coordinator of the Agreement ESAL/DNOS for the irrigation
project of the valleys of Careacu and Bela Vista, M. G. In June, 1974,
he received the degree of Master of Science from the Entomology and
Nematology Department of the University of Florida, Gainesville, Florida.
He has been a member of the American Entomological Society and Florida
Entomological Society, and Georgia Entomological Society.
In 1974, he began to work towards the Degree of Doctor of Philosophy
with a major in Entomology and minor in Plant Pathology.
105


94


PLATE 7
Figure
37.
Thorax of the worker of
Figure
38.
Head, thorax and pedicel
P. dentata (72X)
Figure
39.
Thorax of the worker of
Figure
40.
Head of the worker of P.
Figure
41.
Thorax of the worker of
(106X)
Figure
42.
Head of the worker of P.
. morrisi (72X)
of the worker of
dentata (79X)
crassicornis (54X)
1. metallescens
littoralis (113X)


29
by an amount greater than the first funicular joint; mandibles slender,
curved apically and with two sharp -teeth, the rest of the border
with several small denticles; anterior clypeal border convex; eyes
measuring 0.18 mm in diameter and with more than 65 facets.
In profile, pronotum and anterior mesonotum strongly convex;
mesonotum with a wide transverse impression; propodeal base longer
than the declivity; propodeal spines short and obliquely pointed
dorsad; petiole stout, the node strongly convex in profile; post
petiole strongly convex dorsally in profile and flat ventrally.
Seen from above, postpetiole 1.5 times wider than long and the
sides strongly convex; gaster truncate at the base.
Sculpture: Head finely reticulo-punctate except the irons and
vertex which have smooth and shining areas; frontal area smooth and
shining; clypeus with a median carina and usually smooth or weakly
reticulo-punctate; genae with several short longitudinal striae
emerging from anterior reticulation.
Pleurae of throax reticulo-punctate; petiole and postpetiole
finely reticulo-punctate; gaster smooth and shining.
Dorsum of mesonotum with small smooth and shining areas; basal
face of propodeun strongly reticulo-punctate and the declivity
weakly reticulate and shining.
Pilosity: Hairs yellow, numerous, measuring up to 0.20 mm;
mixed short and long semi-erect hairs present on head; erect, short
and long hairs on the dorsum of thorax; two long semi-erect hairs on
the pedile; numerous short and long, erect and semi-erect hairs on
postpetiole and gaster.


43
7. Pheidole dentigula M. R. Smith
P. dentigula Smith, 1927. Ent. News 38:310. Smith, 1928. Ent.
News 39:245-246. Cole, 1940. Amer. Midland Nat. 24:42, 45. Smith,
1944. Fla. Ent. 27:14. Creighton, 1950. Bull. Mus. Comp. Zool, 104:
178. Gregg, 1958. N.Y. Ent. Soc. 66:21. Wheeler and Wheeler, 1960.
Ent. Soc. Wash., Proc. 62:14.
Type locality: Mississippi State University, Starkville, Mississippi.
Types : Cotypes, National Museum
Range : Tennessee, southward through Alabama and Mississippi,
and north Florida
Diagnosis:
P_. dentigula (fig. 13) is a small yellowish species that differs
from _P. greggi, P_. anastasii, P_. floridana, P_. tysoni, P_. bicarinata
and Ph moerens by the reticulated, opaque occipital lobes. It differs
from P_. metallescens by having a smooth and shining propodeal base and
lacks metallic reflections. The teeth on the anterior ventral border of
the head are prominent.
Discussion:
Smith (1944) reported P. dentigula from Wakulla County (Florida).
Dr. John F. Carroll also collected this species in Marion County (Florida)
on April 3, 1973. According to Smith (1944) this species nests in the
soil and in well rotted stumps in wooded areas, especially where the soil
conatins considerable hummus.


TABLE OF CONTENTS Continued
Page
5. < Pheidole crassicornis Emery 39
Diagnosis 39
Discussion 39
6. Pheidole dentata Mayr 40
Diagnosis 41
Discussion 41
7. Pheidole dentigula M. R. Smith 43
Diagnosis 43
Discussion 43
8. Pheidole diversipilosa Wheeler 44
Diagnosis 44
Discussion 44
9. Pheidole fallax obscurithorax Santschi 46
Diagnosis 46
Discussion 46
10. Pheidole floridana Emery 48
Diagnosis 48
Discussion 48
11. Pheidole greggi n. sp 49
Diagnosis 49
Description 49
Major 49
Measurements 49
Structural characters 49
Sculpture 50
Pilosity 51
Color 51
Worker 51
Measurements 51
Structural characters 51
Sculpture 52
Pilosity 52
Color 52
Female and Male 52
Holotype 52
Paratypes 52
Discussion 52
12. Pheidole lamia Wheeler '55
Diagnosis 55
Discussion 55
13. Pheidole littoralis Cole 57
Diagnosis 57
Discussion 57
14. Pheidole megacephala (Fabricius) 58
Diagnosis.. 58
Discussion 59
vi


la
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
LIST OF FIGURES Continued
Page
84
Head of major of P. greggi n. sp. (68X)
Head of major of EL moerens Wheeler (58X) 84
Clypeus and frontal area of major of P_. vinelandica
Forel (186X) 84
Head of major of P_. bicarinata Mayr (53X) 84
Head of major of P_. adrianoi n. sp. (72X) 86
Thorax, pedicel and gaster of major of _P. adrianoi
n. sp. (50X); white legs are artifacts due to electron
overcharge 86
Head of major of P_. morrisi Forel (37X) 86
Thorax of major of P_. morrisi (56X) 86
Head of major of P. dentata Mayr (48X) 86
Teeth on the anterior ventral head margin of PL
anastasii (Emery (123X) 86
Head of worker of P.. lamia (97X) 88
Dorsal view of worker of PL vinelandica (62X) 88
Dorsal view of worker of P_. adrianoi n. sp. (95X) 88
Worker of P. megacephala (36X) 88
Thorax and pedicel of the worker of P. fallax
obscurithorax (41X) 88
Head of worker of PL fallax obscurithorax (86X) 88
Thorax of the worker of PL morrisi (72X) 90
Head thorax and pedicel of the worker of P. dentata
(72X) r; 90
Thorax of the worker of PL dentata (79X) 90
IX


90


22
Color: Reddish brown; the gaster darker.
Male:
Unknown.
This species is named in honor of my beloved first born son,
Adriano de Resende Naves.
Holotype: Major from Gainesville, Florida, collected by M. A.
Naves.
Paratypes: The type material comprises 30 majors and 60 workers
collected in Gainesville, Florida from several colonies on August 13,
1975 and 25 majors and 33 workers from several colonies from Archbold
Biological Research Station at Lake Placid, Florida, in August, 1975.
All were collected by M. A. Naves.
The holotype and several paratypes will be deposited at the Museum
of Comparative Zoology, Harvard University, Cambridge, Massachussetts.
Paratypes will also be deposited in the National Museum of Natural
History, Washington, D.C., Florida State Museum, Department of Plant
Industry, Gainesville, Florida and in the W. F. Burn and M. A. Naves
collections.
Discussion:
This species is fairly common in Florida. Its range seems to be from
central to northern Florida. It may also occur in southern Georgia. Its
preferable habitat seems to be sandy soils, in clear areas among trees,
the same habitat as in Ij. metallescens, with which it is very similar to
the naked eye.
The colonies have about 60 majors and more than 300 workers. The main
chamber is located 30 to 40 cm deep in the soil and is directly connected
to a vertical tunnel of about 1 to 2 mm in diameter, which usually is the


39
5. Pheidole crassicornis Emery
P. crassicornis Emery, 1985. Zool. Jahrb., Abt. F. System. 8:289,
296. Forel, 1901. Soc. Ent. Belg., Ann. 45:350. Wheeler 1904. Amer.
Mus. Nat. Hist., Bui. 20:302. Creighton, 1950. Bull. Mus. Comp. Zool.
104:175-176. Gregg, 1958. N.Y. Ent. Soc., 66:20.
Type locality: Charlotte, North Carolina
Types : Cotypes, M. C. Z. and A. M. N. H.
Range : Southeastern States
Diagnosis:
P_. crassicornis (fig. 5) differs from P_. diversipilosa and Py cras
sicornis tetra by the absence of erect and semi-erect hairs on the gaster.
It also differs from P. bureni n. sp. by the convex occipital lobes and
the occipital impression which is not sharply V shapped. P_. crassicornis
can be easily distinguished from all Florida Pheidole other than diversipilosa
by the basally strongly bent and flattened scapes.
Discussion:
The finding of P. crassicornis at Gainesville and at Tall Timbers
Research Station will help the understanding of this group because it is
sympatric with P_. diversipilosa at Tall Timbers. P. crassicornis is
apparently confined to the northern part of Florida where it occurs in open
areas, usually among trees. Its nests seem to be in deep soil. After digging
as deep as 60 cm, I was unable to find any chambers of three colonies. The
opening at the soil surface is hard to see. The typical craters common
to other species were never observed. The major and workers were seen carry
ing live termites and other small dead arthropods. The workers and a few
majors were observed foraging approximately 4 meters or more from the nest
opening.


12
b) Mesonotum, with a well developed transverse impression 17
16a) A yellow species; frontal area and middle of front smooth
and shining without striae; anterior ventral margin with
3 teeth; thorax smooth and shining tysoni Forel
b) A black species; anterior half of head with longitudinal
striae; dorsum of pronoturn smooth and shining; pleurae
. of pronotum, mesonotum, and propodeum, heavily sculptured.
(fig. 25, 26) adrianoi n. sp.
17a) Propodeum angular at the junction of the basal and declivous
faces, but the angles not produced into distinct teeth or
spines. (fig. 27, 28) morrisi Forel
b) Propodeum armed with distinct teeth or spines.
(fig. 29)
dentata Mayr


51
Sides of thorax, petiole and postpetiole reticulo-punctate,
opaque to subopaque. Pronotal dorsum with transverse irregular
striae; the interstitial areas weakly reticulo-punctate to smooth
and weakly shining; mesopropodeal impression wide and with longi
tudinal rugulae; the interstitial areas smooth and shining; propodeal
base reticulo-punctate and opaque; postpetiolar dorsum smooth and
shining; gaster smooth and shining.
Pilosity: Erect hair yellow, pointed, numerous; both short
and long hairs on head, thorax, pedicel and gaster; pubescence
sparse on antennae and legs, absent on gaster.
Color: Concolorously yellow
Worker:
Measurements: Body length 1.50 to 1.80 mm. Head length 0.46
to 0.52 mm; head width 0.43 to 0.46 mm. Thoracic length 0.41 to 0.59 mm.
Scape length 0.41 to 0.43 mm. Scape Index 82 to 89.
Structural characters: Head as in fig. 45, slightly longer than
broad, broadest through the eyes; posterior border of head slightly
notched medially; clypeal border curved and even; clypeus with a
median and 2 lateral striae; frontal area depressed and with a
median longitudinal striation; frontal carinae sub-parallel; antennal
scapes slender, more robust apically, reaching or surpassing the
occipital border by an amount less than first funicular joint; eyes
with less than 20 facets.
In profile, pronotum and mesonotum strongly convex and descend
ing sharply to the mesopropodeal impression, the latter with lon
gitudinal rugulae. Propodeal spines sharp and pointed obliquely


49
11. Pheidole greggi n. sp.
P. flavens sculptior M. R. Smith, 1951. Cat. Hym. Amer. No. of
Mex. U.S.D.A. No. 2, p. 802 (nec Forel).
P. flavens Gregg, R. E. 1958. N.Y. Entomol. Soc. 66:21 (nec
Roger).
Diagnosis:
A small yellowish species closely related to P. flavens Roger, P^.
flavens sculptior Forel, _P. floridana Emery, P. anastasii Emery, and
P^. moerens Wheeler. Antennal scrobe weakly depressed but punctate and
opaque. Frontal carinae not surpassing apices of scapes in repose.
Postpetiole without lateral projections. P. greggi has been previously
confounded with P. flavens sculptior and P. flavens.
Description:
Maj or:
Measurement: Body length 2.60 to 2.90 mm. Head length 0.82
to 0.89 mm; head width 0.79 to 0.82. Thoracic length 0.72 to 0.77 mm.
Scape length 0.41 to 0.46 mm. Scape Index 52 to 56.
Structural characters: Head shape as in fig. 21; head, exclud
ing mandibles, usually slightly longer than broad, broadest behind
the eyes. The occipital lobes strongly curved and convex laterally
but less curved or even angular mesally; median occipital sulcus
shallow. Frontal area small and depressed; antennal scrobes weakly
or not at all depressed. Clypeus with weak median carina, its
anterior margin with weak median emargination. Frontal carinae
short and divergent. Scapes slender at base, widening apically and
usually half or more the head length; mandibles stout, each curved


PLATE 2
Figure 7.
Figure 8.
Figure 9.
Figure 10
Figure 11
Scape of major of P_. fallax obscurithorax
Santschi (81X)
Head of major of P. fallax obscurithorax (40X)
Head of major of P. megacephala (Fabricius)
(40X)
Pedicel of major of P_. floridana Emery (128X)
Propodeum, pedicel and gaster of major of P.
metallescens Emery (74X)
Figure 12. Postpetiole of major of P. littoralis Cole (119X)


25
Due to the close resemblance of this species to I?. floridana Emery,
P_. anastasii probably has been misdetermined in several records from
Florida.
The difficulty of having access to Emery's types of _P. anastasii
makes it difficult to determine if the Florida species is conspecific with
the Costa Rican types. Wheeler, however, did have knowledge of the types
of _P. anastasii when he reported it from Florida.
P_. anastasii was usually found nesting under the bark at the base of
pines or along the roots and only rarely in the soil. The colonies are
monogynous and have more than 100 majors and over 500 workers. Several
chambers are constructed interconnected by a string of galleries under
the pine bark. The workers forage over 4 m from the colony and once food
is located majors are recruited to help transport it back to the colony.
It feeds on seeds, fruits, and scavenges on small dead arthropods and is
predaceous on small live arthropods.
Laboratory colonies are very easy to keep and usually accept members
of other colonies. Colonies that have lost their queens will also accept
any other queens of anastasii. Several times I collected two or three
nest founding females found together in one chamber. In the lab one
female always killed the others before the first workers were reared.


55
12. Pheidole lamia Wheeler
P. lamia Wheeler, 1901. Amer. Nat. 35:534. Wheeler, 1908. Amer.
Mus. Nat. Hist., Bui. 24:477-478. Wheeler, 1910. Ants, pp. 212, 248. -
Wheeler, 1926. Ants, pp. 212, 248. Smith, 1931. Ent. News 42:21-22.
Creighton, 1950. Bull. Mus. Comp. Zool. 104:182. Smith, 1951. Cat.
Hym. Amer. No. of Mex. U.S.D.A. Mon 2:803. Gregg, 1956. Ent. News
67:37-39. Gregg, 1958. N.Y. Ent. Soc. 66:23.
Type locality: Austin, Texas
Types : M.C.Z., A.M.N.H.
Range : North Florida to Texas
Diagnosis:
P. lamia is a small, yellowish, shining species with a subcylindrical
head in the major which is truncate anteriorly. The soldier of P. lamia
has the most striking head shape in the genus.
Discussion:
The head shape of P. lamia led Wheeler (1908) to suggest that it was
a phragmotic ant. He also believed it to be a very rare species. Burn
et al. (In Press), showed by laboratory and field observations that phragmosis
does not occur in this species.
At Tall Timbers Research Station where this species is common, I
had the opportunity to observe the biology of this interesting species.
Colonies are monogynous and sometimes have up to about 200 majors and
1,000 workers. The nests are in the ground, usually in grassy areas as
well as among taller vegetation. The nesting sites, at least at Tall
Timbers, can be easily spotted in the morning due to the reddish clay


61
15. Pheidole metallescens Emery
P. metallescens Emery, 1895. Zool. Jahrb., Abst. of System. 8:289,
294. Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:476-477. Smith,
1930. Fla. Ent. 14:3. Wheeler, 1932. N.Y. Ent. Soc. 40:7. Creighton,
S
19/0. Bull. Mus. Comp. Zool. 104:183. Smith, 1951. Cat. Hym. Am. No. of
Mex., U.S.D.A. Mon 2:803. Gregg, 1958. N.Y. Ent. Soc. 66:23. Wheeler
and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:12.
Type locality: St. George, Florida
Types : None in this country
Range : Gulf States
Diagnosis:
.P. metallescens (fig. 16) is a small species usually with reddish
brown head and thorax and black gaster, although completely dark to black
specimens have been found. It differs from _P. greggi n. sp. and P_. moerens
by the distinct lateral connules on the postpetiole. It also differs from
P_. floridana and IP. anastasii by the strong reticulation posteriorly on
the head and from P_. dentigula by having a reticulo-punctate propodeal
base. The workers display strong, metallic, iridescent bluish reflections
over most of the body.
Discussion:
P. metallescens is very common in Florida. The preferable habitat
seems to be under the shade of trees or any high vegetation. Usually it
is easy to find several small craters connected to a central vertical tunnel
to the main chamber, rarely over 40 cm deep in the ground. It is a
monogynous species. The majors usually forage along with the workers.
They feed on small grass seeds and also scavenge dead arthropods.


TABLE OF CONTENTS Continued
Page
3. Pheidole bureni n. sp 26
Diagnosis 26
Description 26
Major 26
Measurements 26
Structural characters 26
Sculpture 27
Pilosity 28
Color 28
Worker 28
Measurements 28
Structural characters 28
Sculpture 29
Pilosity 29
Color 30
Female 30
Measurements 30
Structural characters 30
Sculpture 30
Pilosity 31
Color 31
Male 31
Holotype 31
Paratypes 31
Discussion 32
4. Pheidole carrolli n. sp 33
Diagnosis 33
Description 33
Major 33
Measurements 33
Structural characters 33
Sculpture 34
Pilosity 35
Color 35
Worker 35
Measurements 35
Structural character 35
Sculpture 36
Pilosity 36
Color 36
Holotype 36
Paratypes 36
Discussion 37
v


62
16. Pheidole moerens Wheeler
P. moerens Wheeler, 1908. Bull. Amer. Mus. Nat. Hist. 24:136. -
Smith, 1936. Puerto Rico Univ., Jour. Agr. 20:842-843. Smith, 1967.
Cat. Hym. Amer. No. of Mex. U.S.D.A. Mon 2 (Suppl. 2):354. Wheeler
and Wheeler, 1972. Ga. Ent. Soc. Jour. 7:244. Kempf, 1972. Studia
Ent. 15:196. Wojcik, Banks, and Burn, 1975. Coop. Econ. Ins. Rpt.
U.S.D.A. 25449: 906.
P. moerens dominicensis Wheeler, 1913. Bull. Amer. Mus. Nat. Hist.
32:241. Kempf, 1972. Studia Ent. 15:197. New Synonymy.
Type locality: Antilha Culebra
Types : M.C.Z.
Range : Southern Alabama, Florida, West Indies
Diagnosis:
_P. moerens (fig. 22) is a small reddish yellow to almost black species
closely related to P. greggi n. sp., differing from it by the absence of
the heavily punctate scrobe and in having a carinated shining clypeus. It
differs from P_. dentigula, P_. anastasii, P_. floridana and P. metallescens
by the absence of distinct lateral connules on the postpetiole.
Discussion:
_P. moerens is a common species in north Florida. Its color was
observed in lab colonies to vary from reddish yellow to almost black due
possibly to variation in the diet. Colonies fed with honey and seeds
retained a lighter color than those fed with freshly killed house flies.
Wheeler's subspecies dominicensis appears to represent only a color
variant often found among the normal population. I have examined


6
The ants were placed in snap-cap vials with 70% alcohol for
preservation or kept alive in snap-cap vials with a piece of moisten
ed cotton to ensure adequate survival. The snap-cap vials were placed in
a styrofoam box to protect them against excessive changes in temperature
during transport back to the laboratory.
The colonies were kept in chambers made from petri dishes modified
to assure proper high moisture levels and ease of feeding and observation
without disturbing the colony.
Plastic petri dishes 9 cms in diameter were used. A 0.5 cm hole was
melted by a soldering iron toward theside of the bottom of one dish. A
piece of artificial cotton matting was pushed half way through the hole
and was coated inside with a mixture of plaster of paris (98%) and commercial
cement (2%). The matting below the hole was kept continuously wet with
distilled water and was retained in another petri dish bottom. It did not
need to be replenished with water frequently. The nest chamber was form
ed by applying the petri dish top and sealing with tape. A small feeding
chamber which remained dry was formed by affixing another petri dish
bottom to the petri dish top of the nest chamber with an interconnecting
exit hole. This construction allowed colonies to be maintained in the
laboratory for long periods with only infrequent attention. The dry feed
ing area separated from the nest chamber inhibited the growth of molds.
Large numbers of colonies were thus kept in a small laboratory space.
Colonies were fed with freshly killed house-flies, fresh peanuts,
and honey.
Whenever possible field observations were made concerning the ecology
and biology of each species.


15
11a) Head not completely punctate, the front of the head with
irregular smooth and shining areas. (fig. 42) 12
b) In full face view, head entirely covered with reticulate
punctation or with longitudinal striae. (fig. 43) 13
12a) Pleurae of pronotum heavily reticulate; head, thorax and
gaster usually with distinct violaceous or bluish metallic
reflections; eyes with less than 30 facets.
(fig. 41) metallescens Emery
b) Pleurae of pronotum smooth and shining; body without
violaceous or bluish reflections; eyes with more than
45 facets. (fig. 42) littoralis Cole
13a) Eyes large, with more than 40 facets. (fig. 43) carrolli n. sp.
b) Eyes small, with less than 25 facets. (fig. 50, 51) 14
14a) Postpetiole small, little wider than the petiole.
(fig. -4sr.) moerens Wheeler
or (fig. 4X)..".J'/. greggi n. sp.
b) Postpetiole nearly or more than twice as wide as the
petiole. (fig. 47) 15
15a) Body and mainly the gaster covered with numerous fine
long hairs; gaster enlarged, larger than the thorax;
the propodeal spines very thick and obliquely curved and
pointed laterad; first gastric tergite opalescent and
subopaque. (fig. 46, 47) quiescens n. sp.
b) Not with this combination of characters 16


24
2. Pheidole anastasii Emery
P_. anastasii Emery, 1896. Soc. Ent. Ital., Bol. 28:76. Forel,
1901. Naturhist. Mus. Hamburg, Mitt. 18:78. Wheeler, 1932. N.Y. Ent.
Soc., Journ. 40:6. Smith, 1933. Fla. Ent. 17:23. Creighton, 1950.
Bull. Mus. Comp. Zoll. 104:169. Gregg, 1958. N.Y. Ent. Soc. 66:16.
P,. floridana Van Pelt, 1956. Amer. Midland Nat., 56:376, 384.
Carroll, 1970. Unpublished M.S. thesis, University of Florida, p. 32-34. -
Whitcomb, Denmark, Bhatkar, and Greene, 197^ Fla. Ent., 55:133. (nec
Emery).
Type locality: Jimenes, Costa Rica
Types : None in the U.S.
Range : Florida, southern Alabama and Georgia
Diagnosis:
A small yellowish species differing from ?. floridana in having
opalescent surfaces dorsally on the first gastric tergite; differing
from P_. moerens and P_. greggi n. sp. by having the postpetiole twice as
broad as the petiole. Distinct lateral connules are present. It differs
from P_. dentigula in having smooth and shining occipital lobes.
Discussion:
Wheeler (1932) reported P. anastasii (fig. 17) from Florida. It seems
hard to believe that it is an introduced species as Wheeler suggested. I
have found this species throughout Florida from the keys to the Panhandle.
Its constant nesting at the base of pines and the occurrence of two parasites,
_P. quiescens n. sp., an inquiline ant, and a hymenopteran parasite in the
genus Orasema, seem to contradict Wheeler's suggestion that P. anastasii
is an exotic species.


82


37
The holotype major and 3 worker paratypes will be deposited at
the Museum of Comparative Zoology, Harvard University, Cambridge,
Massachussetts. Paratypes will also be deposited in the National
Museum of Natural History, Washington, D.C., Florida State Museum,
Department of Plant Industry, Gainesville, Florida and in the M. A.
Naves collections.
This species was also collected by me at Tall Timbers Research
Station, but the specimens, unfortunately, were inadvertantly lost.
Discussion:
This species has unusual behavior which is not common to other Pheidole
known to me. It was always found in shaded areas in sandy soils among tall
trees such as pines and oaks. It forages when the temperatures are not
high and avoids direct sunlight. It is easily found at dusk or in the
morning. The workers usually go out at random. Usually there are no
more than 15 workers foraging at one time, and the foraging territory
usually is within 5 meters from the nest entrance. The nest entrance is
small and usually very hard to spot. The worker is sluggish compared with
most other species. The majors are very rare. For a year two colonies
were periodically checked and yielded only one callow major in 1975 and
two majors in March, 1976. Dr. John F. Carroll also collected one major
in the same locality in August, 1973. Periodic excavation usually also
yielded a chamber approximately 4X2X2 cms located 30 cm deep and full of
various seeds, some apparently with a black fungal mycelium. Although the
excavation for one of these colonies at one time was 1.7 meters deep 1.5
meters in diameter, I was unable to locate the main chamber or any majors


56
soil cast up in very small honey-combed hummocks. These clumps consist
of fine soil particles and can have several openings connected to a vertical
tunnel of about 2 mm in diameter. From the vertical tunnel branch single
secondary galleries each directly connected to an oval chamber of about
0.5 to 1.5 cm in height and about 4 to 6 cm in length. Each colony has
several chambers. The first is found usually over 40 cm deep. I have dug
over 1.5 m deep following the main tunnel without reaching the end of it
in four colonies. In the summer of 1973, I found the first female after
digging seven colonies. In July, 1974, I found about thirty males in
tunnels close to the soil surface. In July, 1975, I found 18 females
inside a large chamber of about 50 cm deep. One mating pair that fell on
the ground was collected in the late afternoon on July 15, 1974. The soil
where this species nests usually has the first 30 cm predominantly sandy.
Deeper there is a reddish clay where this species has its chambers.
This species usually forages in the morning or when there is no
sunlight. Majors sometimes are found among debris with the workers.
The pale yellow color of this species also could indicate that this species
is nocturnal. The workers were observed carrying small seeds, small insects
and mites to the nest. Some majors in the chambers might be repletes since
they have quite extended gasters full of liquid. Burn et al. (In Press)
show that this species has stylized defenses against Solenopsis
(Diplorhoptrum) spp.


44
8. Pheidole diversipilosa Wheeler
P. crassicornis var. diversipilosa Wheeler, 1908. Amer. Mus. Nat.
Hist., Bui. 24:467.
P. crassicornis Creighton, 1950. Bull. Mus. Comp. Zool., 104:175-
176.
Type locality: Fort Davis, Texas.
Types : M.C.Z.
Range : Northwestern Florida to western Texas
Diagnosis:
]P. diversipilosa (fig. 3) is a reddish yellow species that differs
from P. crassicornis and P. bureni n. sp. by the numerous erect hairs
on the gaster. In P. crassicornis tetra the gastric hairs are nearly
all of the same short length rather than obviously of mixed lengths
as in diversipilosa.
Discussion:
P. diversipilosa was collected at Tall Timbers Research Station
several times where F\ crassicornis is also present. Morphological
distinction is constant between the two species. There is no real over
lap in the pilosity characters. Therefore, I have treated F\ diversipilosa
as a full species on the basis that it can be sympatric with P. crassicornis
but still retains constant morphological distinction. Creighton (1950)
treated diversipilosa as an intergrade between crassicornis and tetra.
The finding of both crassicornis and diversipilosa in the same locality
in Florida invalidates Creighton's viewpoint.
Specimens of _P. crassicornis collected in Gainesville were within
the variation shown by the Tall Timbers specimens. I was unable to find


98


64
17. Pheidole morrisi Forel
P. morrisii Forel,
1886.
Soc.
Ent.
Belg.
, Bui.
30:46. -
Mayr,
1887.
Verh.
Zool.
- Bot. Ges.
Wien 37:568
. -
Forel,
1901.
Soc. Ent
. Belg.
* >
Ann.
45:350.
- Wheeler,
1904.
Amer.
Mus
. Nat.
Hist.
, Bui 20:
302. -
Wheeler,
1905. Amer. Mus. Nat. Hist., Bui. 21:380. Wheeler, 1908. Amer. Mus.
Nat. Hist., Bui. 24:461. Smith, 1918. Ent. News 29:21. Davis and Bequaert,
1922. Brooklyn Ent. Soc. Bui. 17:8-9. Smith, 1924. Ent. News 35:53. -
Smith, 1930. Fla. Ent. 14:3. Wheeler, 1932. N.Y. Ent. Soc. 40:7. -
Cole, 1940. Amer. Midland Nat. 24:44-45. Creighton, 1950. Bull. Mus.
Comp. Zool. 104:183-184. Smith, 1951. Cat. Hym. Amer. No. of Mex.,
U.S.D.A. Mon 2:803. Gregg, 1958. N.Y. Ent. Soc. 66:24. Wheeler and
Wheeler, 1960. Ent. Soc. Amer., Ann. 53:12.
P. morrisi var. vancea Forel, 1901. Soc. Ent. Belg., Ann. 45:351. -
Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20:302. Wheeler, 1908. Amer.
Mus. Nat. Hist., Bui. 24:461. Smith, 1924. Ent. News 35:53. Smith,
1951. Cat. Hym. Amer. No. of Mex., U.S.D.A. Mon 2:803-804.
Type locality: Vineland, New Jersey
Types : None in this country
Range : Southern New Jersey, south to Florida and eastern
Gulf States.
Diagnosis:
jP morrisi (fig. 27) is a pale to darker yellowish species differing
from P. dentata and JP. vinelandica by the absence of distinct propodeal
teeth or spines. It is larger than P. vinelandica and has a transverse
mesonotal impression.


