Group Title: revision of the genus Paratrechina (Hymenoptera : Formicidae) of the continental United States
Title: A revision of the genus Paratrechina (Hymenoptera : Formicidae) of the continental United States
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Title: A revision of the genus Paratrechina (Hymenoptera : Formicidae) of the continental United States
Physical Description: xiii, 263 leaves : ill. ; 28 cm.
Language: English
Creator: Trager, James C., 1952-
Copyright Date: 1984
Subject: Entomology and Nematology thesis Ph. D
Dissertations, Academic -- Entomology and Nematology -- UF
Genre: bibliography   ( marcgt )
non-fiction   ( marcgt )
Statement of Responsibility: by James C. Trager.
Thesis: Thesis (Ph. D.)--University of Florida, 1984.
Bibliography: Includes bibliographical references (leaves 231-235).
General Note: Typescript.
General Note: Vita.
General Note: Also published as Sociobiology, v.9 no.2, 1984.
 Record Information
Bibliographic ID: UF00099350
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: alephbibnum - 000473011
oclc - 30754829
notis - ACN7942


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According to the International Code of Zoological

Nomenclature (Art. 9), dissertations submitted in partial

fulfillment of requirements for doctoral, master's or other

degrees do not constitute publications in the nomenclatural

sense, even when reproduced on microcards, by microfilm

or other similar methods. Therefore, the novel names or

name-combinations proposed in this dissertation are not

entered into the public domain. The information given in

this work is in effect confidential and cannot ethically be

utilized by anyone in any publication (Art. 8 of Code) until

the author of the dissertation has published it in a legit-

imate outlet, unless the author or the author's adviser

gives permission for someone else to do so, or unless after

a reasonable interval neither can be consulted even with

concerted effort (in conformance with the Code of Ethics,

Appendix A of the International Code of Zoological Nomen-


Protection under the articles and appendices of the

Code is hereby claimed for this dissertation.

I dedicate this study to the memory of Dr. William

F. Buren, who first suggested the subject and gave me the

initial nudging necessary to get it underway. It is my

hope that this will constitute a suitable tribute to his

contributions to ant taxonomy.


As in all such studies, this work could never have

been brought to its present state without the aid of many

people. Foremost among these is my wife, Kim, whose unfail-

ing love and devotion and assistance with the minutiae of

curation and preparation of the manuscript have been immense-

ly important and are thoroughly appreciated. I give heart-

felt thanks to the late Dr. William F. Buren and to Dr.

Jerry L. Stimac for their many discussions with me on this

study and related matters, for their friendship and en-

couragement, and not the least of all for arranging finan-

cial support during much of the course of the work. I

thank Dr. Daniel P. Wojcik and Mr. Ruediger W. Klein for

making extra efforts to collect Paratrechina for me. I

wish to thank Dr. William L. Brown, Jr., and Mr. Roy R.

Selling for their indirect guidance through the example

of their work in any taxonomy and for commenting on early

drafts of the work. For typing the earlier versions, I

thank Anne Keene and Renee Lofgren. For assistance with

the computer program and data input, I thank Dr. Robert J.

O'Neil, Lois Wood and Joan Denicola. Finally, I thank all

the institutions and persons listed in Chapter III for their

cooperation in lending or donating specimens, including

valuable types, without which the study could never have

been carried out.



ACKNOWLEDGMENTS ......................................... iv

LIST OF TABLES ........................................viii

LIST OF FIGURES ......................................... ix

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


I INTRODUCTION ..................................... 1



IV SYNONYMY OF THE SUBGENERA OF Paratrechina ...... 13

V GENERIC DESCRIPTION .............................. 17

Worker ......................................... 17
Queen .......................................... 22
Male .......................................... 23

STATES ......................................... 24

Preface to Keys ................................ 24
Key to Workers ................................. 28
Key to Males ................................... 35


VIII VIVIDULA COMPLEX ................................. 46

Diagnosis of Complex ............................ 46
Paratrechina vividula ....................... 46
Paratrechina terricola ...................... 56

IX PARVULA COMPLEX ................................... 74

Diagnosis of Complex ............................ 74
Paratrechina concinna, New Species ........... 74


Paratrechina faisonensis .. .................. 82
Paratrechina flavipes ....................... 92
Paratrechina parvula ......................... 103
Paratrechina wojciki, New Species ............113
Paratrechina austroccidua, New Species ....... 122

X ARENIVAGA COMPLEX ............................. 134

Diagnosis of Complex ......................... 134
Paratrechina arenivaga ...... ................... 134
Paratrechina phantasma, New Species .......... 146