A MONOGRAPH OF THE GENUS PHEIDOLE IN FLORIDA
(HYMENOPTERA:FORMICIDAE)
By
Marcio A. Naves
A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF THE
UNIVERSITY
OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE
DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1976


78


27
In profile, pronotum and anterior mesonotum convex, mesonotum
with a wide impression in the middle; posteriorly the mesonotum
abruptly descending to the mesopropodeal impression; propodeal base
longer than declivity; propodeal spines short and pointed dorsally;
petiole stout, the anterior face and posterior face sloping, the
node quite large and its crest in profile varying from weakly angular
to strongly convex; the ventral border slightly convex anteriorly;
postpetiole strongly convex dorsally and flat ventrally.
Seen from above, pronotum with pronounced but rounded shoulders;
petiole with parallel sides, the crest of node with a shallow concave
impression in the middle; postpetiole almost two times wider than
long; slightly angular to blunt laterally; gaster truncate at the
base.
Sculpture: Fine longitudinal striae present on the frons, inter
stitial areas finely and weakly reticulo-punctate; vertex and occipital
lobes smooth and shining; genae with longitudinal rugae, the inter
stitial areas weakly punctate; frontal*carinae short and slightly
diverging posteriorly; frontal area smooth and shining; clypeus smooth
and shining, usually with a weak median carinae; frontal furrow with
transverse short rugulae posteriorly; ventral area of head smooth
and shining, its anterior margin with two well developed sharp teeth.
Pronotal pleurae weakly reticulate; mesonotum, propodeum and
petiole finely reticulo-punctate, the floor of each punctation strongly
shining.
Seen from above, pronotum and adjacent area of mesonotum with
weak transverse striae and weakly reticulated; mesonotal dorsum
smooth and strongly shining; mesonotal impression not easily seen;


A MONOGRAPH OF THE GENUS PHEIDOLE IN FLORIDA
(HYMENOPTERA:FORMICIDAE)
By
Marcio A. Naves
A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF THE
UNIVERSITY
OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE
DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1976

To my parents, wife and
and in memory of
son
Father Walter W. Kempf, O.F.M.

ACKNOWLEDGEMENTS
i
I wish to express my gratitude to Dr. William F. Burn, Advisory
Committee Chairman, who patiently guided me and inspired me during every
phase of this research. I also wish to acknowledge my sincere indebteness
to Dr. George E. Allen, Dr. Francis W. Zettler and Dr. Clifford S. Lofgren,
members of my Advisory Committee, for their constructive criticisms and
guidance.
I sincerely thank Mrs. T. Carlysle for providing me with scanning
electron micrographs.
I also wish to thank Dr. John F. Carroll, Dr. J. C. E. Nickerson, and
Dr. D. P. Wojcik, for continuing assistance, and to Dr. Edward 0. Wilson
and Dr. David R. Smith for their permission to study the collections at
the Museum of Comparative Zoology and U.S. National Museum, respectively.
This research was also aided in part by Tall Timbers Research Station,
Leon Co., Florida, through the kindness of Dr. E. V. Komarek, Director of
the Station, and the sponsorship of Dr. Willard H. Whitcomb. The support
of Empresa Brasileira de Pesquisa Agropecuaria is also acknowledged with
gratitude.
To my wife, Lucilia, my utmost thanks and appreciation for her patience
iii
and support.

TABLE OF CONTENTS
Page
ACKNOWLEDGMENTS iii
LIST OF FIGURES viii
ABSTRACT xii
INTRODUCTION 1
REVIEW OF LITERATURE 3
MATERIALS AND METHODS 5
KEY TO SPECIES OF PHEIDOLE OF FLORIDA (MAJORS ONLY) 8
KEY TO PHEIDOLE SPECIES OF FLORIDA (WORKERS ONLY) 13
1. Pheidole adrianoi n. sp 17
Diagnosis 17
Description 17
Major 17
Measurements 17
Structural characters 17
Sculpture 18
Pilosity 19
Color 19
Worker 19
Measurements 19
Structural characters 19
Sculpture 20
Pilosity 20
Color 20
Female 20
Measurements 20
Structural characters 20
Sculpture 21
Pilosity 21
Color 22
Male 22
Holotype 22
Paratypes 22
Discussion 22
2. Pheidole anastasii Emery 24
Diagnosis 24
Discussion 24
lv

TABLE OF CONTENTS Continued
Page
3. Pheidole bureni n. sp 26
Diagnosis 26
Description 26
Major 26
Measurements 26
Structural characters 26
Sculpture 27
Pilosity 28
Color 28
Worker 28
Measurements 28
Structural characters 28
Sculpture 29
Pilosity 29
Color 30
Female 30
Measurements 30
Structural characters 30
Sculpture 30
Pilosity 31
Color 31
Male 31
Holotype 31
Paratypes 31
Discussion 32
4. Pheidole carrolli n. sp 33
Diagnosis 33
Description 33
Major 33
Measurements 33
Structural characters 33
Sculpture 34
Pilosity 35
Color 35
Worker 35
Measurements 35
Structural character 35
Sculpture 36
Pilosity 36
Color 36
Holotype 36
Paratypes 36
Discussion 37
v

TABLE OF CONTENTS Continued
Page
5. < Pheidole crassicornis Emery 39
Diagnosis 39
Discussion 39
6. Pheidole dentata Mayr 40
Diagnosis 41
Discussion 41
7. Pheidole dentigula M. R. Smith 43
Diagnosis 43
Discussion 43
8. Pheidole diversipilosa Wheeler 44
Diagnosis 44
Discussion 44
9. Pheidole fallax obscurithorax Santschi 46
Diagnosis 46
Discussion 46
10. Pheidole floridana Emery 48
Diagnosis 48
Discussion 48
11. Pheidole greggi n. sp 49
Diagnosis 49
Description 49
Major 49
Measurements 49
Structural characters 49
Sculpture 50
Pilosity 51
Color 51
Worker 51
Measurements 51
Structural characters 51
Sculpture 52
Pilosity 52
Color 52
Female and Male 52
Holotype 52
Paratypes 52
Discussion 52
12. Pheidole lamia Wheeler '55
Diagnosis 55
Discussion 55
13. Pheidole littoralis Cole 57
Diagnosis 57
Discussion 57
14. Pheidole megacephala (Fabricius) 58
Diagnosis.. 58
Discussion 59
vi

TABLE OF CONTENTS Continued
Page
15. Pheidole metallescens Emery 61
Diagnosis 61
Discussion 61
16. Pheidole moerens Wheeler 62
Diagnosis 62
Discussion 62
17. Pheidole morrisi Forel 64
Diagnosis 64
Discussion 65
18. Pheidole quiescens n. sp 66
Diagnosis 66
Description 66
Major 66
Worker 66
Measurements 66
Structural characters 66
Sculpture 67
Pilosity 67
Color 68
Female and male 68
Holotype 68
Paratypes 68
Discussion 68
19. Pheidole tysoni Forel 71
Diagnosis 71
Discussion 71
20. Pheidole vinelandica Forel 72
Diagnosis 73
Discussion 73
SPECIES NOT PROPERLY INCLUDED IN THE FLORIDA PHEIDOLE FAUNA .... 75
1. Pheidole pilifera (Roger) 75
Diagnosis 75
Discussion 75
LITERATURE CITED 103
BIOGRAPHICAL SKETCH 105
vii

is
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
LIST OF FIGURES
Page
Head of major of P. lamia Wheeler (33X) 78
Scape of major of P. bureni n. sp. a member of the
crassicornis complex (81X) 78
Head of major of P_. diversipilosa Wheeler (45X) 78
Gaster of major of P_. diversipilosa (47X) 78
Head of major of P_. crassicornis Emery (43X) 78
Gaster of major of crassicornis (58X) 78
Scape of major of P. fallax obscurithorax Santschi
(SIX) 80
Head of major of P. fallax obscurithorax (40X) 80
Head of major of _P. megacephala (Fabricius) (40X) 80
Pedicel of major of ]P. floridana Emery (128X) 80
Propodeum, pedicel and gaster of major of P.
metallescens Emery (74X) 80
Postpetiole of major of ]?. littoralis Cole (119X) 80
Head of major of P_. dentigula M. R. Smith (64X) 82
Head of major of P_. carrolli n. sp. (29X) 82
Pedicel of major of P_. metallescens Emery (163X) 82
Head of major of P_. metallescens (62X) 82
Head of major of P. anastasii Emery (68X) 82
Postpetiole and gaster of P. anastasii (62X) 82
Head of major of P. floridana Emery (72X) 34
Head of major of P. littoralis Cole (37X) 84
viii

la
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
LIST OF FIGURES Continued
Page
84
Head of major of P. greggi n. sp. (68X)
Head of major of EL moerens Wheeler (58X) 84
Clypeus and frontal area of major of P_. vinelandica
Forel (186X) 84
Head of major of P_. bicarinata Mayr (53X) 84
Head of major of P_. adrianoi n. sp. (72X) 86
Thorax, pedicel and gaster of major of _P. adrianoi
n. sp. (50X); white legs are artifacts due to electron
overcharge 86
Head of major of P_. morrisi Forel (37X) 86
Thorax of major of P_. morrisi (56X) 86
Head of major of P. dentata Mayr (48X) 86
Teeth on the anterior ventral head margin of PL
anastasii (Emery (123X) 86
Head of worker of P.. lamia (97X) 88
Dorsal view of worker of PL vinelandica (62X) 88
Dorsal view of worker of P_. adrianoi n. sp. (95X) 88
Worker of P. megacephala (36X) 88
Thorax and pedicel of the worker of P. fallax
obscurithorax (41X) 88
Head of worker of PL fallax obscurithorax (86X) 88
Thorax of the worker of PL morrisi (72X) 90
Head thorax and pedicel of the worker of P. dentata
(72X) r; 90
Thorax of the worker of PL dentata (79X) 90
IX

is
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
LIST OF FIGURES Continued
Page
Head of the worker of P. crassicornis (54X) 90
Thorax of the worker of P_. metallescens (196X) 90
Head of the worker of P. littoralis (113X) 90
Worker of P_. carrolii n. sp. (40X) 92
Head of the worker of P. moerens (102X) 92
Head of worker of P_. greggi n. sp. (102X) 92
Worker of P. quiescens n. sp. (45X) 92
Dorsal view of worker of P. quiescens n. sp. (103X).... 92
Propodeum, pedicel, and gaster of worker of .P.
dentigula (102X) 9?
Dorsal view of thorax of P. dentigula worker (115X) 94
Head of worker of J?. anastasii (109X) 94
Head of worker of P_. floridana (115X) 94
Pedicel of worker of P_. floridana (193X) 94
Dorsal thoracic hairs of worker of P. adrianoi n. sp.
(1,401X and 5,174X, respectively) 94
Dorsal thoracic hairs of worker of P_. adrianoi n. sp.
(1,401X and 5,174X, respectively) 94
Head of major of _P. flavens (95X) 95
Head of major of P_. flavens (87X) 95
Occipital lobes of P_. greggi n. sp. (250X) -95
Postpetiole of the major of P. flavens sculptior (158X) 96
Head and thorax of the minor of P. flavens (83X) 96
Head of major of P. bureni n. sp. (39X) 93
x

LIST OF FIGURES Continued
Figure
61 Gaster and pedicel of P_. bureni n. sp. (40X) .
62 Head and pronotum of worker of P_. bureni n. sp.
(52X)
63 Dorsum of propodeum of maior of P. vinelandica
(272X) 7
64 Thoracic dorsum of P. bicarinata (71X)
65 Postpetiole of P_. bicarinata (186X)
66 Head of P. vinelandica (66X)
67 Map of Florida and the United States
Page
98
100
100
100
100
102
xi

Abstract of Dissertation Presented to the Graduate Council
of the University of Florida in Partial Fulfillment of the Requirements
for the Degree of Doctor of Philosophy
A MONOGRAPH OF THE GENUS PHSIPPLE IN FLORIDA
(HYMENOPTERA:FORMICIDAE)
By
Marcio A. Naves
December, 1976
Chairman: William F. Burn
Major Department: Entomology and Nematology
Until recently only twelve species of Pheidole were reported from
Florida, P. anastasia Emery, P_. bicarinata vinelandica For el, P. dentigula
M. R. Smith, ?. f lavens Roger, P. megacephala (Fabricius), P_. moerens
Wheeler, morrisl Forel, P_. pilifera (Roger), P. dentata Mayr, P.
floridana Emery, P. metallescens Emery and P_. sitarches littoralis Cole.
The last four have Florida as the type locality.
£. pilif era (Roger) is believed to have beer, errouneously reported
from Florida and has been excluded from the Florida fauna. P. flavens
(Gregg, 1958, nec Roger) is considered a misidentification and to be an
undescribed species, P. greggi n. sp.
In addition to P. greggi n. sp. three additional new species of
Pheidole are described from Florida: P_. adrianoi n. sp., P. carrolli n.
sp. and £. qu.iesc.ens n. sp. An exotic species, P. fallax obscurithorax
(= £. fallax ajencola var. obscurithorax Santschi) probably introduced
from South America, is also added to the North American ant fauna.
Pheidole crassicornis Emery and ?. diversipllosa Wheeler are record
ed for the first time in Florida. The latter species is resurrected
from sycornymy and is elevated to full species.

Two additional taxa are elevated to full species: JP. vinelandica
(= JP. bicarinata vinelandica Forel) and P_. littoralis (= JP. sitarches
littoralis Cole) The _P. crassicornis complex is recognized with the
description of 1?. bureni n. sp. collected in Alice, Texas even though
this species does not occur in Florida.
The four new species, P_. adrianoi n. sp., JP. bureni n. sp. JP.
carrolii n. sp., and JP. quiescens n. sp., plus the introduced species
P^. fallax obscurlthorax Santschi and the recognized species 1?. diversi-
££ sa Wheeler increase the total Pheidole fauna of North America north
of Mexico to seventy-seven. The Florida Pheidole fauna is increased
to nineteen species and Pheidole is now the largest ant genus in the
state.
Keys for the identification of both majors and workers are pro
vided. Scanning electron micrographs are also provided as identifica
tion aides. The biology of each species is discussed.
xiii

INTRODUCTION
The genus Pheidole is one of the more important ant genera in
the world. Approximately 1000 taxa are known. Over 400 taxa are known
from the Neotropical region (Kempf, 1972), and about 77 taxa are now known
from North America north of Mexico. Pheidole spp. are abundant in many
areas, and live in varying habitats ranging from the humid tropics to
deserts. They are able to survive in some areas by their habits of
collecting and storing seeds as food resources. They are also scavengers
of dead insects and ether animals, and can be predaceous. Some species
tend aphids and other homopterans, but this food source, so important
to many ants genera, is probably of relatively minor importance to most
species of Pheidole.
Pheidole apparently is a genus of rather recent evolutionary develop
ment. No fossil Pheidole are known previous to the Miocene (Brown, 1973).
There are no Pheidole in the Baltic amber (lower Oligocene) although these
ambers contain large numbers of specimens of other ant genera which have
persisted almost unchanged since that epoch (Formica, Myrmica, Iridomyrmex
Caranonotus. and others). It has been said that Pheidole has undergone
a world wide dispersal since this time and has undergone a large pro
liferation of taxa as well as adaptation to many ecological nitches (Brown
1973) .
A study of the Pheldole of any area is one of the prerequisites to
understanding the formicid interactions of that area and this in turn
could be important in the development of pest management strategies, since

2
ants are probably the most numerous and most important animals within
their size range.
One species, Pheidole megacephala, has become a tropicopoiitan tramp
species, and is a serious pest in various areas (Wheeler. 1910, Flucker
and Beardsley, 1970 and Lieburg, 1975). Some species sting severely
and are comparable with the fire ants in aggressiveness (Burn, personal
communication). Most of the species, however, appear innocuous and are
usually unnoticed by man. Their importance in the ecosystem appears
largely unappreciated.
Due to its size and distribution, Pheidole is a very difficult genus
and has been a challenge to myrmecologists taxonomically as well as
biologically.
Until recently only eleven species were known from Florida, which
suggested that despite its subtropical warmth Florida was deficient in
Pheidole when compared to Texas (36 species) and Arizona (26 species).
The Florida Pheidole fauna has long been known to be more extensive than
that of northern states such as Iowa and New Jersey (3 species each,
Burn, 1944 and Creighton, 1950).
This study was undertaken to increase our knowledge of Pheidole
of Florida. Scanning electron micrographs are provided along with keys
for the identification of both majors and workers.

REVIEW OF LITERATURE
Four out of twelve species reported from Florida have that state as
the type locality: dentata Mayr (1886), P. floridana Emery (1895),
P_. metallescens Emery (1895), and _P. sitarches litorallis, Cole (1952).
Smith (1930) reported P. morrisi; Wheeler (1932) reported P_. anastasil;
Smith (1^33) reported _P. megacephala; Smith (1944) reported J?. dervtigula;
Smith (1951) reported _P. flavens sculptlor (misidentification); Van Pelt
(1956) reported P. pilifera; Smith (1958) reported ?. bicarinata vinelandica,
and Wojcik et al. (1975) reported P_. moerens. Additional information is
discussed under each species by the respective authors.
Creighton (1950) was the first, to try to give a extensive key to the
North America Pheidole; 63 species were listed and he used morphological
characters pertinent to both majors and workers. Creighton's statements
about the North American Pheidole are as follows:
Most of our species of Pheidole posses a dimorphic worker caste
with major and minor workers not connected by intermediates.
In a few species, however, the worker caste is polymorphic (kingi,
instabilis and torpescens, vasliti arizonica, etc.). Most of
the species garner seeds and it is believed that the large-headed
major workers function as seed-huskers. The enlarged head of the
major is mainly filled with mandibular muscles. This enables the
jaws to exert much pressure, which should be useful in cracking
off the husks of seeds. It may be added that sometimes the head
of the major is so large in proportion to its body that if the
insect is turned over on the back of its head it cannot regain
a normal posture without help from other workers. Despite
their preference for a graminivourous diet many species of
Pheidole will accept other food as well. They seem less attract
ed to honey-dew than do many ants but will often feed voraciously
on animal tissue when the opportunity offers. The majority of
our species form small colonies. In many cases there are only
3

4
about two or three hundred individuals in a fully developed
nest. Even in the case of the species which produce comparatively
large nests (morrisi, hyatti, desertorum, etc.) a colony of more
than two or three thousand individuals would be exceptionally
large. By far the majority of our species nest in soil. The
nest may be built under a stone or in open soil without a cover
ing object. In the latter case there is often a mound or craters
of excavated soil surrounding the nest entrance. The eastern
species dentata will nest in rotten logs as well as soil but
such flexibility in nesting habit is exceptional.
(p. 162)
Gregg (1958), published a new revision in which he included the
description of ten new taxa. He also placed ten taxa in synomymy and the
status of eight taxa was revised. Later, Gregg (1969) described another
new species, P_. clementensis.
Smith (1967) reported P_. moerens from Alabama as yet another exotic
species. This was the 71st Pheidole taxon reported for North America north
of Mexico. Wojcik et al. (1975) reported that this species was also
present in Florida.

MATERIALS AND METHODS
Four species of Pheidole (P. dantigula, _P. fallax obscurithorax,
P. bureni, n. sp. and P. greggi n. sp.) included in this research were
received from other collectors. All other specimens were collected by
the author during severa] trips in the state (fig. 67) or while at Tall
Timbers Research Station in Leon Co., Florida, during the summers of
1974 and 1975.
Colonies of Pheidole spp. were located in several ways:
a. Examingiug cavities under the bark of trees or examining grass
behind the leaf bases of palms.
b. Searching for nest craters, mounds, or any other modification
on the soil surface likely to have been produced by ants.
c. Searching for foraging majors or workers and following them to
the nest location.
d. Baiting with honey or dead arthropods and following trails of
workers and majors to the nest location.
3. Overturning stones, boards, or pieces of wood.
The two most successful methods were examining cavities under the
bark of pine trees and looking for craters on the soil surface.
Colonies were collected by aspirating the ants once the chambers were
located. Nests in the soil were excavated by removal of large blocks of
soil, breaking it carefully and aspirating the ants found. The depth
explored was variable, ranging from 30 to 50 era for P. metallescens
to over 1.5 m for P. lamia.
5

6
The ants were placed in snap-cap vials with 70% alcohol for
preservation or kept alive in snap-cap vials with a piece of moisten
ed cotton to ensure adequate survival. The snap-cap vials were placed in
a styrofoam box to protect them against excessive changes in temperature
during transport back to the laboratory.
The colonies were kept in chambers made from petri dishes modified
to assure proper high moisture levels and ease of feeding and observation
without disturbing the colony.
Plastic petri dishes 9 cms in diameter were used. A 0.5 cm hole was
melted by a soldering iron toward theside of the bottom of one dish. A
piece of artificial cotton matting was pushed half way through the hole
and was coated inside with a mixture of plaster of paris (98%) and commercial
cement (2%). The matting below the hole was kept continuously wet with
distilled water and was retained in another petri dish bottom. It did not
need to be replenished with water frequently. The nest chamber was form
ed by applying the petri dish top and sealing with tape. A small feeding
chamber which remained dry was formed by affixing another petri dish
bottom to the petri dish top of the nest chamber with an interconnecting
exit hole. This construction allowed colonies to be maintained in the
laboratory for long periods with only infrequent attention. The dry feed
ing area separated from the nest chamber inhibited the growth of molds.
Large numbers of colonies were thus kept in a small laboratory space.
Colonies were fed with freshly killed house-flies, fresh peanuts,
and honey.
Whenever possible field observations were made concerning the ecology
and biology of each species.

7
A trip was taken to the Museum of Comparative Zoology, Harvard
University, Cambridge, Massachussetcs, and to the National Museum of
Natural History in Washington, D.C., in order to study types of Nearctic
and Neotropical Pheidole.
All specimens were point mounted and studied by using a stereo micro
scope with magnification up to 160X. The pictures were taken using the
scanning electron microscope of the Insects Attractants Laboratory, Agri
cultural Research Service, U.S.D.A., Gainesville, Florida.
Several measurements were taken for the newly described species
including:
a. Body length: the sum of the head length, thoracic length, pedicel
and gastric lengths.
b. Head length: lengths of the. head in full face view (mandibles
excluded).
c. Head width: greatest width of the head in full face view.

d. Thoracic length: greatest length of thorax in lateral view.
e. Scape length: middle of antennal socket to the scape tip.
f. Scape index: found by computing the formula: Scape length X 100
head length
The following abreviations were used throughout this work:
M.C.Z. Museum of Comparative Zoology, Harvard University, Cambridge,
Massachussetts.
U.S. Nat. Mus. United States National Museum, Washington, D.C,
A.M.N.H. American Museum of Natural History, New York, N.Y.

KEY TO SPECIES OF PHEIDOLE OF FLORIDA
(MAJORS ONLY)
la) Head cylindrical in cross section and obliquely truncate
anteriorly, the truncation involving the clypeus, frontal
area, anterior portions of genae, and mandibles.
(fig. 1) lamia Wheeler
b) Head not cylindrical in cross section and not truncate
anteriorly 2
2a) Antennal scape strongly bent, flattened and smooth at the
base, the base nearly as or as broad as the distal portions
of the scape (fig. 2) 3
b) Not as above, scape not flattened at base and usually
distinctly narrower than distal portion; if thickened
and strongly bent at base, obviously not flattened 4
3a) First gastric tergite covered with very numerous long and
short erect hairs; sometimes sparse, coarse, appressed,
pubescence is present, (fig. 3, 4) diversipilosa Wheeler
b) First gastric tergite covered with appressed pubescence
and sometimes with sparse long hairs.
(fig. 5, 6) crassicornis Emery
4a) Large species, total length over 6 mm; head heavily
sculptured; scapes very thick and bent at base. The
scapes seen in full face view are evenly curved measally,
but more angular laterally (fig. 7, 8)..fallax obscurithorax Santschi
8

9
b) Species usually measuring 5 mm or less; if larger the head
never heavily sculptured; the scapes never thick or
strongly bent at base 5
5a) Head cordate, gradually but distinctly narrowed toward
the mandibular insertions; broadest at the occipital
lobes; mesonotum not transversely impressed.
(fig. 9) megacephala (Fabricius)
b) Head not cordate and never strongly narrowed towards
the mandibular insertions; mesonotum may or may not be
transversely impressed 6
6a) Postpetiole, seen from above, bearing distinct lateral
connules; head covered with reticulate punctation or
longitudinally striate. (fig. 10, 11) 7
b) Postpetiole seen from above, without distinct lateral
connules; if indistinct lateral projections present
the posterior half of the head smooth and shining.
(fig. 12) 11
7a) Occipital lobes with reticulate sculpture, with no trace
of transverse or longitudinal striation, the surface
opaque or feebly shining; post-petiolar width two times
length. (fig- 13) dentigula M. R. Smith
b) Occipital lobes smooth and shining or with sparse
punctures or striae; postpetiole width greater than length,
but never two times length. (fig. 10, 11) 8
8a) Sides of head sub-parallel, narrowing slightly anteriorly;
head length 1.82 to 2.0 mm; scape index 41 to 42; pronotum

10
and mesonotum strongly sculptured with transverse and
sometimes with longitudinal rugae; dorsal crest of petiole
notched. (fig. 14) carrolli n. sp.
b) Without all the above characters 9
9a) Erect gastric hairs sparse, tapered and almost all of the
same size; posterior part of the head covered with sculpture;
anterior ventral margin of the head without teeth; body
sometimes with distinct violaceous or bluish metallic
reflections. (fig. 15, 16) metallescens Emery
b) Gaster ocvered with long and short hairs; posterior part
of the head smooth and shining (fig. 17); anterior ventral
margin of the head with two or three teeth, the mesal
tooth usually short and blunt. (fig. 30) 10
10a) Dorsal surface of first gastric tergite finely and densely
granulse, opalescent and opaque; dorsal areas of pronotum
punctate and sometimes with several short and fine transverse
striae; dorsal area of post-petiole punctate.
(fig. 17, 18) anastasii Emery
b) Dorsal surface of first gastric tergite smooth and shining;
dorsum of pronotum partially weakly punctate but also
interdispersed with smooth and shining areas; dorsal area
of post-petiole weakly punctate and shining.
(fig. 10, 19) floridana Emery
11a) Head covered with longitudinal striae; the dorsal area of
the pronotum and usually the mesothorax reticulo-punctate
and with transverse striae or sculpture, (fig. 20)
12

11
b) Posterior areas of the head smooth and shining; the pronotum,
seen from above, smooth and very shining 14
12a) Head usually measuring over 1.4 mm in length and 1.1 mm
in width; sides of the head sub-parallel; scape length
shorter than half the head length (Scape Index around 40);
occipital lobes with distinct circular piligerous punctures.
(fig. 12, 20) littoralis Cole
b) Head measuring less than .9 mm in length and width; occipital
lobes smooth and shining 13
13a) Frontal area concave and bearing 1 or 2 longitudinal
striae. Scrobes strongly punctate and opaque; occipital
lobes smooth and weakly shining; on occiput interstitial
spaces between striae reticulo-punctate. (fig. 21).... greggi n. sp.
b) Frontal area concave, without longitudinal striae; posterior
half of the scrobe weakly punctate and shining; occipital
lobes and most of the head shining; on front, interstitial
spaces between striae smooth and shining. (fig. 22). moerens Wheeler
14a) Anterior clypeal margin bluntly bidentate, with deep
median notch; anterior ventral margin of head without
teeth. (fig. 23) vinelandica Forel
b) Anterior clypeal margin not bidentate and without notch;
anterior ventral margin of the head usually with two or
three teeth; if teeth are absent, shining black species 15
15a) Mesonotum without transverse impression, forming in profile
a continuous smooth curve with the pronotum. (fig. 26) 16

12
b) Mesonotum, with a well developed transverse impression 17
16a) A yellow species; frontal area and middle of front smooth
and shining without striae; anterior ventral margin with
3 teeth; thorax smooth and shining tysoni Forel
b) A black species; anterior half of head with longitudinal
striae; dorsum of pronoturn smooth and shining; pleurae
. of pronotum, mesonotum, and propodeum, heavily sculptured.
(fig. 25, 26) adrianoi n. sp.
17a) Propodeum angular at the junction of the basal and declivous
faces, but the angles not produced into distinct teeth or
spines. (fig. 27, 28) morrisi Forel
b) Propodeum armed with distinct teeth or spines.
(fig. 29)
dentata Mayr