XI BRUESII COMPLEX ............................... 156

Diagnosis of Complex ........ ................. 156
Paratrechina bruesii ......................... 156

XII HYSTRIX COMPLEX ............................... 166

Diagnosis of Complex ......................... 166
Paratrechina hystrix, New Species ........... 166

XIII GUATEMALENSIS COMPLEX ...... ................... 178

Diagnosis of Complex ......................... 178
Paratrechina guatemalensis .................. 178

XIV FULVA COMPLEX ................................. 187

Diagnosis of Complex ......................... 187
Paratrechina pubens ........ ................. 187

XV BOURBONICA COMPLEX ............................. 194

Diagnosis of Complex ......................... 194
Paratrechina bourbonica ..................... 194

XVI LONGICORNIS COMPLEX ........................... 206

Diagnosis of Complex ......................... 206
Paratrechina longicornis ....................... 206

BIBLIOGRAPHY ................ .......................... 231


IN THIS REVISION .............................. 237

APPENDIX A ....................................... 253

BIOGRAPHICAL SKETCH ................................... 263



Table Page

1 Reference collections of Paratrechina,
abbreviations for the collections used in
the text, and individuals responsible for
the loan or gift of specimens ................ 8

2 Synopsis of taxonomy and authorship of
U.S. Paratrechina ............................. 27

3 Selected characteristics of members of the
Guatemalensis Complex of Paratrechina ........ 185


Figure Page

1 Cephalic microsculpture of longicornis
worker ........................................ 14

2 Cephalic macrochaeta of faisonensis male ..... 18

3 Abdominal macrochaeta and sculpture of
faisonensis male .............................. 21

4 Head of vividula worker, dorsal view ......... 71

5 Head of terricola worker, dorsal view ........ 71

6 Head of terricola male, dorsal view .......... 71

7 Male terricola, lateral view .............. ..... 73

8 Male vividula, lateral view .................. 73

9 Worker of terricola, lateral view ............ 73

10 Worker of vividula, lateral view ............. 73

11 Worker of concinna, lateral view ............. 102

12 Worker of faisonensis, lateral view .......... 102

13 Worker of flavipes, lateral view ............. 102

14 Head of concinna worker, dorsal view ......... 102

15 Head of faisonensis worker, dorsal view ...... 102

16 Head of flavipes worker, dorsal view ......... 102

17 Worker of austroccidua, lateral view ......... 133

18 Worker of wojciki, lateral view .............. 133

19 Worker of parvula, lateral view .............. 133

20 Head of austroccidua worker, dorsal view ..... 133

21 Head of wojciki worker, dorsal view .......... 133

22 Head of parvula worker, dorsal view .......... 133

23 Worker of arenivaga, lateral view ............ 155

24 Worker of phantasma, lateral view ............ 155

25 Head of arenivaga worker, dorsal view ........ 155

26 Head of phantasma worker, dorsal view ........ 155

27 Worker of bruesii, lateral view .............. 175

28 Worker of hystrix, lateral view .............. 175

29 Head of bruesii worker, dorsal view .......... 175

30 Head of hystrix worker, dorsal view .......... 175

31 Worker of bourbonica, lateral view ........... 216

32 Worker of longicornis, lateral view .......... 216

33 Worker of pubens, lateral view ............... 216

34 Worker of guatemalensis, lateral view ........ 216

35 Head of guatemalensis worker, dorsal view .... 218

36 Head of pubens worker, dorsal view ........... 218

37 Head of bourbonica worker, dorsal view ....... 218

38 Head of longicornis worker, dorsal view ...... 218

39 Male of bruesii, lateral view ................ 177

40 Male of hystrix, lateral view ................ 177

41 Male of concinna, lateral view ............... 220

42 Male of longicornis, lateral view ............ 220

43 Male of bourbonica, lateral view ............. 220

44 Male of pubens, lateral view ................ 220

45 Genitalia of faisonensis male, dorsal view,
slide mounted ......... ......................... 222

46 Genitalia of guatemalensis male, dorsal view,
slide mounted .................................. 222

47 Genitalia of arenivaga male, dorsal view,
slide mounted .................................. 222

48 Genitalia of phantasma male, dorsal view,
slide mounted ........................ .......... 222

49 Distribution in North America of vividula
( ), terricola ( o ), guatemalensis (A ) and
pubens ( ) .................................. 224

50 Distribution in North America of concinna
(o ), faisonensis ( ), flavipes ( ), hystrix
(A ) and bruesii ( ) ......................... 226