KEY TO PHEIDOLE SPECIES OF FLORIDA
(WORKERS ONLY)
la) Entire head smooth and shining. (fig. 31, 36) 2
b) Head reticulo-punctate or partially reticulate with some
smooth areas. (fig. 40, 41, 44) 9
2a) Scapes failing to reach the posterior border of the head,
or surpassing it by an amount no greater than the first
funicular joint. (fig. 31) 3
b) Scapes surpassing the posterior border of the head by an
amount greater than the first funicular joint. (fig. 34) 6
3a) Thorax smooth, nearly free of sculpture and strongly
shining; yellow 4
b) Pronotum smooth and shining; mesonotum and propodeum
punctate or sculptured; color various. (fig. 32, 33) 5
4a) Propodeum armed with angular teeth which are broad at the
base and do not resemble spines; mesonotum and propodeum
in profile separated by a deep, wide impression.
('MgvBS1) lamia Wheeler
b) Propodeum armed with spines; mesonotum and propodeum in
profile separated by a normal impression tysoni Forel
5a) Yellowish to brownish species, gaster covered with numerous
long fine hairs, (fig. 32) vinelandica Forel
b) Ebony black species, gaster with plumose or subplumose
hairs which are spaced at distances more or less equal to
the hair's length. (fig. 33) adrianoi n. sp.
13

u
6a) Mesonotum without a distinct tranverse impression; in
profile postpetiole longer than petiole and dorsal and
ventral outlines both smoothly convex.
(fig. 34) megacephala (Fabricius)
b) The mesonotum with a distinct transverse impression.
(fig. 35, 37 39) 7
7a) Posteriorly on head a distinct collar visible in full
face view (fig. 36); the mesonotum and base of propodeum
heavily punctuate. (fig. 35) fallax obscurithorax Santschi
b) Posterior collar of head not visible in full face view;
the mesonotal dorsum and propodeal base weakly sculptured
and shining 8
8a) Usually yellowish or pale species, the propodeal spines
absent. (fig. 37) morrisi Forel
b) Dark brown to ferruginous species; propodeal spines well
formed. (fig. 38, 39) dentata Mayr
9a) Scapes surpassing the posterior margin of the head by a
distance greater than the first funicular joint. (fig. 40) 10
b) Scapes either not surpassing posterior margin of the head,
or surpassing less than the length of the first funicular
joint. (fig. 43) 11
10a) First gastric tergite with appressed pubescence and only a
few scattered long hairs crassicornis Emery
b) First gastric tergite covered with numerous long and short
hairs; pubescence usually erect or semi-erect, rarely
appressed
diversipilosa Wheeler

15
11a) Head not completely punctate, the front of the head with
irregular smooth and shining areas. (fig. 42) 12
b) In full face view, head entirely covered with reticulate
punctation or with longitudinal striae. (fig. 43) 13
12a) Pleurae of pronotum heavily reticulate; head, thorax and
gaster usually with distinct violaceous or bluish metallic
reflections; eyes with less than 30 facets.
(fig. 41) metallescens Emery
b) Pleurae of pronotum smooth and shining; body without
violaceous or bluish reflections; eyes with more than
45 facets. (fig. 42) littoralis Cole
13a) Eyes large, with more than 40 facets. (fig. 43) carrolli n. sp.
b) Eyes small, with less than 25 facets. (fig. 50, 51) 14
14a) Postpetiole small, little wider than the petiole.
(fig. -4sr.) moerens Wheeler
or (fig. 4X)..".J'/. greggi n. sp.
b) Postpetiole nearly or more than twice as wide as the
petiole. (fig. 47) 15
15a) Body and mainly the gaster covered with numerous fine
long hairs; gaster enlarged, larger than the thorax;
the propodeal spines very thick and obliquely curved and
pointed laterad; first gastric tergite opalescent and
subopaque. (fig. 46, 47) quiescens n. sp.
b) Not with this combination of characters 16

16
16a) Postpetiole without lateral angles, hairs slender and
pointed; postpetiole and gaster smooth and shining.
(fig. 48) dentigula M.
b) Postpetiole with angular sides, seen from above; hairs
short and thick
17a) Dorsal surface of first gastric tergite finely and densily
granulse, opalescent and subopaque; dorsum of pronotum
and mesonotum punctate; dorsum of postpetiole punctate
and opaque. (fig. 50) anastas
b) First gastric tergite smooth and shining; dorsal area of
postpetiole weakly smooth and shining; dorsum of pronotum
usually with partial smooth shining areas.
R. Smith
17
ii Emery
(fig. 51)
floridana Emery

1. Pheidole ndrianoi n. sp.
Diagnosis:
A small ebony black species differing from IP. metallescens by the
absence of the violaceous or bluish reflections and without the reticulate
sculptured occipital lobes. It also differs from P. littoralis by the
small shining head, the head length usually less than 1.0 mm in the major
as opposed to over 1.4 mm in littoralis; workers with the erect hairs
plumose or semi-plumose at apex.
Description:
Major:
Measurements: Body length 2.70 to 3.10 mm. Head length 0.82
to 0.97 mm; head width 0.79 to 0.95 mm. Thorax length 0.79 to 0.90 mm;
scape length 0.45 to 0.51 mm. Scape Index 51 to 59.
Structural characters: Head shape as in fig. 25. Head, exclud
ing mandibles, usually slightly longer than wide, broadest behind
the eyes from where the sides slightly curve and narrow posteriorly,
the occipital excision gently concave. Frontal area impressed; anterior
border of clypeus with a wide, shallow median notch. A weak, scarcely
depressed antennal semi-scrobe present; a distinct frontal furrow
arising behind the frontal area, widening posteriorly into the
occipital excision. Scapes slender at base, thickening apically and
longer than the half the head length; mandibles stout, weakly curved
laterally, with two apical blunt teeth and two basal denticles. Eyes
small with approximately 35 facets.
Pronotum and mesonotum strongly convex in profile, posteriorly
the mesonotum more abruptly descending into the mesopropodeal impression.
17

18
Mesonotum without transverse impression; spines thick and long but
blunt apically and pointed obliquely dorsad. Petiole stout, anterior
and posterior faces sloping, the crest weakly angular in profile.
Seen from above, the sides of post petiole semi-angular, the
lateral projections blunt and not well developed; gaster truncate
at base and smaller than head.
Sculpture: Strong cephalic longitudinal striae diverging
posteriorly and disappearing in the area posterior to the scape when
in repose; interstitial areas between striae with weak, fine reticulate
punctation; frontal carinae diverging and surpassing the scapes when
in repose; vertex, frontal area as well as the clypeus smooth and
shining; the semi-scrobes not distinct and marked only by lack of
sculpture; the genae with strong longitudinal striae, the interstitial
areas weakly reticulo-punctate; occipital lobes smooth and shining
with sparse small circular piligerous punctures, ventral surface of
head smooth and shining, its anterior margin without teeth; frontal
furrow with its posterior half bearing transverse rugules.
Seen from above, dorsum of the pronotum and anterior mesonotum
nearly smooth and weakly shining, mesopropodeal impression and adjacent
posterior area of mesonotum reticulo-punctate; basal area of propodeum
weakly reticulate and shining; declivous face usually shining.
In profile, pronotum, mesonotum and propodeum heavily reticulate,
the inner areas finely reticulo-punctate and shining; petiole finely
reticulo-punctate. Seen from above, dorsum of pronotum and anterior
mesonotum nearly smooth and weakly shining; mesopropodeal impression
and adjacent posterior area of mesonotum reticulo-punctate; basal

19
area of propodeum reticulate and shining; declivous face with trans
verse striae and usually the interstitial areas finely reticulo-
punctate and shining; several fine interspinal striae present; dorsum
of postpetiole weakly sculptured or almost smooth and shining.
Pilosity: Hairs yellow, fine, numerous and of various lengths,
measuring 0.07 to 0.18 mm; dorsal and ventral surface of head with
semi-erect short and long hairs; dorsum of pronotum and mesonotum
with numerous predominantly long erect hairs; propodeal base with
few scattered short hairs; pedicel with numerous semi-erect long
hairs; gaster with numerous predominantly long semi-erect hairs, most
of them blunt apically. Pubescence on antennae, sparse on legs and
absent on gaster.
Color: Head and gaster ebony black; thorax and legs a little
lighter.
Worker:
Measurements: Body length 1.50 to 1.60 mm. Head length 0.44 to
0.51; head width 0.38 to 0.41 mm. Thoracic length 0.46 to 0.51 mm.
Scape length 0.40 to 0.46 mm. Scape Index 97 to 106.
Structural characters: Head, excluding mandibles, slightly longer
than broad; occipital border slightly concave medianly; frontal area
shallowly depressed; frontal carinae straight, short and slightly
divergent; scapes surpassing the occipital lobes by an amount less
than the first funicular joint; mandibles slender, apically with
two sharp teeth, the remaining masticatory border with several smaller
denticles.

20
Pronotum and mesonotum, in profile, convex; propodeal spines
long but blunt, projecting obliquely dorsad; petiole stout, the
apex of the node angular; postpetiole strongly convex dorsally and
almost flat ventrally; gaster truncate at the base.
Sculpture: Head smooth and strongly shining; pleural surfaces
of pronotum shining; pleurae of mesonotum, propodeum and petiole
strongly reticulo-punctate, postpetiole weakly punctate.
Seen from above dorsum of pronotum and anterior mesonotum smooth
and strongly shining, posterior area of mesonotum, propodeal base
and propodeal declivity reticulo-punctate; dorsum of postpetiole and
gaster smooth and shining.
Pilosity: Hairs white, measuring less than 0.08 mm; sparse all
over the body, semi-erect and of mixed sizes on the head, long and
erect on the dorsum of thorax and long and slender on gaster; under
the stereo microscope the hairs seemed spatulate, but scanning micro
graphs later revealed that such hairs are branched at the tips (fig.
53, 54), therefore, plumose or sub-plumose.
Color: Ebony black, the appendages lighter, mandibles yellowish
to dark brown.
Females:
Measurements: Body length 4.60 mm. Head length 0.90 mm; head
width 0.95 mm. Thoracic length 1.46 mm. Scape length 0.51 mm. Scape
Index 58.
Structural characters: Head broader than long, broadest posteri
orly; occipital border slightly concave; sides slightly convex and

21
narrowing anteriorly toward the mandibular insertions; anterior
border of clypeus weakly notched in the middle.
Frontal area depressed; antennae like those on the major;
scapes longer than half the head length; eyes measuring about 0.24 mm
in diameter and located on anterior half of head; ocelli measuring
about 0.12 mm in diameter.
Thorax flat dorsally, narrower than the head measured through
the wing insertions. Mesoscutum and scutellum well separated; propodeal
base and declivous face not forming a distinct angle. Propodeal spines
stout, blunt and directed posteriorly; petiole stout, the node seen
from above with parallel sides, the crest slightly depressed in the
middle; postpetiole twice as wide as petiole, with rather angular
lateral projections; gaster truncate at the base.
Sculpture: Cephalic rugae strong, covering most of the anterior
part of the head, but disappearing before reaching the occipital border,
the interstitial areas between rugae with widely spaced weak reticulate
punctation; frontal area smooth and shining with two mesal longitudinal
striae; frontal carinae long and diverging posteriorly.
Sides of pronotum and propodeum reticulate; episternum smooth
and shining; propodeal base and declivous face smooth and shining;
petiole and postpetiole punctate and opaque; gaster, seen from above,
with fine longitudinal striae near the base, the interstitial areas
between striae shining but with widely spaced reticulate punctation,
the rest shining and with circular piligerous punctures.
Pilosity: Hairs yellow, pointed; dorsum of thorax, pedicel and
gaster with numerous short and long hairs; gaster usually more pilose
than the remaining parts; pubescence present on the antennae and legs.

22
Color: Reddish brown; the gaster darker.
Male:
Unknown.
This species is named in honor of my beloved first born son,
Adriano de Resende Naves.
Holotype: Major from Gainesville, Florida, collected by M. A.
Naves.
Paratypes: The type material comprises 30 majors and 60 workers
collected in Gainesville, Florida from several colonies on August 13,
1975 and 25 majors and 33 workers from several colonies from Archbold
Biological Research Station at Lake Placid, Florida, in August, 1975.
All were collected by M. A. Naves.
The holotype and several paratypes will be deposited at the Museum
of Comparative Zoology, Harvard University, Cambridge, Massachussetts.
Paratypes will also be deposited in the National Museum of Natural
History, Washington, D.C., Florida State Museum, Department of Plant
Industry, Gainesville, Florida and in the W. F. Burn and M. A. Naves
collections.
Discussion:
This species is fairly common in Florida. Its range seems to be from
central to northern Florida. It may also occur in southern Georgia. Its
preferable habitat seems to be sandy soils, in clear areas among trees,
the same habitat as in Ij. metallescens, with which it is very similar to
the naked eye.
The colonies have about 60 majors and more than 300 workers. The main
chamber is located 30 to 40 cm deep in the soil and is directly connected
to a vertical tunnel of about 1 to 2 mm in diameter, which usually is the

23
center of a small crater on the soil surface. The main flights of sexuals
occur in the summer in July and August. The species is entirely diurnal.
The main sources of food are small seeds, although it also scavenges small
dead arthropods. The majors as well as the workers usually forage for
food and help carry it back to the colony. It is not an agressive species.
The majors when disturbed try to escape to hiding places.
f\ adrianoi differs from P_. metallescens by its ebony black color,
absence of reticulate punctation on the head and by its smooth and strongly
shining occipital lobes and lack of metallic reflections. P. adrianoi
also differs from _P. littoralis by its small head, and the occipital lobes
not strongly rounded, sides of head not parallel and mandibles not stout
and strongly curved. P_. metallescens and P_. littoralis usually have a
reddish brown head and thorax, the gaster always darker or blackish as
opposed to P_. adrianoi which has a shining concolorously ebony black
body.

24
2. Pheidole anastasii Emery
P_. anastasii Emery, 1896. Soc. Ent. Ital., Bol. 28:76. Forel,
1901. Naturhist. Mus. Hamburg, Mitt. 18:78. Wheeler, 1932. N.Y. Ent.
Soc., Journ. 40:6. Smith, 1933. Fla. Ent. 17:23. Creighton, 1950.
Bull. Mus. Comp. Zoll. 104:169. Gregg, 1958. N.Y. Ent. Soc. 66:16.
P,. floridana Van Pelt, 1956. Amer. Midland Nat., 56:376, 384.
Carroll, 1970. Unpublished M.S. thesis, University of Florida, p. 32-34. -
Whitcomb, Denmark, Bhatkar, and Greene, 197^ Fla. Ent., 55:133. (nec
Emery).
Type locality: Jimenes, Costa Rica
Types : None in the U.S.
Range : Florida, southern Alabama and Georgia
Diagnosis:
A small yellowish species differing from ?. floridana in having
opalescent surfaces dorsally on the first gastric tergite; differing
from P_. moerens and P_. greggi n. sp. by having the postpetiole twice as
broad as the petiole. Distinct lateral connules are present. It differs
from P_. dentigula in having smooth and shining occipital lobes.
Discussion:
Wheeler (1932) reported P. anastasii (fig. 17) from Florida. It seems
hard to believe that it is an introduced species as Wheeler suggested. I
have found this species throughout Florida from the keys to the Panhandle.
Its constant nesting at the base of pines and the occurrence of two parasites,
_P. quiescens n. sp., an inquiline ant, and a hymenopteran parasite in the
genus Orasema, seem to contradict Wheeler's suggestion that P. anastasii
is an exotic species.

25
Due to the close resemblance of this species to I?. floridana Emery,
P_. anastasii probably has been misdetermined in several records from
Florida.
The difficulty of having access to Emery's types of _P. anastasii
makes it difficult to determine if the Florida species is conspecific with
the Costa Rican types. Wheeler, however, did have knowledge of the types
of _P. anastasii when he reported it from Florida.
P_. anastasii was usually found nesting under the bark at the base of
pines or along the roots and only rarely in the soil. The colonies are
monogynous and have more than 100 majors and over 500 workers. Several
chambers are constructed interconnected by a string of galleries under
the pine bark. The workers forage over 4 m from the colony and once food
is located majors are recruited to help transport it back to the colony.
It feeds on seeds, fruits, and scavenges on small dead arthropods and is
predaceous on small live arthropods.
Laboratory colonies are very easy to keep and usually accept members
of other colonies. Colonies that have lost their queens will also accept
any other queens of anastasii. Several times I collected two or three
nest founding females found together in one chamber. In the lab one
female always killed the others before the first workers were reared.

26
3. Pheidole bureni n. sp.
Diagnosis:
A medium sized brownish red species with darker gaster, difering from
P. diversipilosa and P. crassicornis tetra by the absence of long erect
hairs on the dorsum of the first gastric tergite. It differs from P_.
crassicornis by having the occipital excision deeper and much more angularly
V shaped, and the occipital lobes sharper.
Description:
Major:
Measurements: Body length 5.30 to 5.70 mm. Head length 1.54
to 1.67 mm; head width 1.43 to 1.54. Thoracic length, 1.44 to 1.54
mm. Scape length, 0.85 to 0.92 mm; Scape Index, 54 to 55. Depth of
occipital excision usually over 0.12 mm.
Structural characters: Head shape as in fig. 60. Head excluding
mandibles longer than wide, broadest above the eyes, from where the
sides curve and narrow posteriorly; the occipital excision deep and
wide and angularly V shaped; occipital lobes much more angular than
in crassicornis or diversipilosa; frontal area impressed; anterior
border of clypeus with shallow median notch; irons strongly pronounc
ed and the vertex inclined posteriorly; genae flat and depressed
laterally. Frontal furrow distinct and extending from frontal area
to the occipital excision; scapes flattened and curved at base, a
little longer than half head length; mandibles curved laterally with
two strong, sharp teeth apically; the rest of the border without
teeth or with several small denticles; eyes measuring about 0.26 mm
in diameter and with over 100 small facets.

27
In profile, pronotum and anterior mesonotum convex, mesonotum
with a wide impression in the middle; posteriorly the mesonotum
abruptly descending to the mesopropodeal impression; propodeal base
longer than declivity; propodeal spines short and pointed dorsally;
petiole stout, the anterior face and posterior face sloping, the
node quite large and its crest in profile varying from weakly angular
to strongly convex; the ventral border slightly convex anteriorly;
postpetiole strongly convex dorsally and flat ventrally.
Seen from above, pronotum with pronounced but rounded shoulders;
petiole with parallel sides, the crest of node with a shallow concave
impression in the middle; postpetiole almost two times wider than
long; slightly angular to blunt laterally; gaster truncate at the
base.
Sculpture: Fine longitudinal striae present on the frons, inter
stitial areas finely and weakly reticulo-punctate; vertex and occipital
lobes smooth and shining; genae with longitudinal rugae, the inter
stitial areas weakly punctate; frontal*carinae short and slightly
diverging posteriorly; frontal area smooth and shining; clypeus smooth
and shining, usually with a weak median carinae; frontal furrow with
transverse short rugulae posteriorly; ventral area of head smooth
and shining, its anterior margin with two well developed sharp teeth.
Pronotal pleurae weakly reticulate; mesonotum, propodeum and
petiole finely reticulo-punctate, the floor of each punctation strongly
shining.
Seen from above, pronotum and adjacent area of mesonotum with
weak transverse striae and weakly reticulated; mesonotal dorsum
smooth and strongly shining; mesonotal impression not easily seen;

28
mesopropodeal impression depressed and shining; prododeal base
weakly reticulo-punctate and shining; propodeal declivity smooth
and strongly shining; petiole and postpetiole finely reticulo-
punctate and opaque; gaster smooth and shining.
Pilosity: Hairs yellow and pointed, measuring up to 0.26 mm;
vertex with two long hairs on each side and several along the frontal
carinae; anterior ventral face of head with semi-erect short hair;
posterior of head and mandibles and scapes with fine appressed
pubescence; a few erect hairs on the scapes; dorsum of pronotum
and mesonotum with a few mixed short and long erect hairs; scattered
appressed pubescence on pronotum; each side of the petiolar node
with long semi-erect hairs; postpetiole with several semi-erect
long hairs; first gastric tergite with numerous fine appressed
pubescent hairs; ventrally the posterior half of gaster with mixed
short and long erect and semi-erect hairs; appendages with sparse
appressed pubescence; funiculus with very fine pubescence.
Color: Reddish brown, the gaster usually darker.
Worker:
Measurements: Body length 3.10 to 3.20 mm. Head length 0.77
to 0.79 mm; head width 0.70 to 0.77mm. Thoracic length 1.05 to 1.13.
Scape length 0.82 to 0.90. Scape Index 106 to 114.
Structural characters: Head, excluding mandibles, a little
longer than broad; occipital border flat to slightly concave; sides
strongly convex; frontal area broader than long and shallow; frontal
carinae short and parallel; scapes surpassing the occipital border

29
by an amount greater than the first funicular joint; mandibles slender,
curved apically and with two sharp -teeth, the rest of the border
with several small denticles; anterior clypeal border convex; eyes
measuring 0.18 mm in diameter and with more than 65 facets.
In profile, pronotum and anterior mesonotum strongly convex;
mesonotum with a wide transverse impression; propodeal base longer
than the declivity; propodeal spines short and obliquely pointed
dorsad; petiole stout, the node strongly convex in profile; post
petiole strongly convex dorsally in profile and flat ventrally.
Seen from above, postpetiole 1.5 times wider than long and the
sides strongly convex; gaster truncate at the base.
Sculpture: Head finely reticulo-punctate except the irons and
vertex which have smooth and shining areas; frontal area smooth and
shining; clypeus with a median carina and usually smooth or weakly
reticulo-punctate; genae with several short longitudinal striae
emerging from anterior reticulation.
Pleurae of throax reticulo-punctate; petiole and postpetiole
finely reticulo-punctate; gaster smooth and shining.
Dorsum of mesonotum with small smooth and shining areas; basal
face of propodeun strongly reticulo-punctate and the declivity
weakly reticulate and shining.
Pilosity: Hairs yellow, numerous, measuring up to 0.20 mm;
mixed short and long semi-erect hairs present on head; erect, short
and long hairs on the dorsum of thorax; two long semi-erect hairs on
the pedile; numerous short and long, erect and semi-erect hairs on
postpetiole and gaster.

30
Color: Concolorously dark brown.
Female:
Measurements: Body length 7.50 mm. Head length 1.20 mm; head
width 1.38 mm. Thoracic length 2.41 mm. Scape length 1.03 mm. Scape
Index 86.
Structural characters: Head broader than long, broadest posterior
ly; occipital border straight; sides slightly convex and narrowing
anteriorly; middle of anterior border of clypeus with an impression;
frontal area broad and depressed; scapes stout, longer than 3/4 the
head length, the basal portions strongly curved and flattened, the
flattening more latero-mesal rather than dorso-ventral as in a major.
Eyes measuring about 0.40 mm in diameter and with over 200 small
facets; ocelli measuring about 0.15 mm in diameter; frontal carina
expanded and raised. Thorax flat dorsally, measured through the
Wing insertions slightly wider than the head; mesoscutum and scutelum
separated by a distinct suture.
Propodeal base and declivity not forming a distinct angle;
propodeal spines very short, thick and pointed dorsad; petiole stout; post
petiole twice as broad as long, the sides produced into distinct,
blunt lateral projections; these slightly convex anteriorly and
posteriorly weakly concave; gaster truncate at the base.
Sculpture: Frontal area and clypeus smooth and shining as well
as a narrow area of the frons between the frontal area and the median
ocellus; frons and lower vertex longitudinally striate laterally;
posterior areas of head smooth and shining; genae with pronounced

31
rugae; frontal carinae short and diverging posteriorly; ventral
surface of the head smooth and shining, its anterior margin with
three distinct teeth, the median tooth short and lobe-like and the
lateral ones long and sharp; sides of pronoturn opaque; esisternum
smooth and shining; propodeum with transverse striae, the inter
stitial areas weakly punctate and shining; propodeal base and
declivity smooth and shining; sides of pedicel finely reticulo-
punctate; gaster smooth and shining.
Pilosity: Hairs yellow and pointed and measuring up to 0.40 mm
in length; head with numerous short and long erect and semi-erect
hairs; dorsum of thorax and petiole with numerous short and long
pointed hairs; postpetiole and gaster with numerous, predominantly
short semi-erect hairs; short sparse fine and pointed semi-erect
hairs on scapes and apendages.
Color: Light reddish brown
Male: Unknown
Holotype: Major, collected at Alice, Texas
Paratypes: 6 majors, 7 workers and 1 female, from the same
colony as the Holotype.
The type material comprises part of a colony collected by William
F. Burn on October 23, 1942 at Alice, Texas.
The holotype, the female paratype and one worker paratype will be
deposited in the Museum of Comparative ^oology, Harvard University,
Cambridge, Massachussetts. Paratypes will also be deposited in the
National Museum of Natural History, Washington, D.C., the Florida State

32
Museum, Department of Plant Industry, Gainesville, Florida, and in the
W. F. Burn and M. A. Naves collections.
Discussion:
This species is named in honor of Dr. William F. Burn, Professor
of Entomology, University of Florida.

33
4. Pheidole carrolli n. sp.
Diagnosis:
A reddish brown species related to the P_. pilifera group. It
differs from P_. pilif era and P_. pilif era artemisia by the absence of
coarse reticulate occipital rugae and blunt postpetiolar connules. It
differs from P_. pilif era coloradensis by its larger head, strongly
convex pronotum and anterior mesonotum, and longer and finer thoracic
and gastric hairs. It differs from P. pilifera pacifica by its larger
size and cephalic rugae extending to the occipital lobes and absence of
transverse occipital rugae.
Description:
Maj or:
Measurements: Body length 3.20 to 3.60 mm. Head length
1.80 to 1.90 mm; head width 1.69 to 1.80 mm. Thoracic length 1.36
to 1.44 mm. Scape length 0.77 to 0.82 mm. Scape Index 42 to 44.
Structural characters: Head shape as in fig. 14. Head, exclud
ing mandibles, longer than broad, occipital lobes prominent and well
rounded, separated by a broad and deep excision; a furrow present,
starting from the frontal area to the occipital excision. Frontal
area depressed and bearing a median longitudinal striation. Clypeus
with a weak median carinae which is continuous with the frontal area
striation; anterior border of clypeus with a shallow weak emargina-
tion; frontal carinae divergent; anterior ventral margin of the head
without teeth; scapes slender at base, thickening apically and measur
ing less than half the head length. Mandibles stout, strongly curved
laterally and with two apical large blunt teeth, the remaining border

34
edentate, except for a small basal tooth-like lobe. Eyes small,
measuring about 0.25 mm in diameter with over 60 facets and located
on the anterior third of the head. Seen from the side, thorax
smaller than head; pronotum and anterior mesonotum strongly convex;
posterior of mesonotum descending to the mesopropodeal impression,
forming an obtuse angle with the propodeal base; base of propodeum
slightly shorter than declivity; propodeal spines short and project
ing dorsad; petiole, in profile, stout and with a short anterior
peduncle, anterior face gently curved and posterior face short; apex
of node weakly angular to moderately convex; seen from above the
sides of the petiole parallel and the node notched in the middle;
postpetiole in profile strongly convex dorsally, slightly concave
and much shorter ventrally; seen from above, postpetiole two times
or more wider than petiole and with lateral connules; gaster oval
and much smaller than the head.
Sculpture: Cephalic longitudinal rugae extending posteriorly
and somewhat divergently; the lateral rugae usually do not reach
the occipital lobes; occipital lobes smooth and shining laterally
and dorsally; rugae present on the genae, extending longitudinally;
posterior to the eyes the surfaces smooth and shining; head posteriorly
with coarse circular piligerous punctures. Dorsum of pronotum and
mesonotum with longitudinal or transverse rugae or both, the inter
stitial areas with fine reticulate punctation. The mesopropodeal
impression wide and with longitudinal rugules, the interstitial
areas smooth and weakly shining; the propodeal base and declivity

35
finely reticulo-punctate and weakly shining. Several transverse
striae present between bases of spines; petiole and postpetiole
opaque and weakly reticulo-punctate; gaster smooth and shining
except in the basalmost area where it is opaque and weakly punctate.
Pilosity: Hairs yellow, pointed, short and semi-erect and sparse
on the anterior face of the head, short and long and semi-erect
ventrally; dorsum of thorax with numerous short and long erect
hairs measuring up to 0.26 mm; pedicel with numerous semi-erect
short and long hair; sparse pubescence confined to antenae and
appendages.
Color: Reddish brown, appendages lighter; dorsum of gaster
slightly darker; anterior margin of clypeus and the border of
mandibles dark.
Worker:
Measurements: Body length 2.30 to 2.40 mm. Head length 0.56
to 0.64 mm; head width 0.56 to 0.58 mm. Thoracic length 0.74 to 0.77
mm. Scape length 0.53 to 0.59 mm. Scape Index 92 to 94 mm.
Structural characters: Head usually a little longer than broad,
occipital borders weakly convex; frontal area impressed and with a
longitudinal striation in the middle; frontal carinae slightly divergent;
clypeus with a weak median carina and 2 longitudinal striae laterally;
anterior border curved and without any notch. Mandibles slender with
2 large and sharp apical teeth and 4 or 5 smaller teeth on the remain
ing masticatory border; scapes usually surpassing the occipital border
by an amount no greater than the first funicular joint; eyes large
with about 45 facets and measuring about 0.15 mm in diameter.