51 Distribution in North America of parvula (e ),
wojciki (o ) and austroccidua (A ) ........... 228

52 Distribution in North America of arenivaga
(* ), phantasma ( o ) and bourbonica ( n ) ..... 230

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




August 1984

Chairman: Jerry L. Stimac
Major Department: Department of Entomology and Nematology

The taxonomy of the ant genus Paratrechina in the con-

tenental United States is revised. Identity of previously

published taxa is determined, and the workers and males are

redescribed. The workers and males of the new species P.

concinna, P. phantasma and P. wojciki from the Southeast and

P. austroccidua and P. hystrix from the Southwest are de-

scribed. P. fulva pubens, P. melanderi arenivaga and P.

arenivaga faisonensis are raised to full species. P. par-

vula grandula is transferred to Conomyrma. The establishment

of P. flavipes of temperate eastern Asia, in Pennsylvania

and New York, and of P. guatemalensis of the Caribbean

region, in southern Florida, is reported for the first time.

The synonomy of P. kincaidi with P. vividula and the sibling

species relationship of vividula to P. terricola are

discussed as are the reasons for resurrecting Buckley's name

terricola for the species described by Wheeler as melanderi.

Biological and distributional data for each species are also

presented. Keys are given for the workers and males of all




Paratrechina is a common though often inconspicuous

element of the ant faunas of almost all continental areas

habitable by ants. Native species are lacking or nearly so

from Europe (Brown, 1973) and colder temperate, Mediterra-

nean climate and desert regions of other continents. The

genus is most diverse in tropical Asia and Australia, which

together have the greatest number of species groups, includ-

ing one-third of the described forms (Emery, 1925). Based

on my acquaintance with the New World forms, I estimate that

perhaps three-fourths of the true number of species has been

described. For the Old World tropics, the proportion may

be much smaller.

A number of species have been transported by shipping

activity to localities well beyond their places of origin.

Some of these species have attained at least minor pest

status through their invasion of nurseries, greenhouses,

laboratories, insectaries, homes, and other buildings (Smith,

1965). Foremost among such species in the New World are

longicornis, vividula, bourbonica, and members of the fulva

group. Other species have been transported by man and become

established in new localities, but the taxonomy of the genus

is so poorly known at present that it would be impossible

to state with certainty which species are involved. Wilson

and Taylor (1967) discussed the taxonomic difficulties in

Paratrechina with special reference to Polynesia.

This study is the first attempt to monograph Paratre-

china and constitutes Part I of a worldwide revision. I

expect that this will lend a degree of clarity to a group

for which the taxonomy "looks very much like a hopeless

muddle" (Creighton, 1950). Apart from the purely academic

interest in unravelling the taxonomic tangle of this diffi-

cult genus, it is hoped that the keys and descriptions will

provide nursery and quarantine inspectors and other applied

entomologists a means to readily identify the species of

this frequently encountered genus.

In the next chapter, I review the nomenclatural history

of Paratrechina and discuss what is known about its rela-

tionships to other ant genera. In the third chapter, I

describe the methods and equipment used in examining the

specimens studied and give a complete listing of abbrevia-

tions used in the text. In Chapter IV, I tentatively synony-

mize the subgenera of Paratrechina and give my reasons for

doing so. In Chapter V, I describe the workers, males, and

queens of Paratrechina at the generic level. Chapter VI

contains a list of the U.S. species of Paratrechina and

those introduced to the region, as well as their authors and

known synonyms. Also in Chapter VI are keys to workers and


males of these species. Chapter VII establishes the format

of the treatment of each species and includes my reasons for

the use of the term "complex" for groups of related species.

The remainder of the chapters contain the descriptions of

the species of each complex. A brief diagnosis of the com-

plex treated is at the beginning of each of these chapters.


The name Paratrechina was first used in the combina-

tion P. currens Motschulsky (1863). The species name was

not used for over sixty years, until Emery (1925) published

his treatise on the formicine genera. In the interim, the

great majority of species known today were described and

were assigned to Prenolepis Mayr (1862), as was Formica

longicornis Latreille (1802), the senior synonym of

Motschulsky's species.

In 1906, Emery published a note, unusual for its time

in that it truly made clear the identity of vividula and

pointed out its probable place of origin and close rela-

tionship to terricola melanderii), shown below to be its

sibling species. More germane to the immediate discussion

is that in his note, Emery erected the subgenera Prenoleois

s.s., Euprenolepis and Nylanderia. The first two of these

are today considered distinct genera. Euprenolepis is not

in the scope of this paper. Prenolepis and Nylanderia are

discussed presently.

Prenolepis s.s. was distinguished by Emery in that all

castes lack erect pilosity on the scapes and tibiae and

instead possess long, dense, oblique (i.e., subdecumbent)


pubescence on these segments. In workers, the mesothorax

is extremely constricted, such that it is less than half as

broad as the pronotum in dorsal view. The pronotum and pro-

podeum are broad and globular, yielding in Prenolepis workers

a "dumbell-shaped" mesosoma. Later, Emery (1925) separated

Prenolepis s.s. as a full genus and added to its distin-

guishing features the placement of the eyes posterior to the

middle of the head and the possession of cerci by the males.

In North America, at least, the short scape of the males of

Prenolepis (about equal to or less than length of head) and

the size of the females (10 or more mm vs. 5 mm or less)

further distinguish it from our species of Paratrechina.

Nylanderia was construed by Emery in 1906 to contain

all of what are now called Paratrechina. In 1925, Emery

established the genus Paratrechina with Paratrechina and

Nylanderia as subgenera, containing longicornis and the

remainder of the species, respectively. Wheeler (1936)

elevated Nylanderia to generic level. Though Donisthorpe

(1943) and Kempf (1972) followed this arrangement in major

taxonomic publications, most other authors have used Emery's

(1925) arrangement. Donisthorpe (1947) proposed a third

subgenus, Paraparatrechina, to include his New Guinean

species pallida. Brown (1973) and Snelling (1981) listed

Nylanderia as a synonym of Paratrechina without discussion.

The validity of the subgenera is discussed in Chapter V.

Paratrechina is said to be most closely related to

Prenolepis, Pseudolasius, Lasius and Formica (Wilson,

1975)--to the last two certainly more distantly than to

the first. Unfortunately, our knowledge of Pseudolasius is

insufficient to state the degree of relation to Paratre-

china. Future investigation should be directed toward

reconstruction of the phylogeny of these apparently related

genera and the rest of the Formicinae.

The relationship of Paratrechina and Prenolepis is

clear. Together they form a monophyletic group. The larvae

of these genera share a plump, dolichoderoid appearance,

indistinct segmentation, lack of a "neck," and relatively

few and short head hairs (G.C. and J. Wheeler, 1953, 1968).

These last authors place the two genera in Forel's tribe

Prenolepidini. (The larvae of Brachymyrmex appear related

to the Prenolepidini, but other characters suggest differ-

ently and the larval similarities may be due to conver-

gence.) Both prenolepidine genera have nude pupae, an

unusual trait in Formicinae. In addition, the workers show

similar reduced sculpture and restricted patterns of

thoracic pilosity (and often of pubescence). The single-

toothed male mandibles and 5- or 6-toothed female mandibles

of the two are virtually indistinguishable, as is the

structure of their proventriculi (Eisner, 1957). Workers

of both groups have a reduced mesothorax, a trait much

further developed in Prenolepis. Finally, both groups con-

tain, with well-known exceptions, mostly smallish, soil or


litter-dwelling inhabitants of more or less undisturbed

mesic areas. All known species in this pair of genera can

readily be placed in one genus or the other.


About one-third of the specimens studied were collected

by myself in Florida, primarily in the vicinity of Gaines-

ville and at Archbold Biological Station in Highlands County.

The remainder of the specimens were obtained by loan or gift

from the collections listed below, thanks to the cooperation

of the individuals listed beside each collection (Table 1).

Table 1. Reference collections of Paratrechina, abbrevia-
tions for the collections used in the text, and
individuals responsible for the loan or gift of

American Museum of Natural

British Museum (Natural

Private Collection

California Academy of Science

Frost Entomological Museum

Florida State Collection of

Helsingfors Zoological Museum

Private Collection

Private Collection

M. Favreau

B. Bolton






& J. Wheeler








Table 1--continued














Private Collection

Natural History Museum of
Los Angeles County

Harvard Museum of Comparative

Museum d'Histoire Naturelle,

North Carolina State

Private Collection

Private Collection

Private Collection

Private Collection

Private Collection

State Biological Survey of
Kansas and Private Collection

Private Collection

Texas Tech University
(Collection funded by Texas
Department of Agriculture)

United States National Museum

University of Arkansas

USDA Fire Ant Project
Collection and Private

Private Collection

Private Collection

A. Newton

C. Besuchet

C. Parron

P. Ward

R. Gregg

R. Lavigne

R. Chew

R. Klein

M. DuBois

T. Nuhn

0. Francke

D. Smith

C. Carlton

D. Wojcik

W. Buren

W. McKay









Point-mounted specimens were measured at 50X with an

ocular micrometer. A computer program was used to convert

the measurements from micrometer units to the nearest 0.01 mm

equivalent, to calculate indices of relationship between the

length of certain body parts and to calculate summary sta-

tistics. Only summarized, rounded-off, data are presented

in the text, but the full raw data set has been reproduced

in Appendix A. All measurements are reported in mm.