36
In profile, pro-mesonotum moderately convex; propodeal spines
small, sharp, and projecting obliquely dorsad; petiole as in the
major, except the crest is proportionately much broader; postpetiole
convex dorsally and flat ventrally, seen from above the sides are
strongly convex and almost 2 times wider than petiole; gaster oval.
Sculpture: Head reticulo-punctate and opaque; a few rugules
on the genae, between the eyes and on the mandibles; thorax, in
profile, reticulo-punctate; dorsum of pronotum and mesonotum usually
with a few longitudinal or transverse striae; petiole reticulo-
punctate; sides of postpetiole weakly punctate, dorsum weakly shin
ing; gaster smooth and shining.
Pilosity: Erect hair yellow, short on the dorsal surface of
head, longer on the ventral surface and anterior clypeal margin;
long and sparse on thorax and pedicel; gaster with rather sparse
hair of mixed lengths; pubescence confined to the antennae and legs.
Color: Reddish brown, sometimes the mandibles and legs are
lighter.
Holotype: Major, collected near Gainesville, Florida by M. A.
Naves.
Paratype: 3 majors and 20 workers probably from the same nest
as the holotype, collected as follows:
Dr. John F. Carroll collected one major and several workers on
August 2, 1973, near Gainesville, Florida. M. A. Naves collected
three majors and several workers in August, 1975, at the same locality.

37
The holotype major and 3 worker paratypes will be deposited at
the Museum of Comparative Zoology, Harvard University, Cambridge,
Massachussetts. Paratypes will also be deposited in the National
Museum of Natural History, Washington, D.C., Florida State Museum,
Department of Plant Industry, Gainesville, Florida and in the M. A.
Naves collections.
This species was also collected by me at Tall Timbers Research
Station, but the specimens, unfortunately, were inadvertantly lost.
Discussion:
This species has unusual behavior which is not common to other Pheidole
known to me. It was always found in shaded areas in sandy soils among tall
trees such as pines and oaks. It forages when the temperatures are not
high and avoids direct sunlight. It is easily found at dusk or in the
morning. The workers usually go out at random. Usually there are no
more than 15 workers foraging at one time, and the foraging territory
usually is within 5 meters from the nest entrance. The nest entrance is
small and usually very hard to spot. The worker is sluggish compared with
most other species. The majors are very rare. For a year two colonies
were periodically checked and yielded only one callow major in 1975 and
two majors in March, 1976. Dr. John F. Carroll also collected one major
in the same locality in August, 1973. Periodic excavation usually also
yielded a chamber approximately 4X2X2 cms located 30 cm deep and full of
various seeds, some apparently with a black fungal mycelium. Although the
excavation for one of these colonies at one time was 1.7 meters deep 1.5
meters in diameter, I was unable to locate the main chamber or any majors

38
or any sexual forms. Probably the main chamber or chambers are located
very deep in the ground. It seems quite obvious that the colonies are
not large. The disproportional size of the major and its rarity also
seems to indicate that it has limited importance in sustaining the colony,
which is not true for most species of the genus.
Another feature of this species when disturbed is the ability of the
workers to flign death. The minors try to flex themselves around particles
of soil which make them very difficult to see. The majors hide themselves
and also seem to avoid light, but their large size and more shining surfaces
compared with the workers make them easier to spot.
Workers of this species, although not many, have been captured by
baiting near Gainesville.
One colony was located at Tall Timbers Research Station in 1974, which
failed to yield any major or sexual form.
Perhaps P. carrolli is a species which may be losing its major in the
evolutionary process. The fewer majors available show rather more mor
phological variation than is normal in most Pheidole species.
This species is named after Dr. John F. Carroll, my friend and colleague
during several years of graduate study at the University of Florida.

39
5. Pheidole crassicornis Emery
P. crassicornis Emery, 1985. Zool. Jahrb., Abt. F. System. 8:289,
296. Forel, 1901. Soc. Ent. Belg., Ann. 45:350. Wheeler 1904. Amer.
Mus. Nat. Hist., Bui. 20:302. Creighton, 1950. Bull. Mus. Comp. Zool.
104:175-176. Gregg, 1958. N.Y. Ent. Soc., 66:20.
Type locality: Charlotte, North Carolina
Types : Cotypes, M. C. Z. and A. M. N. H.
Range : Southeastern States
Diagnosis:
P_. crassicornis (fig. 5) differs from P_. diversipilosa and Py cras
sicornis tetra by the absence of erect and semi-erect hairs on the gaster.
It also differs from P. bureni n. sp. by the convex occipital lobes and
the occipital impression which is not sharply V shapped. P_. crassicornis
can be easily distinguished from all Florida Pheidole other than diversipilosa
by the basally strongly bent and flattened scapes.
Discussion:
The finding of P. crassicornis at Gainesville and at Tall Timbers
Research Station will help the understanding of this group because it is
sympatric with P_. diversipilosa at Tall Timbers. P. crassicornis is
apparently confined to the northern part of Florida where it occurs in open
areas, usually among trees. Its nests seem to be in deep soil. After digging
as deep as 60 cm, I was unable to find any chambers of three colonies. The
opening at the soil surface is hard to see. The typical craters common
to other species were never observed. The major and workers were seen carry
ing live termites and other small dead arthropods. The workers and a few
majors were observed foraging approximately 4 meters or more from the nest
opening.

40
6. Pheidole dentata Mayr
P_. morrisi var. dentata Mayr 1886. Vehr. Zool Bot. Ges. Wein,
36:457.
P. dentata Forel, 1901. Ann. Soc. Ent. Belg., 45:351 Wheeler,
1904. Amer. Mus. Nat. Hist., Bull. 20:302. Mitchell and Pierce, 1912
Ent. Soc. Wash., Proc. 14:71. Smith, 1918. Ent. News, 29:22. Smith,
1924. Ent. News 35:77. Smith, 1930. Fla. Ent. 14:3. Wheeler, 1932.
N.Y. Ent. Soc., 40:6. Dennis, 1938. Ent. Soc. Amer., Ann. 31:281,
304. Van Pelt, 1950. Ent. News 61:161-163. Creighton, 1950. Bull.
Mus. Comp. Zool. 104:177. Wheeler and Wheeler, 1953. Ent. Soc. Wash.
Proc. 55:71. Gregg, 1953. N.Y. Ent. Soc. 66:21-22. Smith, 1965.
U.S. Dept. Agr., Tech. Bui. 1326, pp. 27-28. Wilson, 1975. Science
190:798-800.
P. dentata var. faisonica Forel., 1901. Ann. Ent. Soc. Belg.,
45:352.
P. commutata Mayr 1886. Verb. Zool. Bot. Ges. Wein. 36459. -
Emery, 1895. Zool Jahrb. Syst., 8:289.
Leptothorax tennesseensis Cole, 1938. Amer. Midland Nat. 19:238.
fh dentata var. commutata Wheeler, 1907. Amer. Mus. Nat. Hist.,
Bull. 7:18. Wheeler, 1908. Amer. Mus'. Nat. Hist., Bull. 24:460-461.
Smith, 1924 Ent. News 35:77. Cole, 1940. Amer. Midland Nat. 24:29,
44. Smith, 1951. Cat. Hym. Amer. No. of Mex., U.S.D.A. Mon 2:802.
Type locality: Florida
Types : None in this country
Range : Southeastern States, Tennessee and Gulf states to
Texas.

41
Diagnosis:
]?. dentata (fig. 30) is a reddish yellow to dark brown species
differing from P_. morrisi by the distinct and sharp propodeal spines.
It differs from P_. megacephala in not having a cordate head and differs
from P. vinelandica by the larger size and absence of a longitudinal
striation on the frontal area, and having the mesonotum transversely
impressed in the middle.
Discussion:
P. dentata is one of the more common species in Florida. It is
easy to understand why this species caused so many misunderstandings in
the past. There is much variation, not only in size and color, but
also in habitat preference. I have found colonies of small dark colored
specimens in wooded areas around Gainesville and large specimens that
nest in sandy soil on beaches in south Florida and the Florida Keys. I
also found a yellowish variant that inhabits the marshlands of the keys
and another variant with quite large majors that vary in color from
reddish to very dark brown nesting in open areas around Gainesville. This
morphological variation seems to be without taxonomic significant^
Three mated young queens collected at Gainesville were easily reared
in the laboratory. All three had majors after 50 days. The species is
highly carnivorous and the colonies have a tremendous growth when fed
with living arthropods. One of the colonies was inadvertantly left with
out food and moisture for three weeks. The result was that the adults
formed from the stressed broods were intermediate in size and structure
between workers and majors. Some of the intermediates were not as active

42
as the normal caste. This is a very interesting species for ecological
and laboratory studies. Recent studies by Wilson (1975) have shown that
workers of P. dentata use odor trails to recruit nestmates to food
discoveries and new nest sites. The same pheromone is used also to
recruit major workers to the vicinity of intruders. This type of alarm
recruitment proved rather narrowly specific and works best against some
of the potentially more important enemies of P. dentata in the genus
Solenopsis.

43
7. Pheidole dentigula M. R. Smith
P. dentigula Smith, 1927. Ent. News 38:310. Smith, 1928. Ent.
News 39:245-246. Cole, 1940. Amer. Midland Nat. 24:42, 45. Smith,
1944. Fla. Ent. 27:14. Creighton, 1950. Bull. Mus. Comp. Zool, 104:
178. Gregg, 1958. N.Y. Ent. Soc. 66:21. Wheeler and Wheeler, 1960.
Ent. Soc. Wash., Proc. 62:14.
Type locality: Mississippi State University, Starkville, Mississippi.
Types : Cotypes, National Museum
Range : Tennessee, southward through Alabama and Mississippi,
and north Florida
Diagnosis:
P_. dentigula (fig. 13) is a small yellowish species that differs
from _P. greggi, P_. anastasii, P_. floridana, P_. tysoni, P_. bicarinata
and Ph moerens by the reticulated, opaque occipital lobes. It differs
from P_. metallescens by having a smooth and shining propodeal base and
lacks metallic reflections. The teeth on the anterior ventral border of
the head are prominent.
Discussion:
Smith (1944) reported P. dentigula from Wakulla County (Florida).
Dr. John F. Carroll also collected this species in Marion County (Florida)
on April 3, 1973. According to Smith (1944) this species nests in the
soil and in well rotted stumps in wooded areas, especially where the soil
conatins considerable hummus.

44
8. Pheidole diversipilosa Wheeler
P. crassicornis var. diversipilosa Wheeler, 1908. Amer. Mus. Nat.
Hist., Bui. 24:467.
P. crassicornis Creighton, 1950. Bull. Mus. Comp. Zool., 104:175-
176.
Type locality: Fort Davis, Texas.
Types : M.C.Z.
Range : Northwestern Florida to western Texas
Diagnosis:
]P. diversipilosa (fig. 3) is a reddish yellow species that differs
from P. crassicornis and P. bureni n. sp. by the numerous erect hairs
on the gaster. In P. crassicornis tetra the gastric hairs are nearly
all of the same short length rather than obviously of mixed lengths
as in diversipilosa.
Discussion:
P. diversipilosa was collected at Tall Timbers Research Station
several times where F\ crassicornis is also present. Morphological
distinction is constant between the two species. There is no real over
lap in the pilosity characters. Therefore, I have treated F\ diversipilosa
as a full species on the basis that it can be sympatric with P. crassicornis
but still retains constant morphological distinction. Creighton (1950)
treated diversipilosa as an intergrade between crassicornis and tetra.
The finding of both crassicornis and diversipilosa in the same locality
in Florida invalidates Creighton's viewpoint.
Specimens of _P. crassicornis collected in Gainesville were within
the variation shown by the Tall Timbers specimens. I was unable to find

45
specimens of P_. diversipilosa in Gainesville. Comparison with the types
and cotypes for the two taxa in the M. C. Z. and further material of
diversipilosa collected at Columbus and Mission, Texas by W. F. Burn
as well as the finding of another related species, P_. bureni n. sp.
(fig. 60, 61, 62) emphasize that we are dealing with a species group
that is widely spread and contains several discrete species. P. crassicornis
is predominantly an eastern species. P. diversipilosa, P_. bureni and
P_. crassicornis tetra are western species. Of these apparently only
P_. diversipilosa reaches east into Florida. The lack of better records
for the distribution of these species probably led Creighton (1950) to
synomymizing crassicornis and diversipilosa. _P. crassicornis tetra
appears to be a very distinct form and may be a discrete species also,
but I have not attempted taxonomic treatment in the present paper, since
it does not occur in Florida.

46
9. Pheidole fallax obscurithorax Santschi
Ht4
P. fallax arenicola var. obscurithorax Santschi, 1923. As#n>. Soc.
Ent. Belg. 63:58. Kempf, 1972. Studia Ent. 15:191.
Type locality: Cordoba, Argentina
Types : None in this country
Range : Probably westernmost Florida to at le^st Mobile,
Alabama in U.S. Probably widely ranging in South
America.
Diagnosis:
P_. fallax obscurithorax (fig. 7, 8), is a large, very dark species
over 6 mm in body length. Its characteristics, such as the heavily
sculptured head and thick scape which is curved meslally and more angular
laterally are unique among the Pheidole of North America.
Discussion:
_P. fallax obscurithorax (fig. 8) is a member of the difficult large
neotropical fallax group which has 6 subspecies and 6 varieties. The
specimens collected near the Florida border matched specimens in the
M.C.Z. identified by W. W. Kempf.
This species was collected close to the Florida and Alabama border
in Balbwin County, Alabama by Dr. John F. Carroll. E. 0. Wilson also
(personal communication) collected this species in Mobile, Alabama in
1950 within 1/2 mile of the ship docking area. It is apparent that this
introduced species has been in the U.S. for at least 26 years but, unlike
the imported fire ant, has not spread widely.
It is not a surprise to find another exotic ant species from South
America in the U.S. Brachymyrmex patagonicus Mayr (not published),

47
Iridomyrmex humilis (Mayr), Solenopsis invicta Burn, richteri
Forel, and Pheidole fallax obscurithorax appear to be a group of South
American ants inadvertently imported to the ports of New Orleans, and/or
Mobile, and/or possibly Pensacola.
All five species have ranges .along or near the Paraguay and La Plata
Rivers in Western Brazil, Paraguay and Argentina. The Paraguay River is
navigable by ocean going vessels up river to Asuncion, Paraguay.

48
10. Pheidole floridana Emery
Z* flavens floridana Emery, 1895. Zool. Jahrb., Abt. f. System.
8:293. Smith, 1930. Fla. Ent. 14:3.
£. floridana Emery, 1895. Bull. Soc. Ent. Ital. 28:77. Wheeler,
1932. N.Y. Ent. Soc., 40:6. Creighton, 1950. Bull. Mus. Comp. Zool.
104:179. Smith, 1951. Cat. Hym. Amer. No. of Mex., U.S.D.A. Mon.
2:802. Gregg, 1958. N.Y. Ent. Soc. 66:21-22, 35. Kempf, 1972.
Studia Ent. 15:192.
Type locality: Coconut Grove, Florida
Types : M.C.Z.
Range : Southwest Florida
Diagnosis:
P. floridana (fig. 19) is a small yellowish species differing from
.P. greggi n. sp. and P_. moerens by the distinct lateral connules on the
postpetiole. It differs from P. anastasii by the absence of opalescence
on the dorsum of the first gastric tergite.
Discussion:
jP. floridana seems to be confined to southeast Florida in the Miami
area. This is the only place where I was able to locate this species.
Due to its close relationship to ]?. anastasii the latter has been mis-
identified as P_. floridana many times, thus, mistakenly extending the
supposed range of P. floridana P. anastasii is actually the species
widely distributed in Florida, while floridana is absent or at least
must be rare in most of the state.

49
11. Pheidole greggi n. sp.
P. flavens sculptior M. R. Smith, 1951. Cat. Hym. Amer. No. of
Mex. U.S.D.A. No. 2, p. 802 (nec Forel).
P. flavens Gregg, R. E. 1958. N.Y. Entomol. Soc. 66:21 (nec
Roger).
Diagnosis:
A small yellowish species closely related to P. flavens Roger, P^.
flavens sculptior Forel, _P. floridana Emery, P. anastasii Emery, and
P^. moerens Wheeler. Antennal scrobe weakly depressed but punctate and
opaque. Frontal carinae not surpassing apices of scapes in repose.
Postpetiole without lateral projections. P. greggi has been previously
confounded with P. flavens sculptior and P. flavens.
Description:
Maj or:
Measurement: Body length 2.60 to 2.90 mm. Head length 0.82
to 0.89 mm; head width 0.79 to 0.82. Thoracic length 0.72 to 0.77 mm.
Scape length 0.41 to 0.46 mm. Scape Index 52 to 56.
Structural characters: Head shape as in fig. 21; head, exclud
ing mandibles, usually slightly longer than broad, broadest behind
the eyes. The occipital lobes strongly curved and convex laterally
but less curved or even angular mesally; median occipital sulcus
shallow. Frontal area small and depressed; antennal scrobes weakly
or not at all depressed. Clypeus with weak median carina, its
anterior margin with weak median emargination. Frontal carinae
short and divergent. Scapes slender at base, widening apically and
usually half or more the head length; mandibles stout, each curved

50
apically and bearing two teeth, the remaining border edentate,
except for a small basal tooth. Eyes small and with approximately
23 facets.
In profile, pronotum and mesonotum strongly convex and descend
ing through an abrupt declivity to the mesopropodeal impression
which is rather weakly impressed; propodeal spines small and sharp,
pointed obliquely dorsad; petiole with a short anterior peduncle;
anterior face of node gently sloping, the posterior face sharply
declivous; crest of node convex and blunt in profile but not thicken
ed and sub-truncate as in anastasii and floridana, ventrally the
petiole straight rather than convex as in anastasii and floridana;
postpetiole strongly convex dorsally, flat to slightly concave
ventrally.
Seen from above shoulders of pronotum well rounded; sides of
petiole parallel; postpetiole small; the sides convex and the shape
sub-trapezoidal, without lateral connules, little wider than petiole;
gaster truncate at base and slightly smaller than the head.
Sculpture: Head opaque, cephalic rugae sub-parallel mesally,
slightly diverging laterally and reaching close to occipital lobes
before disappearing; the interstitial areas between rugae with weak
reticulate punctation; the occipital lobes smooth and shining; the
scrobes finely reticulo-punctate; genae and areas posterior to the
eyes, as well as the area anterior to the eyes longitudinally rugose,
the rugae disappearing before reaching the occipital lobes, occipital
lobes with fine circular piligerous punctures.

51
Sides of thorax, petiole and postpetiole reticulo-punctate,
opaque to subopaque. Pronotal dorsum with transverse irregular
striae; the interstitial areas weakly reticulo-punctate to smooth
and weakly shining; mesopropodeal impression wide and with longi
tudinal rugulae; the interstitial areas smooth and shining; propodeal
base reticulo-punctate and opaque; postpetiolar dorsum smooth and
shining; gaster smooth and shining.
Pilosity: Erect hair yellow, pointed, numerous; both short
and long hairs on head, thorax, pedicel and gaster; pubescence
sparse on antennae and legs, absent on gaster.
Color: Concolorously yellow
Worker:
Measurements: Body length 1.50 to 1.80 mm. Head length 0.46
to 0.52 mm; head width 0.43 to 0.46 mm. Thoracic length 0.41 to 0.59 mm.
Scape length 0.41 to 0.43 mm. Scape Index 82 to 89.
Structural characters: Head as in fig. 45, slightly longer than
broad, broadest through the eyes; posterior border of head slightly
notched medially; clypeal border curved and even; clypeus with a
median and 2 lateral striae; frontal area depressed and with a
median longitudinal striation; frontal carinae sub-parallel; antennal
scapes slender, more robust apically, reaching or surpassing the
occipital border by an amount less than first funicular joint; eyes
with less than 20 facets.
In profile, pronotum and mesonotum strongly convex and descend
ing sharply to the mesopropodeal impression, the latter with lon
gitudinal rugulae. Propodeal spines sharp and pointed obliquely

52
dorsad; petiole slender anteriorly. Seen from above, postpetiole
sub-globular and slightly wider than the petiole.
Sculpture: Head reticulo-punctate; frontal area and clypeus
weakly reticulo-punctate and opaque; mandibles with piligerous
punctures; thorax, basal and declivious face of propodeum as well
as the petiole reticulo-punctate; dorsum of postpetiole and gaster
smooth and shining.
Pilosity: Hairs yellow, dorsal surface of head as well as the
ventral surface with numerous mixed short and long erect and semi-
erect hairs; dorsum of thorax with perdominantly long erect hairs;
pedicel and gaster with long semi-erect numerous hairs.
Color: Concolorously yellow
Female and Male:
Unknown
Holotype: Major (collected on December 19, 1945 in Miami,
Florida by W. F. Burn).
Paratypes: 39 majors, 26 workers
Discussion:
The type material comprises23 majors and 16 workers, collected on
December 19, 1945 in Miami, Florida, by W. F. Burn; 6 majors and 6 workers
collected in 1975 in Baldwin County, Florida by D. P. Wojcik; 10 majors
and 8 workers collected on November 25, 1975, in Naples, Florida by M. A.
Naves.
The holotype and several paratypes will be deposited in the Museum
of Comparative Zoology, Harvard University, Cambridge, Massachussetts.
Paratypes will be deposited in the Nation Museum of Natural History,
Washington, D.C., Florida State Museum, Department of Plant Industry,
Gainesville, Florida; and in the W. F. Burn and M. A. Naves collections.

53
The species is named in honor of Dr. Robert E. Gregg, Professor
of Entomology, University of Colorado.
Gregg (1958) expended a great deal of effort trying to identify
a sample of this species that he and Mrs. Gregg collected in Brickel
Hammock, Miami, Florida. In his own words:
The specimens of sculptior from Miami, Florida (Burn
det.) and the sample obtained from Brickel Hammock are
indistinguishable. Despite the similarities of the
scrobes and their sculpture, these ants are not identical,
however, with the examples of sculptior from Puerto Rico
and Martinique, nor with the types from St. Vincent. This
is evident especially from the smooth (almost sculpture
less) and shining occipital lobes of the Florida ants. In
sculptior, the cephalic rugae and punctures completely
cover the head, making it opaque, except at the extreme
posterior margin around the foramen which is smooth and
shining. In addition, the epinotal spines of the Brickel
Hammock ants are longer and sharper than those of sculptior.
These two samples of the flavens group in Florida cannot
represent P_. flavens sculptior (from the West Indies), and
unless this subspecies is known from Florida by other
specimens, or is there but not yet collected, this discovery
requires a revision of the North American list. Pheidole
flavens sculptior must be dropped, and that is the plan
followed in this paper.
(p. 45)
Fig. 21 shows the head of P_. greggi from the original specimens
collected at Miami, Florida, by W. F. Burn. This confirms Gregg's
conclusion that they are not conspecific with sculptior. The occipital
lobes of P.. flavens sculptior from St. Thomas, Virgin Islands, also
collected by Burn on January 18, 1952, are shown in fig. 56.
The specimens collected at Naples were from a colony at the base
of a telephone pole on sandy soil close to the beach. The colony was
quite sizeable as in _P. anastasii, P. floridana and P. moerens.

54
Although Gregg showed that Burn's original identification of
sculptior from Florida specimens was in error, and that apparently
sculptior does not occur in the U.S., he left unresolved the question
of what name should be applied to the Florida species. The present
studies indicate that it is a new, undescribed species distinct from
P_. flavens Roger (fig. 55), P_. flavens sculptior Forel, ]?. floridana
Emery, P^. anastasii Emery and P_. moerens Wheeler.
Much more study is needed in order to fully understand this
difficult group which has several related species, 8 subspecies and
13 varieties in the new world.
This new species differs from _P. anastasii and P.. floridana by
the shape of the postpetiole which is slightly larger than the petiole
and does not have the distinct lateral connules. It differs from P_.
flavens by having the scrobe heavily reticulo-punctate, and the vertex
not smooth and shining. It differs from P. flavens sculptior by
having smooth and shining occipital lobes. It also differs from P_.
moerens by having a reticulo-punctate scrobe, which is opaque as
well as the entire head.

55
12. Pheidole lamia Wheeler
P. lamia Wheeler, 1901. Amer. Nat. 35:534. Wheeler, 1908. Amer.
Mus. Nat. Hist., Bui. 24:477-478. Wheeler, 1910. Ants, pp. 212, 248. -
Wheeler, 1926. Ants, pp. 212, 248. Smith, 1931. Ent. News 42:21-22.
Creighton, 1950. Bull. Mus. Comp. Zool. 104:182. Smith, 1951. Cat.
Hym. Amer. No. of Mex. U.S.D.A. Mon 2:803. Gregg, 1956. Ent. News
67:37-39. Gregg, 1958. N.Y. Ent. Soc. 66:23.
Type locality: Austin, Texas
Types : M.C.Z., A.M.N.H.
Range : North Florida to Texas
Diagnosis:
P. lamia is a small, yellowish, shining species with a subcylindrical
head in the major which is truncate anteriorly. The soldier of P. lamia
has the most striking head shape in the genus.
Discussion:
The head shape of P. lamia led Wheeler (1908) to suggest that it was
a phragmotic ant. He also believed it to be a very rare species. Burn
et al. (In Press), showed by laboratory and field observations that phragmosis
does not occur in this species.
At Tall Timbers Research Station where this species is common, I
had the opportunity to observe the biology of this interesting species.
Colonies are monogynous and sometimes have up to about 200 majors and
1,000 workers. The nests are in the ground, usually in grassy areas as
well as among taller vegetation. The nesting sites, at least at Tall
Timbers, can be easily spotted in the morning due to the reddish clay

56
soil cast up in very small honey-combed hummocks. These clumps consist
of fine soil particles and can have several openings connected to a vertical
tunnel of about 2 mm in diameter. From the vertical tunnel branch single
secondary galleries each directly connected to an oval chamber of about
0.5 to 1.5 cm in height and about 4 to 6 cm in length. Each colony has
several chambers. The first is found usually over 40 cm deep. I have dug
over 1.5 m deep following the main tunnel without reaching the end of it
in four colonies. In the summer of 1973, I found the first female after
digging seven colonies. In July, 1974, I found about thirty males in
tunnels close to the soil surface. In July, 1975, I found 18 females
inside a large chamber of about 50 cm deep. One mating pair that fell on
the ground was collected in the late afternoon on July 15, 1974. The soil
where this species nests usually has the first 30 cm predominantly sandy.
Deeper there is a reddish clay where this species has its chambers.
This species usually forages in the morning or when there is no
sunlight. Majors sometimes are found among debris with the workers.
The pale yellow color of this species also could indicate that this species
is nocturnal. The workers were observed carrying small seeds, small insects
and mites to the nest. Some majors in the chambers might be repletes since
they have quite extended gasters full of liquid. Burn et al. (In Press)
show that this species has stylized defenses against Solenopsis
(Diplorhoptrum) spp.

57
13. Pheidole littoralis Cole
P. sitarches littoralis Cole, 1952. Ann. Ent. Soc. Amer. 45:443. -
Gregg, 1958. N.Y. Ent. Soc. 66:30-39.
Type locality: Lido Beach, Sarasota, Florida
Type : A. C. Cole collection; M.C.Z.: Nat. Mus., A.M.N.H.
and W. S. Creighton collection. Both the A. C. Cole
collection and W. S. Creighton collection are now at
the Los Angeles County Museum of Natural History.
Range : Central Florida
Diagnosis:
P_. littoralis (fig. 20) has a reddish brown head and thorax, the gaster
black and shining. It differs from P. sitarches, P.. sitarches compestris
and ]?. sitarches soritis by the presence of deep, large piligerous foveolae
on the occipital lobes which also have no distinct transverse striae.
Discussion:
P_. littoralis is a distinct species related to the sitarches complex
as Cole (1952) suggested. This species was collected nesting in white
sand at the Archbold Biological Research Station, where it is very common.
It was found nesting side by side with £. metallescens and P. adrianoi n.
sp. The colonies seem to be quite small. Digging to 50 cm deep failed
to produce any brood chamber or any sexual forms. At about 10 to 20 cm
deep there was always a chamber full with small grass seeds and never
more than four majors. The major has a large head which is dispropor-
tional considering the small size of the rest of the body. Majors were
never observed outside the nest foraging. The small black, shining
workers can easily be confounded with P. metallescens and P. adrianoi
n. sp. workers.

58
14. Pheidole megacephala (Fabricius)
Formica megacephala Fabricius, 1793. Ent. Syst. 2:361.
_P. megacephala Emery 1921. Gen. Insect. Fase. 174, p. 85. Phillips
1934. (Hawaii Univ.) Expt. Sta. Pineapple Prod. Coop. Assoc. Bui. 15:5-12. -
Smith, 1936. Puerto Rico Univ., Jour. Agr. 20:843-844. Broekhuysen, 1948.
Union So. Africa Dept. Agr. Bui. 266:1-40. Smith, 1951. Cat. Hym. Am.
No. of Mex. U.S.D.A. Mon 2:803. Wheeler and Wheeler, 1953. Ent. Soc.
Wash., Proc. 55:75. Brown, 1958. Act. Hym. 1:47. Gregg, 1958. N.Y.
Ent. Soc. 66:23. Brown, 1959. Bui. Ent. Res. 50:523. Weber, 1960. Ent.
Soc. Wash., Proc. 62:232. Taylor and Wilson, 1961. Psyche 68:143. Kempf,
1962.Studia Ent. 5:18-19. Haskins and Haskins, 1965. Ecology 46:737. -
Wilson and Taylor, 1967. Pacific. Insects Monog. 14:46-48. Fluker, Huddle
ston, and Beardsley, 1968. Jour. Econ. Ent. 61:474. Fluker and Beardsley,
1970. Ent. Soc. Amer., Ann. 63:1290-1296. Kempf, 1972. Studia Ent. 15:196.
Oecophthora pusilla Heer, 1852. Stueck 66:15. Wheeler, 1922. Bull.
Amer. Mus. Nat. Hist. 45:812.
_P. laevigata Mayr, 1862. Verli. Zool. Bot. Ges. Wien 12:747. -
Mayr, 1870. Ve.hr. Zool. Bot. Ges. Wien 20:981.
Type locality: Isle de France (Mauritius)
Types : None in this country
Range : This species has become a world tramp; in Florida it
apparently is largely confined to South Florida.
Diagnosis:
P- megacephala (fig. 9), is a dark yellow to brownish-ferruginous
species that differs from P. morrisi by having distinct and sharp propodeal
spines. It also differs from P^. morrisi and _P. dentata by having a cordate

59
head narrowing toward the mandibular insertion and the mesonotum is not
distinctly transversely impressed.
Discussion:
P. megacephala is a well known world tramp (Gregg, 1958). It has
been present in Florida for 43 years, at least, being reported by
Smith (1933) from Everglades, Key West and St. Augustine. I collected
this species in downtown Homestead, Florida on November 25, 1975 nesting
under a paved sidewalk and at Archbold Biological Research Station.
Burn observed large colonies of this species trailing in large numbers
from the colony to nearby trees (Personal Communication). Its ecological
impact has been studied in relation to another world tramp, Iridomyrmex
humilis. Flucker et al^. (1970) reported I?. megacephala was losing
territory to I. humilis in Hawaii, although it did not appear to be
evenly remotely approaching extermination. They also found that _?.
megacephala did not seem to occupy territory above 9.15 m, above which I.
humilis was usually found. Dieberburg _et ajL. (1975) reported that in a
7-year period P_. megacephala did not loose much territory to _I. humilis
in Bermuda. He also reported that .P. megacephala prefers clumped food
sources and 1^. humilis prefered dispersed food sources. Based on these
findings they suggested that a equilibrium between the two species was
possible in Bermuda. Brown (1973) suggested that P. megacephala was a
mortal enemy of I. humilis and that, the latter was vanquishing JP.
megacephala outside the tropics.
In Florida the two species seem to occupy two distinct areas. 1.
humilis, although not common, occurs in the northernmost and northwest

60
areas of the state. P_. megacephala, also not common, occurs in the southern
area of the state. In the central area of the state I was able to find
only P. megacephala. At least in Florida the two species are not sympatric
and the inability of either of them to increase must be due to some other
cause, possibly competition with Solenopsis invicta Burn or Solenopsis
geminat.a (Fabricius).

61
15. Pheidole metallescens Emery
P. metallescens Emery, 1895. Zool. Jahrb., Abst. of System. 8:289,
294. Wheeler, 1908. Amer. Mus. Nat. Hist., Bui. 24:476-477. Smith,
1930. Fla. Ent. 14:3. Wheeler, 1932. N.Y. Ent. Soc. 40:7. Creighton,
S
19/0. Bull. Mus. Comp. Zool. 104:183. Smith, 1951. Cat. Hym. Am. No. of
Mex., U.S.D.A. Mon 2:803. Gregg, 1958. N.Y. Ent. Soc. 66:23. Wheeler
and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:12.
Type locality: St. George, Florida
Types : None in this country
Range : Gulf States
Diagnosis:
.P. metallescens (fig. 16) is a small species usually with reddish
brown head and thorax and black gaster, although completely dark to black
specimens have been found. It differs from _P. greggi n. sp. and P_. moerens
by the distinct lateral connules on the postpetiole. It also differs from
P_. floridana and IP. anastasii by the strong reticulation posteriorly on
the head and from P_. dentigula by having a reticulo-punctate propodeal
base. The workers display strong, metallic, iridescent bluish reflections
over most of the body.
Discussion:
P. metallescens is very common in Florida. The preferable habitat
seems to be under the shade of trees or any high vegetation. Usually it
is easy to find several small craters connected to a central vertical tunnel
to the main chamber, rarely over 40 cm deep in the ground. It is a
monogynous species. The majors usually forage along with the workers.
They feed on small grass seeds and also scavenge dead arthropods.

62
16. Pheidole moerens Wheeler
P. moerens Wheeler, 1908. Bull. Amer. Mus. Nat. Hist. 24:136. -
Smith, 1936. Puerto Rico Univ., Jour. Agr. 20:842-843. Smith, 1967.
Cat. Hym. Amer. No. of Mex. U.S.D.A. Mon 2 (Suppl. 2):354. Wheeler
and Wheeler, 1972. Ga. Ent. Soc. Jour. 7:244. Kempf, 1972. Studia
Ent. 15:196. Wojcik, Banks, and Burn, 1975. Coop. Econ. Ins. Rpt.
U.S.D.A. 25449: 906.
P. moerens dominicensis Wheeler, 1913. Bull. Amer. Mus. Nat. Hist.
32:241. Kempf, 1972. Studia Ent. 15:197. New Synonymy.
Type locality: Antilha Culebra
Types : M.C.Z.
Range : Southern Alabama, Florida, West Indies
Diagnosis:
_P. moerens (fig. 22) is a small reddish yellow to almost black species
closely related to P. greggi n. sp., differing from it by the absence of
the heavily punctate scrobe and in having a carinated shining clypeus. It
differs from P_. dentigula, P_. anastasii, P_. floridana and P. metallescens
by the absence of distinct lateral connules on the postpetiole.
Discussion:
_P. moerens is a common species in north Florida. Its color was
observed in lab colonies to vary from reddish yellow to almost black due
possibly to variation in the diet. Colonies fed with honey and seeds
retained a lighter color than those fed with freshly killed house flies.
Wheeler's subspecies dominicensis appears to represent only a color
variant often found among the normal population. I have examined

63
the types of dominicensis and they appear identical with the types of
moerens except in color. I do not consider dominicensis a valid taxon.
This species was found nesting in various places such as under boards,
at base of oak trees and fence posts, along roots, under palm leaves,
inside wall crevices, and rarely in the ground. Usually small chambers
are constructed. It is a monogynous species which has a small blackish
female. Its main flight is usually in July. Several queens may start
founding a nest, but before the first brood emerges, the dominant female
will have killed the others.
The chambers usually are built with small soil or debris particles
and have small openings. A colony may have over 100 majors and over 300
workers. They feed on seeds and scavenge and prey on small dead or live
arthropods. They forage very close to the nesting sites and sometimes a
major is found foraging along with the workers.

64
17. Pheidole morrisi Forel
P. morrisii Forel,
1886.
Soc.
Ent.
Belg.
, Bui.
30:46. -
Mayr,
1887.
Verh.
Zool.
- Bot. Ges.
Wien 37:568
. -
Forel,
1901.
Soc. Ent
. Belg.
* >
Ann.
45:350.
- Wheeler,
1904.
Amer.
Mus
. Nat.
Hist.
, Bui 20:
302. -
Wheeler,
1905. Amer. Mus. Nat. Hist., Bui. 21:380. Wheeler, 1908. Amer. Mus.
Nat. Hist., Bui. 24:461. Smith, 1918. Ent. News 29:21. Davis and Bequaert,
1922. Brooklyn Ent. Soc. Bui. 17:8-9. Smith, 1924. Ent. News 35:53. -
Smith, 1930. Fla. Ent. 14:3. Wheeler, 1932. N.Y. Ent. Soc. 40:7. -
Cole, 1940. Amer. Midland Nat. 24:44-45. Creighton, 1950. Bull. Mus.
Comp. Zool. 104:183-184. Smith, 1951. Cat. Hym. Amer. No. of Mex.,
U.S.D.A. Mon 2:803. Gregg, 1958. N.Y. Ent. Soc. 66:24. Wheeler and
Wheeler, 1960. Ent. Soc. Amer., Ann. 53:12.
P. morrisi var. vancea Forel, 1901. Soc. Ent. Belg., Ann. 45:351. -
Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 20:302. Wheeler, 1908. Amer.
Mus. Nat. Hist., Bui. 24:461. Smith, 1924. Ent. News 35:53. Smith,
1951. Cat. Hym. Amer. No. of Mex., U.S.D.A. Mon 2:803-804.
Type locality: Vineland, New Jersey
Types : None in this country
Range : Southern New Jersey, south to Florida and eastern
Gulf States.
Diagnosis:
jP morrisi (fig. 27) is a pale to darker yellowish species differing
from P. dentata and JP. vinelandica by the absence of distinct propodeal
teeth or spines. It is larger than P. vinelandica and has a transverse
mesonotal impression.

65
Discussion:
_P. morrisi is another common species in Florida. It always nests in
the ground and the colonies are quite large. It is a monogynous species
and is primarily a scavenger.
Several nest founding females displayed singular habits in the
laboratory which may not have been noted previously in this genus. In
constructing the claustral chamber each female used the soil dug from
the bottom of the vertical passageway to form a continuous plug of the
passageway above her rather than bringing the soil particles to the
surface. The claustral chamber at the 20 to 30 cm level below ground thus
is entirely sealed from the surface by a long plug. I interpret this
habit as a defense mechanism against various predators during the claustral
period. The first workers are reared in about 30 days and the first
majors in about 50 days.
The colony increases in size rapidly. In 8 months it can have several
hundred workers and majors. The workers forage alone, although the majors
can help to bring food back to the colony once food is located. Workers
can forage over 8 m from the nesting site. The species is mainly a
scavenger but will also gather seeds.

66
18. Pheidole quiescens n. sp.
Diagnosis:
A small yellowish brown species (fig. 46, 47) parasitic in nests
of P. anastasii. Queens, soldiers and males are still unknown. Soldier
caste probably lacking as in other species parasitic on Pheidole.
Obviously closely related to its host and to other small species in this
species group (P. floridana, P. moerens, and _P. greggi n. sp.) but dis
tinctive in its very long and delicate pilosity and its large and singular
ly shaped postpetiole which seems broadly and almost immovable articulated
with the gaster. In behavior the species seems unique for its habit of
standing motionless with open mandibles for long periods at a distance
from the host workers and their activities.
Description:
Maj or:
Unknown. Probably does not exist in this species.
Worker:
Measurements: Body length 1.90 to 2.08 mm. Head length 0.46
to 0.49 mm; head width 0.41 to 0.46 mm. Thoracic length 0.56 to
0.62 mm. Scape length 0.38 to 0.46 mm. Scape Index 89 to 96.
Structural characters: Head, excluding mandibles, slightly
longer than broad, posteriorly with a median sulcus and weak impression
which forms weak occipital lobes. Frontal area small, slightly depress
ed, with two longitudinal striae. Clypeus with a fine median stria-
tion; anterior margin curved and slightly notched in the middle.
Frontal carinae short and divergent; antenae 12 segmented and with a
3 jointed club as in nonparasitic species; scapes long and slender,

67
the posterior third thicker, usually passing beyond the posterior
margin of the head by an amount less than the length of first funicular
segment. Mandibles slender, curved apically and with two large, sharp
teeth, the remaining border with smaller sharp teeth; eyes convex,
measuring less than 1/4 the head length and with approximately 22
facets. In profile,pronotum and anterior mesonotum convex; posterior
of mesonotum more abruptly descending to the wide mesopropodeal
impression which is slightly depressed below the anterior propodeal
margin. Juncture of base and declivity of propodeum rounded; pro
podeal spines broad and heavy at base and pointed obliquely laterad.
Petiole with gently sloping anterior dorsal face, the posterior face
vertical, the node in profile very blunt apically.
Postpetiole, seen from above, slightly wider than long, two
times or more wider than petiole, the sides slightly angular but not
forming distinct conules, broadest in the posterior half, gaster
truncate at base and always noticeably larger than the head.
Sculpture: Head including irons and clypeus opaque and finely
reticulo-punctate; mandibles smooth and shining. Frontal area small
and finely reticulo-punctate. Sides of thorax and petiole, sub
opaque and reticulo-punctate. Postpetiole weakly reticulo-punctate
and weakly shining. First gastric tergite and usually the postpetiolar
dorsum with opalescent reflections and subopaque.
Pilosity: Hairs yellow, long, very slender and pointed, some
measuring up to 0.15 mm, numerous all over the body, even the ventral
surface of the head; short and sparse hairs on the legs; pubescence
limited to the antennae, absent on gaster and other regions of body.

68
Color: Light reddish brown, appendages lighter.
Female and Male:
Unknown
Holotype: 1 worker, collected at Archbold Research Station.
Paratypes: 3 workers collected in two colonies of the host
Pheidole anastasii at the Archbold Biological Research Station in
Lake Placid, Florida, by M. A. Naves in August, 1975.
The holotype will be deposited in the Museum of Comparative
Zoology, Harvard University, Cambridge, Massachussetts. Paratypes
will also be deposited in the National Museum of Natural History,
Washington, D.C., the Florida State Museum, Department of Plant
Industry, Gainesville, Florida, and M. A. Naves Collection.
Discussion:
Wheeler (1904) described two parasitic species, Epipheidole inquilina
and Sympheidole elecebra. EL inguilina was parasitising colonies of P.
pilifera coloradensis and S^. elecebra was parasitising colonies of P_. ceres.
Both parasitic species were collected in Colorado and up to now have been
the only known ant parasites for Pheidole in the U.S.
E. inquilina was described from males and females collected in three
colonies of its host. The lack of workers and absence of the host queen
led Wheeler to the assumption that the host queen is dispatched by its
own workers once the parasite had secured entry into the nest, and that
this species deserved placement under a separate genus. Later Dr. M. R.
Smith (1941) announced the discovery of a single worker of inquilina^
possibly jeopardizing the status of Epipheidole as separate from Pheidole.

69
S. elecebra was described from one queen and 18 males. Later another
queen was found. The worker caste, if any exists, is still unknown.
Brown (1973) listed both Epipheidole and Sympheidole as synonymous
with Pheidole, but these listings were not annotated and the validity
of these synonymizations is in doubt.
P. quiescens, in contrast with E. inquilina and S. elecebra, still
produces its worker caste. Several attempts to collect sexual forms of
_P. quiescens have failed. In two cases parasitised colonies still retain
ed the ho§t queen.
One of these colonies was brought to the lab for observation. It
was noted that the parasite workers essentially stand in the colony doing
nothing for long periods of time. They obtain their food by trophallaxis
from the host workers. They always stand with the mandibles open, possibly
in a defensive position, at a distance from the host workers. If the
colony happens to be disturbed the parasite will run away, apparently
seeking refuge. Twice they were observed helping to move brood from one
place to another but never were observed doing any other useful work. The
interspecific relationship seems to be inquilinism which means that the
parasite spends its whole life cycle in the host colony.
Subsequent collections of host workers and parasite workers were added
to the lab colony and no rejection of the newcomers was ever observed.
Later on, the host queen died and another queen of P. anastasii collected
in Gainesville, Florida was readily accepted by the colony. The host is
very well distributed all over the state, but collections throughout the
state failed to have parasites, although P. quiescens seems to be common
in colonies of its host at Archbold Biological Research Station.

70
Permanent social parasitsm with a useless worker caste is already
known to occur among ants. Burn (1942), when describing Formica reflexa
suggested that the consistant presence of workers of both _F. subsericea,
the host, and of small, timid workers of F. reflexa in every colony that
he found, was possibly a case of F. reflexa being a permanent inquiline.
King (1949) presented further evidence confirming Buren's suggestions.
King and Sallee (1951) suggested that the absence of pure colonies of F.
reflexa confirmed Buren's suggestions.
_P. guiescens seems to fit Buren's and King's descriptions of a
permanent social parasite which has retained a useless worker caste. P.
quies.cens was collected several times and always the host-workers out-
numered the parasite at least several to one. The new species was never
found as a pure colony.

71
19. Pheidole tysoni Forel
_P. tysoni Forel, 1901. Soc. Ent. Belg., Ann. 45:348. Wheeler,
1904.
Amer.
Mus.
Nat. Hist. 20:302. Smith,
1918.
Ent.
News 29:
22. -
Smith,
1924.
Ent
. News 35:78. Dennis, 1938.
Ent.
Soc.
Amer. 31
: 282,
304. -
Wesson
and
Wesson, 1940. Amer. Midland
Nat.
24:92
. Cole,
1940
Amer. Midland Nat. 24:42, 45. Creighton, 1950. Bull. Mus Comp. Zool.
104:191. Smith, 1951. Cat. Hym. Amer. No. of Mex., US.D.A. Mon 2:805. -
Gregg, 1958. N.Y. Ent. Soc. 66:32.
Type locality: Mt. Mitchell, North Carolina
Types : A. M. N. H., M. C. Z.
Range : North Florida, Georgia, western North Carolina,
southwestern Virginia and eastern Tennessee.
Diagnosis:
P_. tysoni is a small yellowish species closely related to P. vinelandica,
differing from it by the teeth on the anterior ventral margin of head. The
minor has the propodeal pleurae largely free from sculpture and strongly
shining.
Discussion:
_P. tysoni was collected once in a soybean field at Tall Timbers
Research Station in 1973. The colony was quite large and the main chamber
was about 50 cm deep in the ground. Only minors were foraging. This is
the only record known for the state and I am assuming that P. tysoni is
rare in Florida.

72
20. Pheidole vinelandica Forel
P. bicarinata race vinelandica Forel. 1886. Ann. Soc. Ent. Belg.
30:45.
vinelandica Mayr, 1886. Verh. Zool. Bot. Ges. Wien 36:458. -
Emery, 1895. Zool. Jahrb. Syst. 8:292. Forel, 1901. Am. Soc. Ent.
Belg. 45:348. Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 24:458. -
Smith, 1918. Ent. News 29:22. Davis and Beguaert, 1922. Brook. Ent.
Soc. 17:9. Smith, 1924. Ent. News, 35:54. Hayer, 1925. Ent. News
36:42 Cole, 1940. Amer. Midland Nat. 24:42. Wesson and Wesson, 1940
Amer. Midland Nat. 24:92-93.
P. (Allopheidole) vinelandica Forel, 1912 Mem. Soc. Ent. Belg.
19:237.
P. vinelandica laeviuscula Emery, 1895. Zool. Jahrb. Syst. 8:292.
Smith, 1951. Cat. Hym. Amer. No. of Mex. U.S.D.A. Man. 2:805.
P. vinelandica subsp. buccalis Wheeler, 1908. Bull. Amer. Mus.
Nat. Hist. 24:454. Smith, 1951. Cat. Hym. Amer. No. of Mex. U.S.D.A.
Mon 2:805.
_P. vinelandica longula var. castanea. Wheeler, 1915. Amer. Mus.
Nat. Hist. 34:405. Cole, 1937. Ent. News. 48:100.
JP. bicarimata buccalis Creighton, 1950. Bull. Mus. Comp. Zool.
104:171. Cole, 1953. Tenn. Ac. Sc. 28:297.
P. bicarinata vinelandica Creighton. 1950. Bull. Mus. Comp. Zool.
104:17a. Cole, 1956. Tenn. Acad. Sc. 31:114. Gregg, 1958. N.Y. Ent.
Soc. 66:18, 36. Reid and Nugara, 1961. Jour. Parasitology 47:885-889.
Smith, 1965. U.S. Dept. Agr., Tech. Bui. 1326:25-26.

73
P. vinelartdica vinelandica. Smith, 1951. Cat. Hym. Amer. No. of
Mex. U.S.D.A. Mon. 2:806.
Type locality: Vineland, New Jersey.
Types : Cotypes, A.M.N.H.
Range : Southeastern and Gulf States and along the east
coast to New Jersey
Diagnosis:
P_. vinelandica is a small yellowish species differing from P. bicarinata
by having the basal face of propodeum in the major largely punctate (fig.
0
m and having a longitudianl striation on the frontal area (fig. 66). It
differs from P. tysoni by the absence of teeth on the anterior ventral
margin of the head.
Discussion:
I had the opportunity to study specimens of P. vinelandica from New
Jersey, North Carolina, South Carolina, Georgia, Alabama, and Texas. I
have also collected this species at Tall Timbers Research Station as well
as in Putman Co., Florida. I have compared these with specimens of P_.
bicarinata (fig. 24, 64, 65) from Illinois in the M.C.Z. I also have
studied specimens of bicarinata collected at Colorado Springs, Colorado;
Ames and Lewis, Iowa; Devils Tower, Wyoming; Ortonville, Minnesota, and
Corinne, Utah. It was evident that the two forms are not conspecific
and have constant morphological differences, emphasizing that P. vinelandica
deserves full species rank. Gregg (1958) included the var. nebrascensis
Forel and the var. huachucana Smith as synonymus of vinelandica. I have
not studied the types of these taxa but am doubtful about their inclusion
under vinelandica rather than bicarinata. Records of vinelandica from

74
North Dakota and Colorado also seem doubtful, and seem much more likely to
be misdetermined records of bicarinata.
P_. vinelandica is found nesting in grassy areas at Tall Timbers
Research Station. The minors and majors are easily seen foraging close to
the nesting site. The nest has several exits. They were observed carrying
seeds and small dead arthropods.

75
SPECIES NOT PROPERLY INCLUDED IN THE FLORIDA PHEIDOLE FAUNA
1. Pheidole pilifera (Roger)
Leptothorax pilifera Roger, 1863. Berl. Ent. Zeitschr., 7:180.
_P. pilifera var. simulans Wheeler, 1908. Bull. Amer. Mus. Nat. Hist.,
24:436.
P_. pilifera subsp. septentrionalis Wheeler, 1908. Bull. Amer. Mus.
Nat. Hist. 24:436.
P. pennsylvanica Roger, 1863. Berl. Ent. Zeitschr., 7:199. Mayr,
1870. Verh. Zool-Bot. Ges. Wien, 20:981. Mayr, 1886. Verb. Zool-Bot.
Ges. Wien, 36:455. Mayr, 1877. Verh. Zool-Bot. Ges. Wien, 37:588.
P^. pilifera Emery, 1895. Zool. Jahrb. Syst. 8:290. Creighton, 1950.
Bull. Mus. Corap. Z-ol. 104:184-186. Van Pelt, 1956. Amer. Midland Nat.
56:377, 834. Smith, 1958. Cat. Hym. Amer. No. of Mex., U.S.D.A. Mon.
2 (Suppl. 1):123. Gregg, 1958. N.Y. Ent. Soc., 66:24-25.
Type locality: Pennsylvania
Types : None in this country
Range : Massachussetts to North Carolina and West to Iowa
and Nebraska
Diagnosis:
P_. pilifera differs from P. carrolli n. sp. and _P. littoralis Cole
by having the occipital rugae of the major notably reticulate and often
coarse; its occipital lobes are not well rounded and pronounced.
Discussion:
Attempts were made by the author to find P. pilifera in Florida as
reported by Van Pelt (1956). Dr. Van Pelt, however, expressed some reserva
tions about the correctness of this identification. No voucher specimens

76
of P. pilifera are presently available from Dr. Van Pelts work. Although
it seems possible that P. pilifera occurs in Florida, I am unconvinced
that there is any direct evidence. I believe all reported specimens were
probably carrolli n. sp., P. littoralis Cole, or other species.

PLATE 1
Figure
1.
Head of major of P.
lamia Wheeler (33X)
Figure
2.
Scape of major of P
of the crassicornis
. bureni n. sp., a member
complex (81X)
Figure
3.
Head of major of P.
diversipilosa Wheeler (45X)
Figure
4.
Gaster of major of
P. diversipilosa (47X)
Figure
5.
Head of major of P.
crassicornis Emery (43X)
Figure
6.
Gaster of major of
P. crassicornis (58X)

78

PLATE 2
Figure 7.
Figure 8.
Figure 9.
Figure 10
Figure 11
Scape of major of P_. fallax obscurithorax
Santschi (81X)
Head of major of P. fallax obscurithorax (40X)
Head of major of P. megacephala (Fabricius)
(40X)
Pedicel of major of P_. floridana Emery (128X)
Propodeum, pedicel and gaster of major of P.
metallescens Emery (74X)
Figure 12. Postpetiole of major of P. littoralis Cole (119X)

80

PLATE 3
Figure 13.
Figure 14.
Figure 15.
Figure 16.
Figure 17.
Head of major of P. dentigula M. R. Smith
(64X)
Head of major of P. carrolli n. sp. (29X)
Pedicel of major of P. metallescens Emery
(163X)
Head of major of _P. metallescens (62X)
Head of major of P_. anastasii Emery (68X)
Figure 18.
Postpetiole and gaster of P. anastasii (62X)

82

PLATE 4
Figure
19.
Head
of
major
of
p.
floridana Emery
(7 2X)
Figure
20.
Head
of
major
of
p.
littoralis Cole
(37X)
Figure
21.
Head
of
major
of
p.
greggi n. sp. (68X)
Figure
22.
Head
of
major
of
p.
moerens Wheeler
(58X)
Figure
23.
Clypeus and frontal area of major of
vinelandica Forel (186X)
P.
Figure
24.
Head
of
maj or
of
p.
bicarinata Mayr
(53X)


PLATE 5
Figure 25. Head of major of P_. adrianoi n. sp. (72X)
Figure 26. Thorax, pedicel and gaster of major of P_.
adrianoi n. sp. (50X); white legs are
artifacts due to electron overcharge.
Figure 27. Head of major of P_. morrisi Forel (37X)
Figure 28. Thorax of major of P. morrisi (56X)
Figure 29. Head of major of P_. dentata Mayr (48X)
Figure 30. Teeth on the anterior ventral head margin
P. anastasii Emery (123X)


PLATE 6
Figure 31. Head of worker of ]?. lamia (97X)
Figure 32. Dorsal view of worker of P. vinelandica
(62X)
Figure 33. Dorsal view of worker of P. adrianoi n. sp.
(95X)
Figure 34. Worker of P^. megacephala (36X)
Figure 35. Thorax and pedicel of the worker of _P. fallax
obscurithorax (41X)
Head of worker of P. fallax obscurithorax
(86X)
Figure 36.

88

PLATE 7
Figure
37.
Thorax of the worker of
Figure
38.
Head, thorax and pedicel
P. dentata (72X)
Figure
39.
Thorax of the worker of
Figure
40.
Head of the worker of P.
Figure
41.
Thorax of the worker of
(106X)
Figure
42.
Head of the worker of P.
. morrisi (72X)
of the worker of
dentata (79X)
crassicornis (54X)
1. metallescens
littoralis (113X)

90

PLATE 8
Figure 43.
Figure 44.
Figure 45.
Figure 46.
Figure 47.
Figure 48.
Worker of 1?. carrolli n. sp. (40X)
Head of the worker of P. moerefrfe (102X)
Head of worker of P. greggi n. sp. (102X)
Worker of _P. quiescens n. sp. (45X)
Dorsal view of worker of P. quiescens n. sp.
(103X)
Propodeum, pedicel and gaster of worker of
£. dentigula (102X)

92

PLATE 9
Figure 49. Dorsal view of thorax of ]?. dentigula worker
(115X)
Figure 50. Head of worker of P. anastasii (109X)
Figure 51. Head of worker of P. floridana (115X)
Figure 52. Pedicel of worker of _P. floridana (193X)
Figure 53. Dorsal thoracic hairs of worker of P_. adrianoi
n. sp. (1,401X and 5,174X, respectively)
Dorsal thoracic hairs of worker of P^. adrianoi
n. sp. (1,401X and 5,174X, respectively)
Figure 54.

94

PLATE 10
Figure 55.
Figure 56.
Figure 57.
Figure 58.
Head of major of P_. flavens (95X)
Head of major of P_. flavens sculptior (87X)
Occipital lobes of P_. greggi n. sp. (250X0
Postpetiole of the major of P. flavens sculptior
(158X)
Figure 59.
Head and thorax of the minor of P. flavens (83X)

96

PLATE 11
Figure 60. Head of major of P. bureni n. sp.
Figure 61. Gaster and pedicel of £. bureni n
Figure 62. Head and pronotum of worker of P.
(52X)
(39X)
. sp. (40X)
bureni n.