Measurements are counts taken were as follows:

TL--HL + WL + GL (see below)

HL--Head length in full face view from a line perpen-

dicular to sagittal axis and tangent to the pos-

teriormost parts of the rear border to such a line

tangent to the anterior most parts of the clypeal


HW--Maximum width of head (including eyes) in full face


EL--Maximum diameter of compound eye

SL--Length of scape (chord if curved) from base (exclu-

sive of basal radicle) to its terminus

PW--Maximum width of pronotum

WL--Weber's length of thorax, from anterior edge of

pronotum (exclusive of anterior flange) to posterior

corner of metapleuron

MCL--Length (chord if curved) of largest pronotal macro-


FL--Length of fore femur

GL--Length of gaster in dorsal view (exclusive of cone

of acidopore in worker or of genitalia in males)

SM--Number of standing barbulate macrochaetae on scape

PM--Number of barbulate macrochaetae on pronotum to

one side of sagittal plane

MM--Number of barbulate macrochaetae on mesonotum to

one side of sagittal plane

CI = x 100

01 = E- x 100

SI = |- x 100


FI = x 100

Illustrations were prepared with the aid of a Nikon

SMZ10 stereo dissecting microscope with drawing tube attach-

ment. All are diagrammatic and are modelled on more than

one specimen; proportions and macrochaetal counts were drawn

at or near their mean values for the species. The pubescence

patterns indicated on the head drawings are actual setal maps

of the middorsal portion of the head of a specimen chosen

for its "typical" pubescence.

Slide preparations of genitalia prepared by Dr. Arnold

Van Pelt, loaned by USNM, were consulted in preparation of

the drawings of the genitalia of faisonensis and arenivaga.


The drawings of the genitalia of guatemalensis and phantasma

are based on my own dissections but were positioned as

though based on slide mounts.


As has been shown by Wilson (1955) and Snelling (1976),

subgenera may be useful taxonomic tools when they represent

clearcut phyletic trends comprising major monophyletic group-

ings of species within a genus. It seems clear to me that

the nominate subgenera of Paratrechina do not live up to

this criterion. Indeed, some of the species groups described

in 1925 by Emery such as his caledonica group, and microps

and its relatives in the Caribbean region come closer to

meeting it, but it would be premature to describe subgenera

for such groups without a better knowledge of the world fauna.

I have studied workers of longicornis, pallida and

cisipa to determine their relationship and to determine the

validity of the subgenera. Longicornis is an unusual mem-

ber of the genus in many respects, including its large eyes,

great elongation of the scapes and tibiae, reduction of pil-

osity, unusual microsculpture (see Fig. 1 and description)

and unique biology. Pallida appears related to longicornis.

Pallida, too, has 5-tooth mandibles, long scapes and tibiae

and even longer maxillary palps, further reduced pilosity

(except that there are two pairs of macrochaetae on the

propodeum not found on longicornis), fine very short



Fig. 1. Cephalic microsculpture of
longicornis worker.


pubescence, and slight bluish reflections. The thorax of

the two species is of virtually identical conformation.

The single type specimen of pallida available to me could

not be subjected to study in the SEM to determine micro-

sculptural details, but the macroscopic appearance of the

integument and the thick, blunt appearance of the macro-

chaetae are similar to those of longicornis.

Smith and Lavigne (1973) described cisipa from Puerto

Rico, noting that its elongated, flattened thorax and long

legs and scapes resembled those of longicornis but that its

6-toothed mandibles and bristly scapes forced them to place

cisipa in Nylanderia. In fact, cisipa, while not related

to longicornis, is not close to any other member of Paratre-

china either. Some cisipa workers I have seen (RJL, RWK),

including one in the type colony, have 5-toothed mandibles.

The bristles on the scapes and tibiae are not the usual

barbulate macrochaetae but are in fact thickened, oblique

pubescence very much like that of Prenolepis. Interestingly,

the eyes of cisipa are placed further back on the head than

usual for a Paratrechina (but not as far back as in Prenol-

epis), and the mesothorax is quite constricted. On the other

hand, the lack of cerci on the male and the pattern of

thoracic pilosity on the worker of cisipa are typical of

Paratrechina. The microsculpture and nature of the barbula-

tion of the macrochaetae are as found in "Nylanderia," but

this species cannot otherwise be to closely linked to that

grouping of species.