98

PLATE 12
Figure 63. Dorsum of propodeum of major of P. vinelandica
(272X)
Figure 64. Thoracic dorsum of P_. bicarinata (71X)
Figure 65. Postpetiole of P. bicarinata (186X)
Figure 66.
Head of P. vinelandica (66X)

100

PLATE 13
Figure 67. Maps of Florida and the United States
showing:
Tall Timbers Research Station
O Gainesville
T Archbold Biological Research Station
Locations where collections of Pheidole
spp. were made in Florida
V -
Alice, Texas

102
?*T
J /
f NiSi'Aj'"'--
i j L t-L-. \ > }-^srV

LITERATURE CITED
Brown, W. L. 1973. A comparison of the Hylean and Congo-West African
rain forest ant faunas, p. 161-185. In: B. J. Meggers, E. S.
Ayensu, and W. D. Duckworth (ed.). Tropical forest ecosystems in
Africa and South America:A comparative review. Smithsonian Institution
Press, Washington, D.C.
Burn, W. F. 1942. New ants from Minnesota, Iowa, and Wisonsin. Iowa
State College Jour. Sci. 16:399-40.
Burn, W. F. 1944. A List of Iowa Ants. Iowa State College Jour. Sci.
18:277-312.
Burn, W. F., M. A. Naves, and T. Carlysle. False Phragmosis and Apparent
Specialization for Subterranean Warfare in Pheidole lamia Wheeler
(Hymenoptera:Formicidae) In press.
Cole, A. C. 1952. A New Pheidole (Hymenoptera:Formicidae) From Florida.
Ent. Soc. Amer., Ann. 45:443-444.
Creighton, W. S. 1950. The ants of North America. Bull of the Mus. Comp.
Zool, Harvard, 104:1-585.
Emery, C., 1895. Beitraege Zur Kenntnis der nordamerikanischen Ameisenfauna.
II. Zool. Jahrb. Syst. 8:257-360.
Fluker, S. S., and J. W. Beardsley, 1970. Sympatric Associations of Three
Ants:Iridomyrmex humilis, Pheidole megacephala, and Anoplolepis
longipes in Hawaii. Ent. Soc. Amer., Ann. 63:1290-1296.
Gregg, R. E. 1958. Key to The Species of Pheidole (Hymenoptera:Formicidae)
in the United States. N.Y. Ent. Soc., Jour. 66:7-48.
Gregg, R. E. 1969. New Species of Pheidole from Pacific Coast Islands
(Hymenoptera¡Formicidae) Entom. News 80:93-101.
Kempf, W. W. 1972. Catalogo abreviado das Formigas da Regiao Neotropical
(Hymenoptera:Formicidae). Studia Ent. 15:3-344.
King, R. L. 1949. Mixed Colonies in Ants. Iowa Acad. Sci. 56:367-370.
King, R. L., and R. M. Sallee. 1951. More Mixed Colonies in Ants. Iowa
Acad. Sci. 58:487-489.
Liebergurg, I., P. M. Krans, and A. Seip. 1975. Bermudian Ants revisited.
The Status and Interaction of Pheidole megacephala and Iridomyrmex
humilis. Ecology. 56:473-478.
103

104
Mayr, G. L. 1886. Die Formicidin Der Vereinigten Staaten Von Nordamericana
Verhandlungen Der. Zool-Bot. Gesell. Wein, Verh. 36:419-464.
Smith, M. R. 1930. A List of Florida Ants. Fla. Ent. 14:1-6.
Smith, M. R. 1933. Additional species of Florida ants, with remarks.
Fla. Ent. 17:21-26.
Smith, M. R. 1941. The discovery of the worker caste of an inquilinous
ant, Epipheidole inquilina Wheeler. Proc. Ent. Soc. Washington
24:104-109.
Smith, M. R. 1944. Additional ants recorded from Florida, with descriptions
of two new subspecies. Fla. Ent. 27:14-17.
Smith, M. R. 1951. Formicidae. In: K. V. Krombein and B. D. Burks,
eds., Hymenoptera of America North of Mexico, Synoptic Catalog, U.S.D.A.,
Agr. Mon. 2:799-806.
Smith, M. R. 1958. Formicidae. In: K. V. Krombein and B. D. Burks,
eds., Hymenoptera of America North of Mexico, Synoptic Catalog,
U.S.D.A., Agr. Mon 2 (Suppl. 1):120.
Smith, M. R. 1967. Formicidae. In: K. V. Krombein and B. D. Burks, eds.,
Hymenoptera of America North of Mexico, Synoptic Catalog, U.S.D.A.,
Agr. Mon 2 (Suppl. 2):343-474.
Van Pelt, A. F., Jr. 1956. The Ecology of the ants of the Welaka Reserve,
Florida (Hymenoptera:Formic.idae). Amer. Midland Nat. 56:387.
Wheeler, W. M. 1904. Three new genera of Inquiline ants from Utah and
Colorado, Bull. Amer. Mus. Nat. Hist. 20:1-17.
Wheeler, WT. M. 1908. The ants of Texas, New Mexico and Arizona. Bull.
Amer. Mus. of Nat. Hist. 24:399-485.
Wheeler, W. M. 1910. Ants, their structure, development and behavior,
Columbia University Press, 663 pp.
Wheeler, W. M. 1932. A list of the ants of Florida with descriptions of
nev; forms. N.Y. Ent. Soc. Jour. 40:1-171.
Wilson, E.0. 1975. Enemy Specification in the Alarm-Recruitment System
of the Ant. Science 190:798-800.
Wojcik, D. P., W. A. Banks and W. F. Burn, 1975. First Report of Pheidole
moerens in Florida (Hymenoptera:Formicidae) U.S.D.A. Coop. Econ.
Ins. Rpt. 25(49-52):906.

BIOGRAPHICAL SKETCH
Marcio Antonio Naves was born February 26, 1943, in Lavras, Minas
Gerais, Brazil. He is married to Lucilia de Rezende Naves, and has a
son, Adriano de Rezende Naves.
In December, 1970, he graduated from Escola Superior de Agricultura
de Lavras, Lavras, M. G., where he received a BS in Agronomy. In 1971
he was hired as coordinator of the Agreement ESAL/DNOS for the irrigation
project of the valleys of Careacu and Bela Vista, M. G. In June, 1974,
he received the degree of Master of Science from the Entomology and
Nematology Department of the University of Florida, Gainesville, Florida.
He has been a member of the American Entomological Society and Florida
Entomological Society, and Georgia Entomological Society.
In 1974, he began to work towards the Degree of Doctor of Philosophy
with a major in Entomology and minor in Plant Pathology.
105

I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality as a dissertation for the degree of
Doctor of Philosophy.
William F. Burn, Chairman
Professor of Entomology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality as a dissertation for the degree of
Doctor of Philosophy.
George E. Allen
Professor of Entomology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality as a dissertation for the degree of
Doctor of Philosophy.
Professor of Plant Pathology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality as a dissertation for the degree of
Doctor of Philosophy.
x// >y n Q /
TZr/r: -4.1
Lofgren
Professor of Entomology

This dissertation was submitted to the Graduate Faculty of the College
of Agriculture and to the Graduate Council, and was accepted as partial
fulfillment of the requirements for the degree of Doctor of Philosophy.
December, 1976
Dean, Graduate School



36
In profile, pro-mesonotum moderately convex; propodeal spines
small, sharp, and projecting obliquely dorsad; petiole as in the
major, except the crest is proportionately much broader; postpetiole
convex dorsally and flat ventrally, seen from above the sides are
strongly convex and almost 2 times wider than petiole; gaster oval.
Sculpture: Head reticulo-punctate and opaque; a few rugules
on the genae, between the eyes and on the mandibles; thorax, in
profile, reticulo-punctate; dorsum of pronotum and mesonotum usually
with a few longitudinal or transverse striae; petiole reticulo-
punctate; sides of postpetiole weakly punctate, dorsum weakly shin
ing; gaster smooth and shining.
Pilosity: Erect hair yellow, short on the dorsal surface of
head, longer on the ventral surface and anterior clypeal margin;
long and sparse on thorax and pedicel; gaster with rather sparse
hair of mixed lengths; pubescence confined to the antennae and legs.
Color: Reddish brown, sometimes the mandibles and legs are
lighter.
Holotype: Major, collected near Gainesville, Florida by M. A.
Naves.
Paratype: 3 majors and 20 workers probably from the same nest
as the holotype, collected as follows:
Dr. John F. Carroll collected one major and several workers on
August 2, 1973, near Gainesville, Florida. M. A. Naves collected
three majors and several workers in August, 1975, at the same locality.


65
Discussion:
_P. morrisi is another common species in Florida. It always nests in
the ground and the colonies are quite large. It is a monogynous species
and is primarily a scavenger.
Several nest founding females displayed singular habits in the
laboratory which may not have been noted previously in this genus. In
constructing the claustral chamber each female used the soil dug from
the bottom of the vertical passageway to form a continuous plug of the
passageway above her rather than bringing the soil particles to the
surface. The claustral chamber at the 20 to 30 cm level below ground thus
is entirely sealed from the surface by a long plug. I interpret this
habit as a defense mechanism against various predators during the claustral
period. The first workers are reared in about 30 days and the first
majors in about 50 days.
The colony increases in size rapidly. In 8 months it can have several
hundred workers and majors. The workers forage alone, although the majors
can help to bring food back to the colony once food is located. Workers
can forage over 8 m from the nesting site. The species is mainly a
scavenger but will also gather seeds.


42
as the normal caste. This is a very interesting species for ecological
and laboratory studies. Recent studies by Wilson (1975) have shown that
workers of P. dentata use odor trails to recruit nestmates to food
discoveries and new nest sites. The same pheromone is used also to
recruit major workers to the vicinity of intruders. This type of alarm
recruitment proved rather narrowly specific and works best against some
of the potentially more important enemies of P. dentata in the genus
Solenopsis.


Abstract of Dissertation Presented to the Graduate Council
of the University of Florida in Partial Fulfillment of the Requirements
for the Degree of Doctor of Philosophy
A MONOGRAPH OF THE GENUS PHSIPPLE IN FLORIDA
(HYMENOPTERA:FORMICIDAE)
By
Marcio A. Naves
December, 1976
Chairman: William F. Burn
Major Department: Entomology and Nematology
Until recently only twelve species of Pheidole were reported from
Florida, P. anastasia Emery, P_. bicarinata vinelandica For el, P. dentigula
M. R. Smith, ?. f lavens Roger, P. megacephala (Fabricius), P_. moerens
Wheeler, morrisl Forel, P_. pilifera (Roger), P. dentata Mayr, P.
floridana Emery, P. metallescens Emery and P_. sitarches littoralis Cole.
The last four have Florida as the type locality.
£. pilif era (Roger) is believed to have beer, errouneously reported
from Florida and has been excluded from the Florida fauna. P. flavens
(Gregg, 1958, nec Roger) is considered a misidentification and to be an
undescribed species, P. greggi n. sp.
In addition to P. greggi n. sp. three additional new species of
Pheidole are described from Florida: P_. adrianoi n. sp., P. carrolli n.
sp. and £. qu.iesc.ens n. sp. An exotic species, P. fallax obscurithorax
(= £. fallax ajencola var. obscurithorax Santschi) probably introduced
from South America, is also added to the North American ant fauna.
Pheidole crassicornis Emery and ?. diversipllosa Wheeler are record
ed for the first time in Florida. The latter species is resurrected
from sycornymy and is elevated to full species.


I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality as a dissertation for the degree of
Doctor of Philosophy.
William F. Burn, Chairman
Professor of Entomology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality as a dissertation for the degree of
Doctor of Philosophy.
George E. Allen
Professor of Entomology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality as a dissertation for the degree of
Doctor of Philosophy.
Professor of Plant Pathology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate, in scope and quality as a dissertation for the degree of
Doctor of Philosophy.
x// >y n Q /
TZr/r: -4.1
Lofgren
Professor of Entomology


PLATE 6
Figure 31. Head of worker of ]?. lamia (97X)
Figure 32. Dorsal view of worker of P. vinelandica
(62X)
Figure 33. Dorsal view of worker of P. adrianoi n. sp.
(95X)
Figure 34. Worker of P^. megacephala (36X)
Figure 35. Thorax and pedicel of the worker of _P. fallax
obscurithorax (41X)
Head of worker of P. fallax obscurithorax
(86X)
Figure 36.


31
rugae; frontal carinae short and diverging posteriorly; ventral
surface of the head smooth and shining, its anterior margin with
three distinct teeth, the median tooth short and lobe-like and the
lateral ones long and sharp; sides of pronoturn opaque; esisternum
smooth and shining; propodeum with transverse striae, the inter
stitial areas weakly punctate and shining; propodeal base and
declivity smooth and shining; sides of pedicel finely reticulo-
punctate; gaster smooth and shining.
Pilosity: Hairs yellow and pointed and measuring up to 0.40 mm
in length; head with numerous short and long erect and semi-erect
hairs; dorsum of thorax and petiole with numerous short and long
pointed hairs; postpetiole and gaster with numerous, predominantly
short semi-erect hairs; short sparse fine and pointed semi-erect
hairs on scapes and apendages.
Color: Light reddish brown
Male: Unknown
Holotype: Major, collected at Alice, Texas
Paratypes: 6 majors, 7 workers and 1 female, from the same
colony as the Holotype.
The type material comprises part of a colony collected by William
F. Burn on October 23, 1942 at Alice, Texas.
The holotype, the female paratype and one worker paratype will be
deposited in the Museum of Comparative ^oology, Harvard University,
Cambridge, Massachussetts. Paratypes will also be deposited in the
National Museum of Natural History, Washington, D.C., the Florida State


PLATE 9
Figure 49. Dorsal view of thorax of ]?. dentigula worker
(115X)
Figure 50. Head of worker of P. anastasii (109X)
Figure 51. Head of worker of P. floridana (115X)
Figure 52. Pedicel of worker of _P. floridana (193X)
Figure 53. Dorsal thoracic hairs of worker of P_. adrianoi
n. sp. (1,401X and 5,174X, respectively)
Dorsal thoracic hairs of worker of P^. adrianoi
n. sp. (1,401X and 5,174X, respectively)
Figure 54.


To my parents, wife and
and in memory of
son
Father Walter W. Kempf, O.F.M.


REVIEW OF LITERATURE
Four out of twelve species reported from Florida have that state as
the type locality: dentata Mayr (1886), P. floridana Emery (1895),
P_. metallescens Emery (1895), and _P. sitarches litorallis, Cole (1952).
Smith (1930) reported P. morrisi; Wheeler (1932) reported P_. anastasil;
Smith (1^33) reported _P. megacephala; Smith (1944) reported J?. dervtigula;
Smith (1951) reported _P. flavens sculptlor (misidentification); Van Pelt
(1956) reported P. pilifera; Smith (1958) reported ?. bicarinata vinelandica,
and Wojcik et al. (1975) reported P_. moerens. Additional information is
discussed under each species by the respective authors.
Creighton (1950) was the first, to try to give a extensive key to the
North America Pheidole; 63 species were listed and he used morphological
characters pertinent to both majors and workers. Creighton's statements
about the North American Pheidole are as follows:
Most of our species of Pheidole posses a dimorphic worker caste
with major and minor workers not connected by intermediates.
In a few species, however, the worker caste is polymorphic (kingi,
instabilis and torpescens, vasliti arizonica, etc.). Most of
the species garner seeds and it is believed that the large-headed
major workers function as seed-huskers. The enlarged head of the
major is mainly filled with mandibular muscles. This enables the
jaws to exert much pressure, which should be useful in cracking
off the husks of seeds. It may be added that sometimes the head
of the major is so large in proportion to its body that if the
insect is turned over on the back of its head it cannot regain
a normal posture without help from other workers. Despite
their preference for a graminivourous diet many species of
Pheidole will accept other food as well. They seem less attract
ed to honey-dew than do many ants but will often feed voraciously
on animal tissue when the opportunity offers. The majority of
our species form small colonies. In many cases there are only
3


96


26
3. Pheidole bureni n. sp.
Diagnosis:
A medium sized brownish red species with darker gaster, difering from
P. diversipilosa and P. crassicornis tetra by the absence of long erect
hairs on the dorsum of the first gastric tergite. It differs from P_.
crassicornis by having the occipital excision deeper and much more angularly
V shaped, and the occipital lobes sharper.
Description:
Major:
Measurements: Body length 5.30 to 5.70 mm. Head length 1.54
to 1.67 mm; head width 1.43 to 1.54. Thoracic length, 1.44 to 1.54
mm. Scape length, 0.85 to 0.92 mm; Scape Index, 54 to 55. Depth of
occipital excision usually over 0.12 mm.
Structural characters: Head shape as in fig. 60. Head excluding
mandibles longer than wide, broadest above the eyes, from where the
sides curve and narrow posteriorly; the occipital excision deep and
wide and angularly V shaped; occipital lobes much more angular than
in crassicornis or diversipilosa; frontal area impressed; anterior
border of clypeus with shallow median notch; irons strongly pronounc
ed and the vertex inclined posteriorly; genae flat and depressed
laterally. Frontal furrow distinct and extending from frontal area
to the occipital excision; scapes flattened and curved at base, a
little longer than half head length; mandibles curved laterally with
two strong, sharp teeth apically; the rest of the border without
teeth or with several small denticles; eyes measuring about 0.26 mm
in diameter and with over 100 small facets.


30
Color: Concolorously dark brown.
Female:
Measurements: Body length 7.50 mm. Head length 1.20 mm; head
width 1.38 mm. Thoracic length 2.41 mm. Scape length 1.03 mm. Scape
Index 86.
Structural characters: Head broader than long, broadest posterior
ly; occipital border straight; sides slightly convex and narrowing
anteriorly; middle of anterior border of clypeus with an impression;
frontal area broad and depressed; scapes stout, longer than 3/4 the
head length, the basal portions strongly curved and flattened, the
flattening more latero-mesal rather than dorso-ventral as in a major.
Eyes measuring about 0.40 mm in diameter and with over 200 small
facets; ocelli measuring about 0.15 mm in diameter; frontal carina
expanded and raised. Thorax flat dorsally, measured through the
Wing insertions slightly wider than the head; mesoscutum and scutelum
separated by a distinct suture.
Propodeal base and declivity not forming a distinct angle;
propodeal spines very short, thick and pointed dorsad; petiole stout; post
petiole twice as broad as long, the sides produced into distinct,
blunt lateral projections; these slightly convex anteriorly and
posteriorly weakly concave; gaster truncate at the base.
Sculpture: Frontal area and clypeus smooth and shining as well
as a narrow area of the frons between the frontal area and the median
ocellus; frons and lower vertex longitudinally striate laterally;
posterior areas of head smooth and shining; genae with pronounced


102
?*T
J /
f NiSi'Aj'"'--
i j L t-L-. \ > }-^srV


38
or any sexual forms. Probably the main chamber or chambers are located
very deep in the ground. It seems quite obvious that the colonies are
not large. The disproportional size of the major and its rarity also
seems to indicate that it has limited importance in sustaining the colony,
which is not true for most species of the genus.
Another feature of this species when disturbed is the ability of the
workers to flign death. The minors try to flex themselves around particles
of soil which make them very difficult to see. The majors hide themselves
and also seem to avoid light, but their large size and more shining surfaces
compared with the workers make them easier to spot.
Workers of this species, although not many, have been captured by
baiting near Gainesville.
One colony was located at Tall Timbers Research Station in 1974, which
failed to yield any major or sexual form.
Perhaps P. carrolli is a species which may be losing its major in the
evolutionary process. The fewer majors available show rather more mor
phological variation than is normal in most Pheidole species.
This species is named after Dr. John F. Carroll, my friend and colleague
during several years of graduate study at the University of Florida.


68
Color: Light reddish brown, appendages lighter.
Female and Male:
Unknown
Holotype: 1 worker, collected at Archbold Research Station.
Paratypes: 3 workers collected in two colonies of the host
Pheidole anastasii at the Archbold Biological Research Station in
Lake Placid, Florida, by M. A. Naves in August, 1975.
The holotype will be deposited in the Museum of Comparative
Zoology, Harvard University, Cambridge, Massachussetts. Paratypes
will also be deposited in the National Museum of Natural History,
Washington, D.C., the Florida State Museum, Department of Plant
Industry, Gainesville, Florida, and M. A. Naves Collection.
Discussion:
Wheeler (1904) described two parasitic species, Epipheidole inquilina
and Sympheidole elecebra. EL inguilina was parasitising colonies of P.
pilifera coloradensis and S^. elecebra was parasitising colonies of P_. ceres.
Both parasitic species were collected in Colorado and up to now have been
the only known ant parasites for Pheidole in the U.S.
E. inquilina was described from males and females collected in three
colonies of its host. The lack of workers and absence of the host queen
led Wheeler to the assumption that the host queen is dispatched by its
own workers once the parasite had secured entry into the nest, and that
this species deserved placement under a separate genus. Later Dr. M. R.
Smith (1941) announced the discovery of a single worker of inquilina^
possibly jeopardizing the status of Epipheidole as separate from Pheidole.


18
Mesonotum without transverse impression; spines thick and long but
blunt apically and pointed obliquely dorsad. Petiole stout, anterior
and posterior faces sloping, the crest weakly angular in profile.
Seen from above, the sides of post petiole semi-angular, the
lateral projections blunt and not well developed; gaster truncate
at base and smaller than head.
Sculpture: Strong cephalic longitudinal striae diverging
posteriorly and disappearing in the area posterior to the scape when
in repose; interstitial areas between striae with weak, fine reticulate
punctation; frontal carinae diverging and surpassing the scapes when
in repose; vertex, frontal area as well as the clypeus smooth and
shining; the semi-scrobes not distinct and marked only by lack of
sculpture; the genae with strong longitudinal striae, the interstitial
areas weakly reticulo-punctate; occipital lobes smooth and shining
with sparse small circular piligerous punctures, ventral surface of
head smooth and shining, its anterior margin without teeth; frontal
furrow with its posterior half bearing transverse rugules.
Seen from above, dorsum of the pronotum and anterior mesonotum
nearly smooth and weakly shining, mesopropodeal impression and adjacent
posterior area of mesonotum reticulo-punctate; basal area of propodeum
weakly reticulate and shining; declivous face usually shining.
In profile, pronotum, mesonotum and propodeum heavily reticulate,
the inner areas finely reticulo-punctate and shining; petiole finely
reticulo-punctate. Seen from above, dorsum of pronotum and anterior
mesonotum nearly smooth and weakly shining; mesopropodeal impression
and adjacent posterior area of mesonotum reticulo-punctate; basal


46
9. Pheidole fallax obscurithorax Santschi
Ht4
P. fallax arenicola var. obscurithorax Santschi, 1923. As#n>. Soc.
Ent. Belg. 63:58. Kempf, 1972. Studia Ent. 15:191.
Type locality: Cordoba, Argentina
Types : None in this country
Range : Probably westernmost Florida to at le^st Mobile,
Alabama in U.S. Probably widely ranging in South
America.
Diagnosis:
P_. fallax obscurithorax (fig. 7, 8), is a large, very dark species
over 6 mm in body length. Its characteristics, such as the heavily
sculptured head and thick scape which is curved meslally and more angular
laterally are unique among the Pheidole of North America.
Discussion:
_P. fallax obscurithorax (fig. 8) is a member of the difficult large
neotropical fallax group which has 6 subspecies and 6 varieties. The
specimens collected near the Florida border matched specimens in the
M.C.Z. identified by W. W. Kempf.
This species was collected close to the Florida and Alabama border
in Balbwin County, Alabama by Dr. John F. Carroll. E. 0. Wilson also
(personal communication) collected this species in Mobile, Alabama in
1950 within 1/2 mile of the ship docking area. It is apparent that this
introduced species has been in the U.S. for at least 26 years but, unlike
the imported fire ant, has not spread widely.
It is not a surprise to find another exotic ant species from South
America in the U.S. Brachymyrmex patagonicus Mayr (not published),


66
18. Pheidole quiescens n. sp.
Diagnosis:
A small yellowish brown species (fig. 46, 47) parasitic in nests
of P. anastasii. Queens, soldiers and males are still unknown. Soldier
caste probably lacking as in other species parasitic on Pheidole.
Obviously closely related to its host and to other small species in this
species group (P. floridana, P. moerens, and _P. greggi n. sp.) but dis
tinctive in its very long and delicate pilosity and its large and singular
ly shaped postpetiole which seems broadly and almost immovable articulated
with the gaster. In behavior the species seems unique for its habit of
standing motionless with open mandibles for long periods at a distance
from the host workers and their activities.
Description:
Maj or:
Unknown. Probably does not exist in this species.
Worker:
Measurements: Body length 1.90 to 2.08 mm. Head length 0.46
to 0.49 mm; head width 0.41 to 0.46 mm. Thoracic length 0.56 to
0.62 mm. Scape length 0.38 to 0.46 mm. Scape Index 89 to 96.
Structural characters: Head, excluding mandibles, slightly
longer than broad, posteriorly with a median sulcus and weak impression
which forms weak occipital lobes. Frontal area small, slightly depress
ed, with two longitudinal striae. Clypeus with a fine median stria-
tion; anterior margin curved and slightly notched in the middle.
Frontal carinae short and divergent; antenae 12 segmented and with a
3 jointed club as in nonparasitic species; scapes long and slender,


20
Pronotum and mesonotum, in profile, convex; propodeal spines
long but blunt, projecting obliquely dorsad; petiole stout, the
apex of the node angular; postpetiole strongly convex dorsally and
almost flat ventrally; gaster truncate at the base.
Sculpture: Head smooth and strongly shining; pleural surfaces
of pronotum shining; pleurae of mesonotum, propodeum and petiole
strongly reticulo-punctate, postpetiole weakly punctate.
Seen from above dorsum of pronotum and anterior mesonotum smooth
and strongly shining, posterior area of mesonotum, propodeal base
and propodeal declivity reticulo-punctate; dorsum of postpetiole and
gaster smooth and shining.
Pilosity: Hairs white, measuring less than 0.08 mm; sparse all
over the body, semi-erect and of mixed sizes on the head, long and
erect on the dorsum of thorax and long and slender on gaster; under
the stereo microscope the hairs seemed spatulate, but scanning micro
graphs later revealed that such hairs are branched at the tips (fig.
53, 54), therefore, plumose or sub-plumose.
Color: Ebony black, the appendages lighter, mandibles yellowish
to dark brown.
Females:
Measurements: Body length 4.60 mm. Head length 0.90 mm; head
width 0.95 mm. Thoracic length 1.46 mm. Scape length 0.51 mm. Scape
Index 58.
Structural characters: Head broader than long, broadest posteri
orly; occipital border slightly concave; sides slightly convex and


100


92


23
center of a small crater on the soil surface. The main flights of sexuals
occur in the summer in July and August. The species is entirely diurnal.
The main sources of food are small seeds, although it also scavenges small
dead arthropods. The majors as well as the workers usually forage for
food and help carry it back to the colony. It is not an agressive species.
The majors when disturbed try to escape to hiding places.
f\ adrianoi differs from P_. metallescens by its ebony black color,
absence of reticulate punctation on the head and by its smooth and strongly
shining occipital lobes and lack of metallic reflections. P. adrianoi
also differs from _P. littoralis by its small head, and the occipital lobes
not strongly rounded, sides of head not parallel and mandibles not stout
and strongly curved. P_. metallescens and P_. littoralis usually have a
reddish brown head and thorax, the gaster always darker or blackish as
opposed to P_. adrianoi which has a shining concolorously ebony black
body.


57
13. Pheidole littoralis Cole
P. sitarches littoralis Cole, 1952. Ann. Ent. Soc. Amer. 45:443. -
Gregg, 1958. N.Y. Ent. Soc. 66:30-39.
Type locality: Lido Beach, Sarasota, Florida
Type : A. C. Cole collection; M.C.Z.: Nat. Mus., A.M.N.H.
and W. S. Creighton collection. Both the A. C. Cole
collection and W. S. Creighton collection are now at
the Los Angeles County Museum of Natural History.
Range : Central Florida
Diagnosis:
P_. littoralis (fig. 20) has a reddish brown head and thorax, the gaster
black and shining. It differs from P. sitarches, P.. sitarches compestris
and ]?. sitarches soritis by the presence of deep, large piligerous foveolae
on the occipital lobes which also have no distinct transverse striae.
Discussion:
P_. littoralis is a distinct species related to the sitarches complex
as Cole (1952) suggested. This species was collected nesting in white
sand at the Archbold Biological Research Station, where it is very common.
It was found nesting side by side with £. metallescens and P. adrianoi n.
sp. The colonies seem to be quite small. Digging to 50 cm deep failed
to produce any brood chamber or any sexual forms. At about 10 to 20 cm
deep there was always a chamber full with small grass seeds and never
more than four majors. The major has a large head which is dispropor-
tional considering the small size of the rest of the body. Majors were
never observed outside the nest foraging. The small black, shining
workers can easily be confounded with P. metallescens and P. adrianoi
n. sp. workers.


33
4. Pheidole carrolli n. sp.
Diagnosis:
A reddish brown species related to the P_. pilifera group. It
differs from P_. pilif era and P_. pilif era artemisia by the absence of
coarse reticulate occipital rugae and blunt postpetiolar connules. It
differs from P_. pilif era coloradensis by its larger head, strongly
convex pronotum and anterior mesonotum, and longer and finer thoracic
and gastric hairs. It differs from P. pilifera pacifica by its larger
size and cephalic rugae extending to the occipital lobes and absence of
transverse occipital rugae.
Description:
Maj or:
Measurements: Body length 3.20 to 3.60 mm. Head length
1.80 to 1.90 mm; head width 1.69 to 1.80 mm. Thoracic length 1.36
to 1.44 mm. Scape length 0.77 to 0.82 mm. Scape Index 42 to 44.
Structural characters: Head shape as in fig. 14. Head, exclud
ing mandibles, longer than broad, occipital lobes prominent and well
rounded, separated by a broad and deep excision; a furrow present,
starting from the frontal area to the occipital excision. Frontal
area depressed and bearing a median longitudinal striation. Clypeus
with a weak median carinae which is continuous with the frontal area
striation; anterior border of clypeus with a shallow weak emargina-
tion; frontal carinae divergent; anterior ventral margin of the head
without teeth; scapes slender at base, thickening apically and measur
ing less than half the head length. Mandibles stout, strongly curved
laterally and with two apical large blunt teeth, the remaining border


INTRODUCTION
The genus Pheidole is one of the more important ant genera in
the world. Approximately 1000 taxa are known. Over 400 taxa are known
from the Neotropical region (Kempf, 1972), and about 77 taxa are now known
from North America north of Mexico. Pheidole spp. are abundant in many
areas, and live in varying habitats ranging from the humid tropics to
deserts. They are able to survive in some areas by their habits of
collecting and storing seeds as food resources. They are also scavengers
of dead insects and ether animals, and can be predaceous. Some species
tend aphids and other homopterans, but this food source, so important
to many ants genera, is probably of relatively minor importance to most
species of Pheidole.
Pheidole apparently is a genus of rather recent evolutionary develop
ment. No fossil Pheidole are known previous to the Miocene (Brown, 1973).
There are no Pheidole in the Baltic amber (lower Oligocene) although these
ambers contain large numbers of specimens of other ant genera which have
persisted almost unchanged since that epoch (Formica, Myrmica, Iridomyrmex
Caranonotus. and others). It has been said that Pheidole has undergone
a world wide dispersal since this time and has undergone a large pro
liferation of taxa as well as adaptation to many ecological nitches (Brown
1973) .
A study of the Pheldole of any area is one of the prerequisites to
understanding the formicid interactions of that area and this in turn
could be important in the development of pest management strategies, since


72
20. Pheidole vinelandica Forel
P. bicarinata race vinelandica Forel. 1886. Ann. Soc. Ent. Belg.
30:45.
vinelandica Mayr, 1886. Verh. Zool. Bot. Ges. Wien 36:458. -
Emery, 1895. Zool. Jahrb. Syst. 8:292. Forel, 1901. Am. Soc. Ent.
Belg. 45:348. Wheeler, 1904. Amer. Mus. Nat. Hist., Bui. 24:458. -
Smith, 1918. Ent. News 29:22. Davis and Beguaert, 1922. Brook. Ent.
Soc. 17:9. Smith, 1924. Ent. News, 35:54. Hayer, 1925. Ent. News
36:42 Cole, 1940. Amer. Midland Nat. 24:42. Wesson and Wesson, 1940
Amer. Midland Nat. 24:92-93.
P. (Allopheidole) vinelandica Forel, 1912 Mem. Soc. Ent. Belg.
19:237.
P. vinelandica laeviuscula Emery, 1895. Zool. Jahrb. Syst. 8:292.
Smith, 1951. Cat. Hym. Amer. No. of Mex. U.S.D.A. Man. 2:805.
P. vinelandica subsp. buccalis Wheeler, 1908. Bull. Amer. Mus.
Nat. Hist. 24:454. Smith, 1951. Cat. Hym. Amer. No. of Mex. U.S.D.A.
Mon 2:805.
_P. vinelandica longula var. castanea. Wheeler, 1915. Amer. Mus.
Nat. Hist. 34:405. Cole, 1937. Ent. News. 48:100.
JP. bicarimata buccalis Creighton, 1950. Bull. Mus. Comp. Zool.
104:171. Cole, 1953. Tenn. Ac. Sc. 28:297.
P. bicarinata vinelandica Creighton. 1950. Bull. Mus. Comp. Zool.
104:17a. Cole, 1956. Tenn. Acad. Sc. 31:114. Gregg, 1958. N.Y. Ent.
Soc. 66:18, 36. Reid and Nugara, 1961. Jour. Parasitology 47:885-889.
Smith, 1965. U.S. Dept. Agr., Tech. Bui. 1326:25-26.


70
Permanent social parasitsm with a useless worker caste is already
known to occur among ants. Burn (1942), when describing Formica reflexa
suggested that the consistant presence of workers of both _F. subsericea,
the host, and of small, timid workers of F. reflexa in every colony that
he found, was possibly a case of F. reflexa being a permanent inquiline.
King (1949) presented further evidence confirming Buren's suggestions.
King and Sallee (1951) suggested that the absence of pure colonies of F.
reflexa confirmed Buren's suggestions.
_P. guiescens seems to fit Buren's and King's descriptions of a
permanent social parasite which has retained a useless worker caste. P.
quies.cens was collected several times and always the host-workers out-
numered the parasite at least several to one. The new species was never
found as a pure colony.


74
North Dakota and Colorado also seem doubtful, and seem much more likely to
be misdetermined records of bicarinata.
P_. vinelandica is found nesting in grassy areas at Tall Timbers
Research Station. The minors and majors are easily seen foraging close to
the nesting site. The nest has several exits. They were observed carrying
seeds and small dead arthropods.


104
Mayr, G. L. 1886. Die Formicidin Der Vereinigten Staaten Von Nordamericana
Verhandlungen Der. Zool-Bot. Gesell. Wein, Verh. 36:419-464.
Smith, M. R. 1930. A List of Florida Ants. Fla. Ent. 14:1-6.
Smith, M. R. 1933. Additional species of Florida ants, with remarks.
Fla. Ent. 17:21-26.
Smith, M. R. 1941. The discovery of the worker caste of an inquilinous
ant, Epipheidole inquilina Wheeler. Proc. Ent. Soc. Washington
24:104-109.
Smith, M. R. 1944. Additional ants recorded from Florida, with descriptions
of two new subspecies. Fla. Ent. 27:14-17.
Smith, M. R. 1951. Formicidae. In: K. V. Krombein and B. D. Burks,
eds., Hymenoptera of America North of Mexico, Synoptic Catalog, U.S.D.A.,
Agr. Mon. 2:799-806.
Smith, M. R. 1958. Formicidae. In: K. V. Krombein and B. D. Burks,
eds., Hymenoptera of America North of Mexico, Synoptic Catalog,
U.S.D.A., Agr. Mon 2 (Suppl. 1):120.
Smith, M. R. 1967. Formicidae. In: K. V. Krombein and B. D. Burks, eds.,
Hymenoptera of America North of Mexico, Synoptic Catalog, U.S.D.A.,
Agr. Mon 2 (Suppl. 2):343-474.
Van Pelt, A. F., Jr. 1956. The Ecology of the ants of the Welaka Reserve,
Florida (Hymenoptera:Formic.idae). Amer. Midland Nat. 56:387.
Wheeler, W. M. 1904. Three new genera of Inquiline ants from Utah and
Colorado, Bull. Amer. Mus. Nat. Hist. 20:1-17.
Wheeler, WT. M. 1908. The ants of Texas, New Mexico and Arizona. Bull.
Amer. Mus. of Nat. Hist. 24:399-485.
Wheeler, W. M. 1910. Ants, their structure, development and behavior,
Columbia University Press, 663 pp.
Wheeler, W. M. 1932. A list of the ants of Florida with descriptions of
nev; forms. N.Y. Ent. Soc. Jour. 40:1-171.
Wilson, E.0. 1975. Enemy Specification in the Alarm-Recruitment System
of the Ant. Science 190:798-800.
Wojcik, D. P., W. A. Banks and W. F. Burn, 1975. First Report of Pheidole
moerens in Florida (Hymenoptera:Formicidae) U.S.D.A. Coop. Econ.
Ins. Rpt. 25(49-52):906.


MATERIALS AND METHODS
Four species of Pheidole (P. dantigula, _P. fallax obscurithorax,
P. bureni, n. sp. and P. greggi n. sp.) included in this research were
received from other collectors. All other specimens were collected by
the author during severa] trips in the state (fig. 67) or while at Tall
Timbers Research Station in Leon Co., Florida, during the summers of
1974 and 1975.
Colonies of Pheidole spp. were located in several ways:
a. Examingiug cavities under the bark of trees or examining grass
behind the leaf bases of palms.
b. Searching for nest craters, mounds, or any other modification
on the soil surface likely to have been produced by ants.
c. Searching for foraging majors or workers and following them to
the nest location.
d. Baiting with honey or dead arthropods and following trails of
workers and majors to the nest location.
3. Overturning stones, boards, or pieces of wood.
The two most successful methods were examining cavities under the
bark of pine trees and looking for craters on the soil surface.
Colonies were collected by aspirating the ants once the chambers were
located. Nests in the soil were excavated by removal of large blocks of
soil, breaking it carefully and aspirating the ants found. The depth
explored was variable, ranging from 30 to 50 era for P. metallescens
to over 1.5 m for P. lamia.
5


41
Diagnosis:
]?. dentata (fig. 30) is a reddish yellow to dark brown species
differing from P_. morrisi by the distinct and sharp propodeal spines.
It differs from P_. megacephala in not having a cordate head and differs
from P. vinelandica by the larger size and absence of a longitudinal
striation on the frontal area, and having the mesonotum transversely
impressed in the middle.
Discussion:
P. dentata is one of the more common species in Florida. It is
easy to understand why this species caused so many misunderstandings in
the past. There is much variation, not only in size and color, but
also in habitat preference. I have found colonies of small dark colored
specimens in wooded areas around Gainesville and large specimens that
nest in sandy soil on beaches in south Florida and the Florida Keys. I
also found a yellowish variant that inhabits the marshlands of the keys
and another variant with quite large majors that vary in color from
reddish to very dark brown nesting in open areas around Gainesville. This
morphological variation seems to be without taxonomic significant^
Three mated young queens collected at Gainesville were easily reared
in the laboratory. All three had majors after 50 days. The species is
highly carnivorous and the colonies have a tremendous growth when fed
with living arthropods. One of the colonies was inadvertantly left with
out food and moisture for three weeks. The result was that the adults
formed from the stressed broods were intermediate in size and structure
between workers and majors. Some of the intermediates were not as active


1. Pheidole ndrianoi n. sp.
Diagnosis:
A small ebony black species differing from IP. metallescens by the
absence of the violaceous or bluish reflections and without the reticulate
sculptured occipital lobes. It also differs from P. littoralis by the
small shining head, the head length usually less than 1.0 mm in the major
as opposed to over 1.4 mm in littoralis; workers with the erect hairs
plumose or semi-plumose at apex.
Description:
Major:
Measurements: Body length 2.70 to 3.10 mm. Head length 0.82
to 0.97 mm; head width 0.79 to 0.95 mm. Thorax length 0.79 to 0.90 mm;
scape length 0.45 to 0.51 mm. Scape Index 51 to 59.
Structural characters: Head shape as in fig. 25. Head, exclud
ing mandibles, usually slightly longer than wide, broadest behind
the eyes from where the sides slightly curve and narrow posteriorly,
the occipital excision gently concave. Frontal area impressed; anterior
border of clypeus with a wide, shallow median notch. A weak, scarcely
depressed antennal semi-scrobe present; a distinct frontal furrow
arising behind the frontal area, widening posteriorly into the
occipital excision. Scapes slender at base, thickening apically and
longer than the half the head length; mandibles stout, weakly curved
laterally, with two apical blunt teeth and two basal denticles. Eyes
small with approximately 35 facets.
Pronotum and mesonotum strongly convex in profile, posteriorly
the mesonotum more abruptly descending into the mesopropodeal impression.
17


32
Museum, Department of Plant Industry, Gainesville, Florida, and in the
W. F. Burn and M. A. Naves collections.
Discussion:
This species is named in honor of Dr. William F. Burn, Professor
of Entomology, University of Florida.


7
A trip was taken to the Museum of Comparative Zoology, Harvard
University, Cambridge, Massachussetcs, and to the National Museum of
Natural History in Washington, D.C., in order to study types of Nearctic
and Neotropical Pheidole.
All specimens were point mounted and studied by using a stereo micro
scope with magnification up to 160X. The pictures were taken using the
scanning electron microscope of the Insects Attractants Laboratory, Agri
cultural Research Service, U.S.D.A., Gainesville, Florida.
Several measurements were taken for the newly described species
including:
a. Body length: the sum of the head length, thoracic length, pedicel
and gastric lengths.
b. Head length: lengths of the. head in full face view (mandibles
excluded).
c. Head width: greatest width of the head in full face view.

d. Thoracic length: greatest length of thorax in lateral view.
e. Scape length: middle of antennal socket to the scape tip.
f. Scape index: found by computing the formula: Scape length X 100
head length
The following abreviations were used throughout this work:
M.C.Z. Museum of Comparative Zoology, Harvard University, Cambridge,
Massachussetts.
U.S. Nat. Mus. United States National Museum, Washington, D.C,
A.M.N.H. American Museum of Natural History, New York, N.Y.


PLATE 1
Figure
1.
Head of major of P.
lamia Wheeler (33X)
Figure
2.
Scape of major of P
of the crassicornis
. bureni n. sp., a member
complex (81X)
Figure
3.
Head of major of P.
diversipilosa Wheeler (45X)
Figure
4.
Gaster of major of
P. diversipilosa (47X)
Figure
5.
Head of major of P.
crassicornis Emery (43X)
Figure
6.
Gaster of major of
P. crassicornis (58X)




50
apically and bearing two teeth, the remaining border edentate,
except for a small basal tooth. Eyes small and with approximately
23 facets.
In profile, pronotum and mesonotum strongly convex and descend
ing through an abrupt declivity to the mesopropodeal impression
which is rather weakly impressed; propodeal spines small and sharp,
pointed obliquely dorsad; petiole with a short anterior peduncle;
anterior face of node gently sloping, the posterior face sharply
declivous; crest of node convex and blunt in profile but not thicken
ed and sub-truncate as in anastasii and floridana, ventrally the
petiole straight rather than convex as in anastasii and floridana;
postpetiole strongly convex dorsally, flat to slightly concave
ventrally.
Seen from above shoulders of pronotum well rounded; sides of
petiole parallel; postpetiole small; the sides convex and the shape
sub-trapezoidal, without lateral connules, little wider than petiole;
gaster truncate at base and slightly smaller than the head.
Sculpture: Head opaque, cephalic rugae sub-parallel mesally,
slightly diverging laterally and reaching close to occipital lobes
before disappearing; the interstitial areas between rugae with weak
reticulate punctation; the occipital lobes smooth and shining; the
scrobes finely reticulo-punctate; genae and areas posterior to the
eyes, as well as the area anterior to the eyes longitudinally rugose,
the rugae disappearing before reaching the occipital lobes, occipital
lobes with fine circular piligerous punctures.


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PLATE 12
Figure 63. Dorsum of propodeum of major of P. vinelandica
(272X)
Figure 64. Thoracic dorsum of P_. bicarinata (71X)
Figure 65. Postpetiole of P. bicarinata (186X)
Figure 66.
Head of P. vinelandica (66X)


54
Although Gregg showed that Burn's original identification of
sculptior from Florida specimens was in error, and that apparently
sculptior does not occur in the U.S., he left unresolved the question
of what name should be applied to the Florida species. The present
studies indicate that it is a new, undescribed species distinct from
P_. flavens Roger (fig. 55), P_. flavens sculptior Forel, ]?. floridana
Emery, P^. anastasii Emery and P_. moerens Wheeler.
Much more study is needed in order to fully understand this
difficult group which has several related species, 8 subspecies and
13 varieties in the new world.
This new species differs from _P. anastasii and P.. floridana by
the shape of the postpetiole which is slightly larger than the petiole
and does not have the distinct lateral connules. It differs from P_.
flavens by having the scrobe heavily reticulo-punctate, and the vertex
not smooth and shining. It differs from P. flavens sculptior by
having smooth and shining occipital lobes. It also differs from P_.
moerens by having a reticulo-punctate scrobe, which is opaque as
well as the entire head.


67
the posterior third thicker, usually passing beyond the posterior
margin of the head by an amount less than the length of first funicular
segment. Mandibles slender, curved apically and with two large, sharp
teeth, the remaining border with smaller sharp teeth; eyes convex,
measuring less than 1/4 the head length and with approximately 22
facets. In profile,pronotum and anterior mesonotum convex; posterior
of mesonotum more abruptly descending to the wide mesopropodeal
impression which is slightly depressed below the anterior propodeal
margin. Juncture of base and declivity of propodeum rounded; pro
podeal spines broad and heavy at base and pointed obliquely laterad.
Petiole with gently sloping anterior dorsal face, the posterior face
vertical, the node in profile very blunt apically.
Postpetiole, seen from above, slightly wider than long, two
times or more wider than petiole, the sides slightly angular but not
forming distinct conules, broadest in the posterior half, gaster
truncate at base and always noticeably larger than the head.
Sculpture: Head including irons and clypeus opaque and finely
reticulo-punctate; mandibles smooth and shining. Frontal area small
and finely reticulo-punctate. Sides of thorax and petiole, sub
opaque and reticulo-punctate. Postpetiole weakly reticulo-punctate
and weakly shining. First gastric tergite and usually the postpetiolar
dorsum with opalescent reflections and subopaque.
Pilosity: Hairs yellow, long, very slender and pointed, some
measuring up to 0.15 mm, numerous all over the body, even the ventral
surface of the head; short and sparse hairs on the legs; pubescence
limited to the antennae, absent on gaster and other regions of body.


is
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
LIST OF FIGURES Continued
Page
Head of the worker of P. crassicornis (54X) 90
Thorax of the worker of P_. metallescens (196X) 90
Head of the worker of P. littoralis (113X) 90
Worker of P_. carrolii n. sp. (40X) 92
Head of the worker of P. moerens (102X) 92
Head of worker of P_. greggi n. sp. (102X) 92
Worker of P. quiescens n. sp. (45X) 92
Dorsal view of worker of P. quiescens n. sp. (103X).... 92
Propodeum, pedicel, and gaster of worker of .P.
dentigula (102X) 9?
Dorsal view of thorax of P. dentigula worker (115X) 94
Head of worker of J?. anastasii (109X) 94
Head of worker of P_. floridana (115X) 94
Pedicel of worker of P_. floridana (193X) 94
Dorsal thoracic hairs of worker of P. adrianoi n. sp.
(1,401X and 5,174X, respectively) 94
Dorsal thoracic hairs of worker of P_. adrianoi n. sp.
(1,401X and 5,174X, respectively) 94
Head of major of _P. flavens (95X) 95
Head of major of P_. flavens (87X) 95
Occipital lobes of P_. greggi n. sp. (250X) -95
Postpetiole of the major of P. flavens sculptior (158X) 96
Head and thorax of the minor of P. flavens (83X) 96
Head of major of P. bureni n. sp. (39X) 93
x


TABLE OF CONTENTS
Page
ACKNOWLEDGMENTS iii
LIST OF FIGURES viii
ABSTRACT xii
INTRODUCTION 1
REVIEW OF LITERATURE 3
MATERIALS AND METHODS 5
KEY TO SPECIES OF PHEIDOLE OF FLORIDA (MAJORS ONLY) 8
KEY TO PHEIDOLE SPECIES OF FLORIDA (WORKERS ONLY) 13
1. Pheidole adrianoi n. sp 17
Diagnosis 17
Description 17
Major 17
Measurements 17
Structural characters 17
Sculpture 18
Pilosity 19
Color 19
Worker 19
Measurements 19
Structural characters 19
Sculpture 20
Pilosity 20
Color 20
Female 20
Measurements 20
Structural characters 20
Sculpture 21
Pilosity 21
Color 22
Male 22
Holotype 22
Paratypes 22
Discussion 22
2. Pheidole anastasii Emery 24
Diagnosis 24
Discussion 24
lv


35
finely reticulo-punctate and weakly shining. Several transverse
striae present between bases of spines; petiole and postpetiole
opaque and weakly reticulo-punctate; gaster smooth and shining
except in the basalmost area where it is opaque and weakly punctate.
Pilosity: Hairs yellow, pointed, short and semi-erect and sparse
on the anterior face of the head, short and long and semi-erect
ventrally; dorsum of thorax with numerous short and long erect
hairs measuring up to 0.26 mm; pedicel with numerous semi-erect
short and long hair; sparse pubescence confined to antenae and
appendages.
Color: Reddish brown, appendages lighter; dorsum of gaster
slightly darker; anterior margin of clypeus and the border of
mandibles dark.
Worker:
Measurements: Body length 2.30 to 2.40 mm. Head length 0.56
to 0.64 mm; head width 0.56 to 0.58 mm. Thoracic length 0.74 to 0.77
mm. Scape length 0.53 to 0.59 mm. Scape Index 92 to 94 mm.
Structural characters: Head usually a little longer than broad,
occipital borders weakly convex; frontal area impressed and with a
longitudinal striation in the middle; frontal carinae slightly divergent;
clypeus with a weak median carina and 2 longitudinal striae laterally;
anterior border curved and without any notch. Mandibles slender with
2 large and sharp apical teeth and 4 or 5 smaller teeth on the remain
ing masticatory border; scapes usually surpassing the occipital border
by an amount no greater than the first funicular joint; eyes large
with about 45 facets and measuring about 0.15 mm in diameter.


47
Iridomyrmex humilis (Mayr), Solenopsis invicta Burn, richteri
Forel, and Pheidole fallax obscurithorax appear to be a group of South
American ants inadvertently imported to the ports of New Orleans, and/or
Mobile, and/or possibly Pensacola.
All five species have ranges .along or near the Paraguay and La Plata
Rivers in Western Brazil, Paraguay and Argentina. The Paraguay River is
navigable by ocean going vessels up river to Asuncion, Paraguay.


is
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
LIST OF FIGURES
Page
Head of major of P. lamia Wheeler (33X) 78
Scape of major of P. bureni n. sp. a member of the
crassicornis complex (81X) 78
Head of major of P_. diversipilosa Wheeler (45X) 78
Gaster of major of P_. diversipilosa (47X) 78
Head of major of P_. crassicornis Emery (43X) 78
Gaster of major of crassicornis (58X) 78
Scape of major of P. fallax obscurithorax Santschi
(SIX) 80
Head of major of P. fallax obscurithorax (40X) 80
Head of major of _P. megacephala (Fabricius) (40X) 80
Pedicel of major of ]P. floridana Emery (128X) 80
Propodeum, pedicel and gaster of major of P.
metallescens Emery (74X) 80
Postpetiole of major of ]?. littoralis Cole (119X) 80
Head of major of P_. dentigula M. R. Smith (64X) 82
Head of major of P_. carrolli n. sp. (29X) 82
Pedicel of major of P_. metallescens Emery (163X) 82
Head of major of P_. metallescens (62X) 82
Head of major of P. anastasii Emery (68X) 82
Postpetiole and gaster of P. anastasii (62X) 82
Head of major of P. floridana Emery (72X) 34
Head of major of P. littoralis Cole (37X) 84
viii


69
S. elecebra was described from one queen and 18 males. Later another
queen was found. The worker caste, if any exists, is still unknown.
Brown (1973) listed both Epipheidole and Sympheidole as synonymous
with Pheidole, but these listings were not annotated and the validity
of these synonymizations is in doubt.
P. quiescens, in contrast with E. inquilina and S. elecebra, still
produces its worker caste. Several attempts to collect sexual forms of
_P. quiescens have failed. In two cases parasitised colonies still retain
ed the ho§t queen.
One of these colonies was brought to the lab for observation. It
was noted that the parasite workers essentially stand in the colony doing
nothing for long periods of time. They obtain their food by trophallaxis
from the host workers. They always stand with the mandibles open, possibly
in a defensive position, at a distance from the host workers. If the
colony happens to be disturbed the parasite will run away, apparently
seeking refuge. Twice they were observed helping to move brood from one
place to another but never were observed doing any other useful work. The
interspecific relationship seems to be inquilinism which means that the
parasite spends its whole life cycle in the host colony.
Subsequent collections of host workers and parasite workers were added
to the lab colony and no rejection of the newcomers was ever observed.
Later on, the host queen died and another queen of P. anastasii collected
in Gainesville, Florida was readily accepted by the colony. The host is
very well distributed all over the state, but collections throughout the
state failed to have parasites, although P. quiescens seems to be common
in colonies of its host at Archbold Biological Research Station.


75
SPECIES NOT PROPERLY INCLUDED IN THE FLORIDA PHEIDOLE FAUNA
1. Pheidole pilifera (Roger)
Leptothorax pilifera Roger, 1863. Berl. Ent. Zeitschr., 7:180.
_P. pilifera var. simulans Wheeler, 1908. Bull. Amer. Mus. Nat. Hist.,
24:436.
P_. pilifera subsp. septentrionalis Wheeler, 1908. Bull. Amer. Mus.
Nat. Hist. 24:436.
P. pennsylvanica Roger, 1863. Berl. Ent. Zeitschr., 7:199. Mayr,
1870. Verh. Zool-Bot. Ges. Wien, 20:981. Mayr, 1886. Verb. Zool-Bot.
Ges. Wien, 36:455. Mayr, 1877. Verh. Zool-Bot. Ges. Wien, 37:588.
P^. pilifera Emery, 1895. Zool. Jahrb. Syst. 8:290. Creighton, 1950.
Bull. Mus. Corap. Z-ol. 104:184-186. Van Pelt, 1956. Amer. Midland Nat.
56:377, 834. Smith, 1958. Cat. Hym. Amer. No. of Mex., U.S.D.A. Mon.
2 (Suppl. 1):123. Gregg, 1958. N.Y. Ent. Soc., 66:24-25.
Type locality: Pennsylvania
Types : None in this country
Range : Massachussetts to North Carolina and West to Iowa
and Nebraska
Diagnosis:
P_. pilifera differs from P. carrolli n. sp. and _P. littoralis Cole
by having the occipital rugae of the major notably reticulate and often
coarse; its occipital lobes are not well rounded and pronounced.
Discussion:
Attempts were made by the author to find P. pilifera in Florida as
reported by Van Pelt (1956). Dr. Van Pelt, however, expressed some reserva
tions about the correctness of this identification. No voucher specimens


40
6. Pheidole dentata Mayr
P_. morrisi var. dentata Mayr 1886. Vehr. Zool Bot. Ges. Wein,
36:457.
P. dentata Forel, 1901. Ann. Soc. Ent. Belg., 45:351 Wheeler,
1904. Amer. Mus. Nat. Hist., Bull. 20:302. Mitchell and Pierce, 1912
Ent. Soc. Wash., Proc. 14:71. Smith, 1918. Ent. News, 29:22. Smith,
1924. Ent. News 35:77. Smith, 1930. Fla. Ent. 14:3. Wheeler, 1932.
N.Y. Ent. Soc., 40:6. Dennis, 1938. Ent. Soc. Amer., Ann. 31:281,
304. Van Pelt, 1950. Ent. News 61:161-163. Creighton, 1950. Bull.
Mus. Comp. Zool. 104:177. Wheeler and Wheeler, 1953. Ent. Soc. Wash.
Proc. 55:71. Gregg, 1953. N.Y. Ent. Soc. 66:21-22. Smith, 1965.
U.S. Dept. Agr., Tech. Bui. 1326, pp. 27-28. Wilson, 1975. Science
190:798-800.
P. dentata var. faisonica Forel., 1901. Ann. Ent. Soc. Belg.,
45:352.
P. commutata Mayr 1886. Verb. Zool. Bot. Ges. Wein. 36459. -
Emery, 1895. Zool Jahrb. Syst., 8:289.
Leptothorax tennesseensis Cole, 1938. Amer. Midland Nat. 19:238.
fh dentata var. commutata Wheeler, 1907. Amer. Mus. Nat. Hist.,
Bull. 7:18. Wheeler, 1908. Amer. Mus'. Nat. Hist., Bull. 24:460-461.
Smith, 1924 Ent. News 35:77. Cole, 1940. Amer. Midland Nat. 24:29,
44. Smith, 1951. Cat. Hym. Amer. No. of Mex., U.S.D.A. Mon 2:802.
Type locality: Florida
Types : None in this country
Range : Southeastern States, Tennessee and Gulf states to
Texas.