Thus, longicornis and pallida appear to form a real

species-group but hardly constitute a major phyletic trend.

Nylanderia is a paraphyletic group including all the re-

maining diversity within the genus, including some species-

groups at least as removed phenetically from the mainstream

of the genus as are longicornis and pallida. I, therefore,

consider the subgenera Paraparatrechina and Nylanderia

synonymous with Paratrechina and recommend that the names

no longer be used. The only feasible alternative is a more

restricted Nylanderia and the erection of several new sub-

genera, but this must await further study.


The males and workers of Paratrechina have been

characterized by M.R. Smith (1943, 1947, 1965). Some

important points from his diagnoses are included below.

In addition, characters not discussed by him and a de-

scription of the female are included.


Small, 1.5-3.5 mm (4 mm in one Australian form). An-

tenna 12-segmented, arising from a fossa placed close to but

not confluent with rear border of clypeus; scape longer than

head, usually with two to four rows of subdecmebent to erect

barbulate macrochaetae (as in Fig. 2) (lacking in Darvula,

opaca, longicornis, etc.); funiculus filiform to very

weakly clavate, segments longer than broad, terminal one

almost twice as long as the others. Eye typically well-

developed, about one/fifth or more as long as head (much

smaller in subterranean microps group of Caribbean region,

caledonica, etc.). Ocelli usually indistinct, or absent;

often only median ocellus is visible. Mandible striate 6-

or 5-toothed; typical dentition as in vividula (Fig. 4)

(Smith, 1939, was in error in stating that myops had four

Fig. 2. Cephalic macrochaeta of faisonensis


mandibular teeth.) Palpal formula 6, 4; palpal segments most

often subequal in length. Clypeus bears 6-20 slender, barbed

macrochaetae of varying lengths and a smoothly rounded median

longitudinal angle. Clypeus emarginate or evenly rounded

anteriorly; in side view with a declinate anterior face.

Frontal carinae very short, extending little beyond antennal

fossae, thus, the frontal area often indicated only by its

possession of four longitudinal rows of regularly spaced

thick, barbulate macrochaetae (the inner two, often, and the

outer rows, occasionally, reduced to a pair or so of

bristles); the rows extending back through the ocellar area

to the rear border. Sides of head in full face view nearly

parallel, or more often, rounded and weakly convergent

anteriad; rarely broader before than behind the eyes. Sides

of head with macrochaetae shorter than those on preocellar

area, subdecumbent and often greatly reduced in number.

Posterior border straight or rounded, often weakly emarginate

medially, with somewhat longer, more curved, erect pilosity

than on sides. Venter of head with fine flexuous pilosity

similar to that on clypeus, or none.

Pronotum convex or subangular. Mesonotum usually nearly

flat in profile, the latter with a very short anterior and

posterior face in many species such that the flat dorsal

surface is slightly raised above pronotum, at least anterior-

ly. Mesonotum somewhat reduced, about 1/2 to 3/5 as broad

as pronotum seen from above. Metanotum a narrow band usually

in a distinct furrow, bordered laterally by mesothoracic

spiracles which are nearly dorsal in position and orienta-

tion of their openings. Pronotum and mesonotum each with

two pairs of large macrochaetae like those on head (Fig. 2)

and usually one or more (rarely up to 16) pairs of smaller

macrochaetae on pronotum and usually one or 2 (up to 8)

such ancillary pairs on mesonotum.

Propodeum weakly angular with a short, low, dorsal face,

or convex and of varying height relative to thoracic dorsum.

Propodeum without pilosity except in caledonica group, and in

glabra, dugasi (Emery, 1925), microps, hystrix and pallida.

Some caledonica-group species may have only one pair of

macrochaetae on mesonotum. Petiole cuneate with crest vary-

ing from sharp to truncate or rounded in profile; in dorsal

view, crest flat, angular or convex, only rarely notched in

workers (e.g., sakurae). Petiole glabrous or with one or a

few short erect macrochaetae on the crest.

Gaster ovate, acuminate terminally, relatively volumi-

nous, with more or less abundant flexuous pilosity with

less barbs per macrochaetae (Fig. 3) than on those of thorax.

Anterior face of gastral tergite 1 impressed and overhang-

ing petiole, concealing it partially from dorsal view. Bor-

der of impression distinctly angular. Acidopore conspicuous

and conical, with a distinct corona.

Pubescence typically dense on the head and appendages,

shorter and more dilute or lacking on much of thorax and

gaster, the patterns of vestiture forming important diag-

nostic characters of some species or species-groups.


Fig. 3. Abdominal macrochaeta and sculpture
of faisonensis male.

Sculpture usually limited to shallow pubigerous

foveolation on head, delicate shagreening on gaster, and

honeycomb-like reticulation on terminal segments. Some

species in caledonica group are heavily punctate, opaque

forms which contrast strongly with the typical shining

appearance (when not obscured by pubescence) of other



In general, fitting the worker description but dif-

fering as follows. TL 4-7 mm, scape relatively shorter and

with less pilosity than in conspecific workers. Eye a

little larger to much larger in subterranean forms, 1/4 or

more HL. Ocelli fully developed.

Thorax of normal alate formicine type. Pronotum and

other dorsal surfaces invested with barbulate macrochaetae

like those found on workers but not in so regular an array.

Propodeum with relatively shorter dorsal face than in

worker. Petiole typically broader, sharper in profile, and

often with at least a weak median notch.

Gaster much more massive and more thoroughly hiding

petiole in its impressed anterior face.

Pubescence longer, thicker and more dense, usually

covering entire body, except petiole and rear face of



With the important generic characters noted for

females, i.e., barbulate pilosity; the impressed first

gastric tergite and antennal scapes longer than head. In

overall size, approximating that of conspecific worker.

Mandibles well-developed, though smaller and less

heavily sclerotized than in workers; with a single apical

tooth and occasionally a small adjacent cleft, and one or

more inconspicuous subapical denticles. Antennal scapes

with less macrochaetae than conspecific workers or queens.

Antennae 13-segmented, with conformation of segments

generally resembling that of female castes.

Petiole of same conformation as in workers but broader

and more blunt, sometimes notched like that of queen.

External genitalia prominent, varying greatly between

species-groups and interspecifically and, thus, useful in

taxonomy. Cerci (as in Prenolepis) lacking.

Pubescence distributed as in queens, but in length and

density more like that of workers. Thoracic pilosity

usually limited to mesometanotum, rarely more generally

distributed; typically sparse and decumbent on gastral



Preface to Keys

In this chapter, I provide what will undoubtedly be the

most frequently read portion of this study, namely the keys

for identification of the workers and males of Paratrechina

in the United States. Also included is a taxonomic synposis

of the species included in the keys and formally treated in

the later chapters (see Table 2).

The key contains a number of species which occur out-

side of the United States but in many cases cannot be used

to identify them outside of this country. There are many

Paratrechina species not treated in this revision, which

makes the use of these keys futile outside of the geographic

region for which they are designed.

The characters which separate Paratrechina species are

more subtle than those used to separate species in some

other insect groups. One will not find here the simple,

one-feature alternatives of older keys to Paratrechina. My

keys assume the user has possession of a stereo dissecting

microscope fitted with a calibrated ocular micrometer and

of a calculator to expedite the calculation of indices.

The reader is urged to read carefully the definitions of

the measurements in Chapter III and the following remarks

on possible difficulties one may encounter with certain

measurements and counts before attempting to use the keys.

Remarks. Certain difficulties were encountered in

taking some of the measurements, and the following comments

may help the user of the keys in selecting the appropriate

alternative at a given couplet (and in reading the descrip-


The posteroventral portion of the head is excised and,

thus, in certain positions obscures the front of the pronotum

making it difficult or impossible to measure WL from the

side. In such cases, WL was taken from a dorsal view. As

a check, the measurement was taken on a number of specimens

in which WL was visible from both views. The two measure-

ments never differed by more than 0.01 mm and usually by

no more than 0.005 mm.

Macrochaeta counts proved to have certain unforeseen pit-

falls. Only barbulate setae of the sort shown in Figs. 2 and

3 were counted. These were usually discerned by their great-

er length and diameter and their different color (usually

darker than pubescence and body color), but in some species

the macrochaetae are nearly the color of the substrate and are

rather short. Compounding the problem in such cases is that

there may occur simple setae of nearly the same macroscopic

appearance. Through observation at high magnification and

checking back at lower power, I learned to discriminate

these setal types. The discrimination turned out to have

a slightly different "Gestalt" for each of the "difficult"

species. In addition, the macrochaetae are all visible only

at certain angles of viewing and illumination or where SM is

large, only by rotating the scape. Though this particular

count is undoubtedly the one for which the most errors were

made, I feel confident that the high end of the ranges re-

ported are close to correct. The count recorded was the

highest repeatable one from among either of the two scapes

resulting in an inherent bias against low counts, whether

natural or due to damage. The scapes of a single specimen

often differed in SM, even on fresh, undamaged specimens.