PLATE 10
Figure 55.
Figure 56.
Figure 57.
Figure 58.
Head of major of P_. flavens (95X)
Head of major of P_. flavens sculptior (87X)
Occipital lobes of P_. greggi n. sp. (250X0
Postpetiole of the major of P. flavens sculptior
(158X)
Figure 59.
Head and thorax of the minor of P. flavens (83X)


9
b) Species usually measuring 5 mm or less; if larger the head
never heavily sculptured; the scapes never thick or
strongly bent at base 5
5a) Head cordate, gradually but distinctly narrowed toward
the mandibular insertions; broadest at the occipital
lobes; mesonotum not transversely impressed.
(fig. 9) megacephala (Fabricius)
b) Head not cordate and never strongly narrowed towards
the mandibular insertions; mesonotum may or may not be
transversely impressed 6
6a) Postpetiole, seen from above, bearing distinct lateral
connules; head covered with reticulate punctation or
longitudinally striate. (fig. 10, 11) 7
b) Postpetiole seen from above, without distinct lateral
connules; if indistinct lateral projections present
the posterior half of the head smooth and shining.
(fig. 12) 11
7a) Occipital lobes with reticulate sculpture, with no trace
of transverse or longitudinal striation, the surface
opaque or feebly shining; post-petiolar width two times
length. (fig- 13) dentigula M. R. Smith
b) Occipital lobes smooth and shining or with sparse
punctures or striae; postpetiole width greater than length,
but never two times length. (fig. 10, 11) 8
8a) Sides of head sub-parallel, narrowing slightly anteriorly;
head length 1.82 to 2.0 mm; scape index 41 to 42; pronotum


63
the types of dominicensis and they appear identical with the types of
moerens except in color. I do not consider dominicensis a valid taxon.
This species was found nesting in various places such as under boards,
at base of oak trees and fence posts, along roots, under palm leaves,
inside wall crevices, and rarely in the ground. Usually small chambers
are constructed. It is a monogynous species which has a small blackish
female. Its main flight is usually in July. Several queens may start
founding a nest, but before the first brood emerges, the dominant female
will have killed the others.
The chambers usually are built with small soil or debris particles
and have small openings. A colony may have over 100 majors and over 300
workers. They feed on seeds and scavenge and prey on small dead or live
arthropods. They forage very close to the nesting sites and sometimes a
major is found foraging along with the workers.


45
specimens of P_. diversipilosa in Gainesville. Comparison with the types
and cotypes for the two taxa in the M. C. Z. and further material of
diversipilosa collected at Columbus and Mission, Texas by W. F. Burn
as well as the finding of another related species, P_. bureni n. sp.
(fig. 60, 61, 62) emphasize that we are dealing with a species group
that is widely spread and contains several discrete species. P. crassicornis
is predominantly an eastern species. P. diversipilosa, P_. bureni and
P_. crassicornis tetra are western species. Of these apparently only
P_. diversipilosa reaches east into Florida. The lack of better records
for the distribution of these species probably led Creighton (1950) to
synomymizing crassicornis and diversipilosa. _P. crassicornis tetra
appears to be a very distinct form and may be a discrete species also,
but I have not attempted taxonomic treatment in the present paper, since
it does not occur in Florida.




PLATE 11
Figure 60. Head of major of P. bureni n. sp.
Figure 61. Gaster and pedicel of £. bureni n
Figure 62. Head and pronotum of worker of P.
(52X)
(39X)
. sp. (40X)
bureni n.


88


KEY TO SPECIES OF PHEIDOLE OF FLORIDA
(MAJORS ONLY)
la) Head cylindrical in cross section and obliquely truncate
anteriorly, the truncation involving the clypeus, frontal
area, anterior portions of genae, and mandibles.
(fig. 1) lamia Wheeler
b) Head not cylindrical in cross section and not truncate
anteriorly 2
2a) Antennal scape strongly bent, flattened and smooth at the
base, the base nearly as or as broad as the distal portions
of the scape (fig. 2) 3
b) Not as above, scape not flattened at base and usually
distinctly narrower than distal portion; if thickened
and strongly bent at base, obviously not flattened 4
3a) First gastric tergite covered with very numerous long and
short erect hairs; sometimes sparse, coarse, appressed,
pubescence is present, (fig. 3, 4) diversipilosa Wheeler
b) First gastric tergite covered with appressed pubescence
and sometimes with sparse long hairs.
(fig. 5, 6) crassicornis Emery
4a) Large species, total length over 6 mm; head heavily
sculptured; scapes very thick and bent at base. The
scapes seen in full face view are evenly curved measally,
but more angular laterally (fig. 7, 8)..fallax obscurithorax Santschi
8


PLATE 5
Figure 25. Head of major of P_. adrianoi n. sp. (72X)
Figure 26. Thorax, pedicel and gaster of major of P_.
adrianoi n. sp. (50X); white legs are
artifacts due to electron overcharge.
Figure 27. Head of major of P_. morrisi Forel (37X)
Figure 28. Thorax of major of P. morrisi (56X)
Figure 29. Head of major of P_. dentata Mayr (48X)
Figure 30. Teeth on the anterior ventral head margin
P. anastasii Emery (123X)


KEY TO PHEIDOLE SPECIES OF FLORIDA
(WORKERS ONLY)
la) Entire head smooth and shining. (fig. 31, 36) 2
b) Head reticulo-punctate or partially reticulate with some
smooth areas. (fig. 40, 41, 44) 9
2a) Scapes failing to reach the posterior border of the head,
or surpassing it by an amount no greater than the first
funicular joint. (fig. 31) 3
b) Scapes surpassing the posterior border of the head by an
amount greater than the first funicular joint. (fig. 34) 6
3a) Thorax smooth, nearly free of sculpture and strongly
shining; yellow 4
b) Pronotum smooth and shining; mesonotum and propodeum
punctate or sculptured; color various. (fig. 32, 33) 5
4a) Propodeum armed with angular teeth which are broad at the
base and do not resemble spines; mesonotum and propodeum
in profile separated by a deep, wide impression.
('MgvBS1) lamia Wheeler
b) Propodeum armed with spines; mesonotum and propodeum in
profile separated by a normal impression tysoni Forel
5a) Yellowish to brownish species, gaster covered with numerous
long fine hairs, (fig. 32) vinelandica Forel
b) Ebony black species, gaster with plumose or subplumose
hairs which are spaced at distances more or less equal to
the hair's length. (fig. 33) adrianoi n. sp.
13


71
19. Pheidole tysoni Forel
_P. tysoni Forel, 1901. Soc. Ent. Belg., Ann. 45:348. Wheeler,
1904.
Amer.
Mus.
Nat. Hist. 20:302. Smith,
1918.
Ent.
News 29:
22. -
Smith,
1924.
Ent
. News 35:78. Dennis, 1938.
Ent.
Soc.
Amer. 31
: 282,
304. -
Wesson
and
Wesson, 1940. Amer. Midland
Nat.
24:92
. Cole,
1940
Amer. Midland Nat. 24:42, 45. Creighton, 1950. Bull. Mus Comp. Zool.
104:191. Smith, 1951. Cat. Hym. Amer. No. of Mex., US.D.A. Mon 2:805. -
Gregg, 1958. N.Y. Ent. Soc. 66:32.
Type locality: Mt. Mitchell, North Carolina
Types : A. M. N. H., M. C. Z.
Range : North Florida, Georgia, western North Carolina,
southwestern Virginia and eastern Tennessee.
Diagnosis:
P_. tysoni is a small yellowish species closely related to P. vinelandica,
differing from it by the teeth on the anterior ventral margin of head. The
minor has the propodeal pleurae largely free from sculpture and strongly
shining.
Discussion:
_P. tysoni was collected once in a soybean field at Tall Timbers
Research Station in 1973. The colony was quite large and the main chamber
was about 50 cm deep in the ground. Only minors were foraging. This is
the only record known for the state and I am assuming that P. tysoni is
rare in Florida.


u
6a) Mesonotum without a distinct tranverse impression; in
profile postpetiole longer than petiole and dorsal and
ventral outlines both smoothly convex.
(fig. 34) megacephala (Fabricius)
b) The mesonotum with a distinct transverse impression.
(fig. 35, 37 39) 7
7a) Posteriorly on head a distinct collar visible in full
face view (fig. 36); the mesonotum and base of propodeum
heavily punctuate. (fig. 35) fallax obscurithorax Santschi
b) Posterior collar of head not visible in full face view;
the mesonotal dorsum and propodeal base weakly sculptured
and shining 8
8a) Usually yellowish or pale species, the propodeal spines
absent. (fig. 37) morrisi Forel
b) Dark brown to ferruginous species; propodeal spines well
formed. (fig. 38, 39) dentata Mayr
9a) Scapes surpassing the posterior margin of the head by a
distance greater than the first funicular joint. (fig. 40) 10
b) Scapes either not surpassing posterior margin of the head,
or surpassing less than the length of the first funicular
joint. (fig. 43) 11
10a) First gastric tergite with appressed pubescence and only a
few scattered long hairs crassicornis Emery
b) First gastric tergite covered with numerous long and short
hairs; pubescence usually erect or semi-erect, rarely
appressed
diversipilosa Wheeler


Two additional taxa are elevated to full species: JP. vinelandica
(= JP. bicarinata vinelandica Forel) and P_. littoralis (= JP. sitarches
littoralis Cole) The _P. crassicornis complex is recognized with the
description of 1?. bureni n. sp. collected in Alice, Texas even though
this species does not occur in Florida.
The four new species, P_. adrianoi n. sp., JP. bureni n. sp. JP.
carrolii n. sp., and JP. quiescens n. sp., plus the introduced species
P^. fallax obscurlthorax Santschi and the recognized species 1?. diversi-
££ sa Wheeler increase the total Pheidole fauna of North America north
of Mexico to seventy-seven. The Florida Pheidole fauna is increased
to nineteen species and Pheidole is now the largest ant genus in the
state.
Keys for the identification of both majors and workers are pro
vided. Scanning electron micrographs are also provided as identifica
tion aides. The biology of each species is discussed.
xiii


2
ants are probably the most numerous and most important animals within
their size range.
One species, Pheidole megacephala, has become a tropicopoiitan tramp
species, and is a serious pest in various areas (Wheeler. 1910, Flucker
and Beardsley, 1970 and Lieburg, 1975). Some species sting severely
and are comparable with the fire ants in aggressiveness (Burn, personal
communication). Most of the species, however, appear innocuous and are
usually unnoticed by man. Their importance in the ecosystem appears
largely unappreciated.
Due to its size and distribution, Pheidole is a very difficult genus
and has been a challenge to myrmecologists taxonomically as well as
biologically.
Until recently only eleven species were known from Florida, which
suggested that despite its subtropical warmth Florida was deficient in
Pheidole when compared to Texas (36 species) and Arizona (26 species).
The Florida Pheidole fauna has long been known to be more extensive than
that of northern states such as Iowa and New Jersey (3 species each,
Burn, 1944 and Creighton, 1950).
This study was undertaken to increase our knowledge of Pheidole
of Florida. Scanning electron micrographs are provided along with keys
for the identification of both majors and workers.


PLATE 4
Figure
19.
Head
of
major
of
p.
floridana Emery
(7 2X)
Figure
20.
Head
of
major
of
p.
littoralis Cole
(37X)
Figure
21.
Head
of
major
of
p.
greggi n. sp. (68X)
Figure
22.
Head
of
major
of
p.
moerens Wheeler
(58X)
Figure
23.
Clypeus and frontal area of major of
vinelandica Forel (186X)
P.
Figure
24.
Head
of
maj or
of
p.
bicarinata Mayr
(53X)


21
narrowing anteriorly toward the mandibular insertions; anterior
border of clypeus weakly notched in the middle.
Frontal area depressed; antennae like those on the major;
scapes longer than half the head length; eyes measuring about 0.24 mm
in diameter and located on anterior half of head; ocelli measuring
about 0.12 mm in diameter.
Thorax flat dorsally, narrower than the head measured through
the wing insertions. Mesoscutum and scutellum well separated; propodeal
base and declivous face not forming a distinct angle. Propodeal spines
stout, blunt and directed posteriorly; petiole stout, the node seen
from above with parallel sides, the crest slightly depressed in the
middle; postpetiole twice as wide as petiole, with rather angular
lateral projections; gaster truncate at the base.
Sculpture: Cephalic rugae strong, covering most of the anterior
part of the head, but disappearing before reaching the occipital border,
the interstitial areas between rugae with widely spaced weak reticulate
punctation; frontal area smooth and shining with two mesal longitudinal
striae; frontal carinae long and diverging posteriorly.
Sides of pronotum and propodeum reticulate; episternum smooth
and shining; propodeal base and declivous face smooth and shining;
petiole and postpetiole punctate and opaque; gaster, seen from above,
with fine longitudinal striae near the base, the interstitial areas
between striae shining but with widely spaced reticulate punctation,
the rest shining and with circular piligerous punctures.
Pilosity: Hairs yellow, pointed; dorsum of thorax, pedicel and
gaster with numerous short and long hairs; gaster usually more pilose
than the remaining parts; pubescence present on the antennae and legs.


76
of P. pilifera are presently available from Dr. Van Pelts work. Although
it seems possible that P. pilifera occurs in Florida, I am unconvinced
that there is any direct evidence. I believe all reported specimens were
probably carrolli n. sp., P. littoralis Cole, or other species.


53
The species is named in honor of Dr. Robert E. Gregg, Professor
of Entomology, University of Colorado.
Gregg (1958) expended a great deal of effort trying to identify
a sample of this species that he and Mrs. Gregg collected in Brickel
Hammock, Miami, Florida. In his own words:
The specimens of sculptior from Miami, Florida (Burn
det.) and the sample obtained from Brickel Hammock are
indistinguishable. Despite the similarities of the
scrobes and their sculpture, these ants are not identical,
however, with the examples of sculptior from Puerto Rico
and Martinique, nor with the types from St. Vincent. This
is evident especially from the smooth (almost sculpture
less) and shining occipital lobes of the Florida ants. In
sculptior, the cephalic rugae and punctures completely
cover the head, making it opaque, except at the extreme
posterior margin around the foramen which is smooth and
shining. In addition, the epinotal spines of the Brickel
Hammock ants are longer and sharper than those of sculptior.
These two samples of the flavens group in Florida cannot
represent P_. flavens sculptior (from the West Indies), and
unless this subspecies is known from Florida by other
specimens, or is there but not yet collected, this discovery
requires a revision of the North American list. Pheidole
flavens sculptior must be dropped, and that is the plan
followed in this paper.
(p. 45)
Fig. 21 shows the head of P_. greggi from the original specimens
collected at Miami, Florida, by W. F. Burn. This confirms Gregg's
conclusion that they are not conspecific with sculptior. The occipital
lobes of P.. flavens sculptior from St. Thomas, Virgin Islands, also
collected by Burn on January 18, 1952, are shown in fig. 56.
The specimens collected at Naples were from a colony at the base
of a telephone pole on sandy soil close to the beach. The colony was
quite sizeable as in _P. anastasii, P. floridana and P. moerens.


52
dorsad; petiole slender anteriorly. Seen from above, postpetiole
sub-globular and slightly wider than the petiole.
Sculpture: Head reticulo-punctate; frontal area and clypeus
weakly reticulo-punctate and opaque; mandibles with piligerous
punctures; thorax, basal and declivious face of propodeum as well
as the petiole reticulo-punctate; dorsum of postpetiole and gaster
smooth and shining.
Pilosity: Hairs yellow, dorsal surface of head as well as the
ventral surface with numerous mixed short and long erect and semi-
erect hairs; dorsum of thorax with perdominantly long erect hairs;
pedicel and gaster with long semi-erect numerous hairs.
Color: Concolorously yellow
Female and Male:
Unknown
Holotype: Major (collected on December 19, 1945 in Miami,
Florida by W. F. Burn).
Paratypes: 39 majors, 26 workers
Discussion:
The type material comprises23 majors and 16 workers, collected on
December 19, 1945 in Miami, Florida, by W. F. Burn; 6 majors and 6 workers
collected in 1975 in Baldwin County, Florida by D. P. Wojcik; 10 majors
and 8 workers collected on November 25, 1975, in Naples, Florida by M. A.
Naves.
The holotype and several paratypes will be deposited in the Museum
of Comparative Zoology, Harvard University, Cambridge, Massachussetts.
Paratypes will be deposited in the Nation Museum of Natural History,
Washington, D.C., Florida State Museum, Department of Plant Industry,
Gainesville, Florida; and in the W. F. Burn and M. A. Naves collections.


59
head narrowing toward the mandibular insertion and the mesonotum is not
distinctly transversely impressed.
Discussion:
P. megacephala is a well known world tramp (Gregg, 1958). It has
been present in Florida for 43 years, at least, being reported by
Smith (1933) from Everglades, Key West and St. Augustine. I collected
this species in downtown Homestead, Florida on November 25, 1975 nesting
under a paved sidewalk and at Archbold Biological Research Station.
Burn observed large colonies of this species trailing in large numbers
from the colony to nearby trees (Personal Communication). Its ecological
impact has been studied in relation to another world tramp, Iridomyrmex
humilis. Flucker et al^. (1970) reported I?. megacephala was losing
territory to I. humilis in Hawaii, although it did not appear to be
evenly remotely approaching extermination. They also found that _?.
megacephala did not seem to occupy territory above 9.15 m, above which I.
humilis was usually found. Dieberburg _et ajL. (1975) reported that in a
7-year period P_. megacephala did not loose much territory to _I. humilis
in Bermuda. He also reported that .P. megacephala prefers clumped food
sources and 1^. humilis prefered dispersed food sources. Based on these
findings they suggested that a equilibrium between the two species was
possible in Bermuda. Brown (1973) suggested that P. megacephala was a
mortal enemy of I. humilis and that, the latter was vanquishing JP.
megacephala outside the tropics.
In Florida the two species seem to occupy two distinct areas. 1.
humilis, although not common, occurs in the northernmost and northwest


60
areas of the state. P_. megacephala, also not common, occurs in the southern
area of the state. In the central area of the state I was able to find
only P. megacephala. At least in Florida the two species are not sympatric
and the inability of either of them to increase must be due to some other
cause, possibly competition with Solenopsis invicta Burn or Solenopsis
geminat.a (Fabricius).


LITERATURE CITED
Brown, W. L. 1973. A comparison of the Hylean and Congo-West African
rain forest ant faunas, p. 161-185. In: B. J. Meggers, E. S.
Ayensu, and W. D. Duckworth (ed.). Tropical forest ecosystems in
Africa and South America:A comparative review. Smithsonian Institution
Press, Washington, D.C.
Burn, W. F. 1942. New ants from Minnesota, Iowa, and Wisonsin. Iowa
State College Jour. Sci. 16:399-40.
Burn, W. F. 1944. A List of Iowa Ants. Iowa State College Jour. Sci.
18:277-312.
Burn, W. F., M. A. Naves, and T. Carlysle. False Phragmosis and Apparent
Specialization for Subterranean Warfare in Pheidole lamia Wheeler
(Hymenoptera:Formicidae) In press.
Cole, A. C. 1952. A New Pheidole (Hymenoptera:Formicidae) From Florida.
Ent. Soc. Amer., Ann. 45:443-444.
Creighton, W. S. 1950. The ants of North America. Bull of the Mus. Comp.
Zool, Harvard, 104:1-585.
Emery, C., 1895. Beitraege Zur Kenntnis der nordamerikanischen Ameisenfauna.
II. Zool. Jahrb. Syst. 8:257-360.
Fluker, S. S., and J. W. Beardsley, 1970. Sympatric Associations of Three
Ants:Iridomyrmex humilis, Pheidole megacephala, and Anoplolepis
longipes in Hawaii. Ent. Soc. Amer., Ann. 63:1290-1296.
Gregg, R. E. 1958. Key to The Species of Pheidole (Hymenoptera:Formicidae)
in the United States. N.Y. Ent. Soc., Jour. 66:7-48.
Gregg, R. E. 1969. New Species of Pheidole from Pacific Coast Islands
(Hymenoptera¡Formicidae) Entom. News 80:93-101.
Kempf, W. W. 1972. Catalogo abreviado das Formigas da Regiao Neotropical
(Hymenoptera:Formicidae). Studia Ent. 15:3-344.
King, R. L. 1949. Mixed Colonies in Ants. Iowa Acad. Sci. 56:367-370.
King, R. L., and R. M. Sallee. 1951. More Mixed Colonies in Ants. Iowa
Acad. Sci. 58:487-489.
Liebergurg, I., P. M. Krans, and A. Seip. 1975. Bermudian Ants revisited.
The Status and Interaction of Pheidole megacephala and Iridomyrmex
humilis. Ecology. 56:473-478.
103


58
14. Pheidole megacephala (Fabricius)
Formica megacephala Fabricius, 1793. Ent. Syst. 2:361.
_P. megacephala Emery 1921. Gen. Insect. Fase. 174, p. 85. Phillips
1934. (Hawaii Univ.) Expt. Sta. Pineapple Prod. Coop. Assoc. Bui. 15:5-12. -
Smith, 1936. Puerto Rico Univ., Jour. Agr. 20:843-844. Broekhuysen, 1948.
Union So. Africa Dept. Agr. Bui. 266:1-40. Smith, 1951. Cat. Hym. Am.
No. of Mex. U.S.D.A. Mon 2:803. Wheeler and Wheeler, 1953. Ent. Soc.
Wash., Proc. 55:75. Brown, 1958. Act. Hym. 1:47. Gregg, 1958. N.Y.
Ent. Soc. 66:23. Brown, 1959. Bui. Ent. Res. 50:523. Weber, 1960. Ent.
Soc. Wash., Proc. 62:232. Taylor and Wilson, 1961. Psyche 68:143. Kempf,
1962.Studia Ent. 5:18-19. Haskins and Haskins, 1965. Ecology 46:737. -
Wilson and Taylor, 1967. Pacific. Insects Monog. 14:46-48. Fluker, Huddle
ston, and Beardsley, 1968. Jour. Econ. Ent. 61:474. Fluker and Beardsley,
1970. Ent. Soc. Amer., Ann. 63:1290-1296. Kempf, 1972. Studia Ent. 15:196.
Oecophthora pusilla Heer, 1852. Stueck 66:15. Wheeler, 1922. Bull.
Amer. Mus. Nat. Hist. 45:812.
_P. laevigata Mayr, 1862. Verli. Zool. Bot. Ges. Wien 12:747. -
Mayr, 1870. Ve.hr. Zool. Bot. Ges. Wien 20:981.
Type locality: Isle de France (Mauritius)
Types : None in this country
Range : This species has become a world tramp; in Florida it
apparently is largely confined to South Florida.
Diagnosis:
P- megacephala (fig. 9), is a dark yellow to brownish-ferruginous
species that differs from P. morrisi by having distinct and sharp propodeal
spines. It also differs from P^. morrisi and _P. dentata by having a cordate


73
P. vinelartdica vinelandica. Smith, 1951. Cat. Hym. Amer. No. of
Mex. U.S.D.A. Mon. 2:806.
Type locality: Vineland, New Jersey.
Types : Cotypes, A.M.N.H.
Range : Southeastern and Gulf States and along the east
coast to New Jersey
Diagnosis:
P_. vinelandica is a small yellowish species differing from P. bicarinata
by having the basal face of propodeum in the major largely punctate (fig.
0
m and having a longitudianl striation on the frontal area (fig. 66). It
differs from P. tysoni by the absence of teeth on the anterior ventral
margin of the head.
Discussion:
I had the opportunity to study specimens of P. vinelandica from New
Jersey, North Carolina, South Carolina, Georgia, Alabama, and Texas. I
have also collected this species at Tall Timbers Research Station as well
as in Putman Co., Florida. I have compared these with specimens of P_.
bicarinata (fig. 24, 64, 65) from Illinois in the M.C.Z. I also have
studied specimens of bicarinata collected at Colorado Springs, Colorado;
Ames and Lewis, Iowa; Devils Tower, Wyoming; Ortonville, Minnesota, and
Corinne, Utah. It was evident that the two forms are not conspecific
and have constant morphological differences, emphasizing that P. vinelandica
deserves full species rank. Gregg (1958) included the var. nebrascensis
Forel and the var. huachucana Smith as synonymus of vinelandica. I have
not studied the types of these taxa but am doubtful about their inclusion
under vinelandica rather than bicarinata. Records of vinelandica from


48
10. Pheidole floridana Emery
Z* flavens floridana Emery, 1895. Zool. Jahrb., Abt. f. System.
8:293. Smith, 1930. Fla. Ent. 14:3.
£. floridana Emery, 1895. Bull. Soc. Ent. Ital. 28:77. Wheeler,
1932. N.Y. Ent. Soc., 40:6. Creighton, 1950. Bull. Mus. Comp. Zool.
104:179. Smith, 1951. Cat. Hym. Amer. No. of Mex., U.S.D.A. Mon.
2:802. Gregg, 1958. N.Y. Ent. Soc. 66:21-22, 35. Kempf, 1972.
Studia Ent. 15:192.
Type locality: Coconut Grove, Florida
Types : M.C.Z.
Range : Southwest Florida
Diagnosis:
P. floridana (fig. 19) is a small yellowish species differing from
.P. greggi n. sp. and P_. moerens by the distinct lateral connules on the
postpetiole. It differs from P. anastasii by the absence of opalescence
on the dorsum of the first gastric tergite.
Discussion:
jP. floridana seems to be confined to southeast Florida in the Miami
area. This is the only place where I was able to locate this species.
Due to its close relationship to ]?. anastasii the latter has been mis-
identified as P_. floridana many times, thus, mistakenly extending the
supposed range of P. floridana P. anastasii is actually the species
widely distributed in Florida, while floridana is absent or at least
must be rare in most of the state.


TABLE OF CONTENTS Continued
Page
15. Pheidole metallescens Emery 61
Diagnosis 61
Discussion 61
16. Pheidole moerens Wheeler 62
Diagnosis 62
Discussion 62
17. Pheidole morrisi Forel 64
Diagnosis 64
Discussion 65
18. Pheidole quiescens n. sp 66
Diagnosis 66
Description 66
Major 66
Worker 66
Measurements 66
Structural characters 66
Sculpture 67
Pilosity 67
Color 68
Female and male 68
Holotype 68
Paratypes 68
Discussion 68
19. Pheidole tysoni Forel 71
Diagnosis 71
Discussion 71
20. Pheidole vinelandica Forel 72
Diagnosis 73
Discussion 73
SPECIES NOT PROPERLY INCLUDED IN THE FLORIDA PHEIDOLE FAUNA .... 75
1. Pheidole pilifera (Roger) 75
Diagnosis 75
Discussion 75
LITERATURE CITED 103
BIOGRAPHICAL SKETCH 105
vii


PLATE 8
Figure 43.
Figure 44.
Figure 45.
Figure 46.
Figure 47.
Figure 48.
Worker of 1?. carrolli n. sp. (40X)
Head of the worker of P. moerefrfe (102X)
Head of worker of P. greggi n. sp. (102X)
Worker of _P. quiescens n. sp. (45X)
Dorsal view of worker of P. quiescens n. sp.
(103X)
Propodeum, pedicel and gaster of worker of
£. dentigula (102X)


16
16a) Postpetiole without lateral angles, hairs slender and
pointed; postpetiole and gaster smooth and shining.
(fig. 48) dentigula M.
b) Postpetiole with angular sides, seen from above; hairs
short and thick
17a) Dorsal surface of first gastric tergite finely and densily
granulse, opalescent and subopaque; dorsum of pronotum
and mesonotum punctate; dorsum of postpetiole punctate
and opaque. (fig. 50) anastas
b) First gastric tergite smooth and shining; dorsal area of
postpetiole weakly smooth and shining; dorsum of pronotum
usually with partial smooth shining areas.
R. Smith
17
ii Emery
(fig. 51)
floridana Emery


ACKNOWLEDGEMENTS
i
I wish to express my gratitude to Dr. William F. Burn, Advisory
Committee Chairman, who patiently guided me and inspired me during every
phase of this research. I also wish to acknowledge my sincere indebteness
to Dr. George E. Allen, Dr. Francis W. Zettler and Dr. Clifford S. Lofgren,
members of my Advisory Committee, for their constructive criticisms and
guidance.
I sincerely thank Mrs. T. Carlysle for providing me with scanning
electron micrographs.
I also wish to thank Dr. John F. Carroll, Dr. J. C. E. Nickerson, and
Dr. D. P. Wojcik, for continuing assistance, and to Dr. Edward 0. Wilson
and Dr. David R. Smith for their permission to study the collections at
the Museum of Comparative Zoology and U.S. National Museum, respectively.
This research was also aided in part by Tall Timbers Research Station,
Leon Co., Florida, through the kindness of Dr. E. V. Komarek, Director of
the Station, and the sponsorship of Dr. Willard H. Whitcomb. The support
of Empresa Brasileira de Pesquisa Agropecuaria is also acknowledged with
gratitude.
To my wife, Lucilia, my utmost thanks and appreciation for her patience
iii
and support.