PM and MM also proved to be asymmetrical in many spec-

imens. Always present (or at least indicated by a socket)

were the 8 largest hairs (the "major thoracic macrochaetae"),

but the number of ancillaries might be different on each

side. The highest repeatable count of hairs and/or unmis-

takable hair-sockets was the one recorded. On occasion,

smaller barbulate setae resembled pubescence, but I learned

to discriminate these as above. The discrimination was

much easier on pale species with dark macrochaetae, and

most counting errors were certainly made on dark species.

I refer loosely to the middle tagma of ants as the

thorax which is the current usage by many hymenopterists

for the homologous body region of ants and other Hymenop-

tera. In practice, there is little need to refer specifi-

cally to the thorax and propodeum collectively, for which

more accurate terms would be alitrunk or mesosoma.

Table 2. Synopsis of taxonomy and authorship of
U.S. Paratrechina.

Vividula Complex
vividula (Nylander)
--kincaidi (Wheeler) syn. n.
terricola (Buckley) stat. rev.
--melanderi (Wheeler) syn. n.

Parvula Complex
concinna sp. n.
faisonensis (Forel)
flavipes (F. Smith)
parvula (Mayr)
wojciki sp. n.
austroccidua sp. n.

Arenivaga Complex
arenivaga (Wheeler)
phantasma sp. n.

Bruessi Complex
bruesii (Wheeler)

Hystrix Complex
hystrix sp. n.

Section Guatemalensis
guatemalensis (Forel)

Section Fulva
pubens (Forel) stat. n.

stat. n. & rev.

stat. rev.

& rev.

Section Bourbonica
bourbonica (Forel)

Section Longicornis
longicornis (Latreille)

Excluded from Paratrechina
parvula grandula (Forel)
--becomes Conomvrma grandula (Forel) comb. n.

Pilosity refers to the large, standing, barbulate setae

(macrochaetae) characteristic of this group (though by no

means exclusive to it). The unbarbed macrochaetae on the

parameres of males are also referred to as pilosity. Simple,

appressed to subdecumbent setae of small diameter are referred

to as Dubescence. Otherwise, terminology used here is that

which has become standard in recent revisions of formicine

genera (Wilson, 1955; Wing, 1968; Bolton, 1973; Snelling,


Key to Workers

la. Species of eastern U.S. and Great

Plains, including pains and woodlands

of east Texas ....................................... 2

lb. Species of Southwest including west

Texas mountains and deserts ....................... 14

2a. Scapes and legs unusually long (SI>165,

FI>115); weakly shining black or gray

with bluish reflections; with sparse,

short, barely visible pubescence; near

human dwellings, Gulf Coast states,

urban areas, sporadic elsewhere .......... longicornis

2b. Scapes and legs of usual proportions

(SI<130, FI<105); colored variously

but never with bluish reflections;

shiny, or if dull, this due to dense

pubescence; habitat various ........................ 3

3a. Yellow to pale whitish; nests in sandy

soil or dunes with entrances surrounded

by conspicuous crater of subsoil in

clearings between vegetation ...................... 14

3b. Uniformly dark-colored or bicolored;

nests inconspicuous; in more mesic

microhabitats under moss, rocks, logs,

in litter; or near man-made structures

(parvula may have crater nests in sandy

areas but is never uniformly yellow) ............... 5

4a. Yellow with gaster infuscated posteri-

orly; thoracic pilosity flexuous and

dark brown (notably darker than body

color); scapes with 5-17 (usually 7-12)

macrochaetae and suberect pubescence;

New Jersey, Florida, Gulf Coast, sand-

hills of Midwest .......................

4b. Yellow or whitish with gaster, at most,

only slightly darker, thoracic pilosity

nearly straight, nearly the same color

or only slightly darker than body color;

scapes with 0-4 (usually 1-3) standing

macrochaetae and short, appressed

pubescence (Florida scrub and dunes) ...

5a. Scapes with not more than 4 standing

macrochaetae ...........................

5b. Scapes with at least 4 (usually 7 or

more) standing macrochaetae ............

.... arenivaga

.... phantasma

............ 6

......... ... 7

6a. Typically bicolored, with thorax

yellowish to reddish-brown, head and

gaster darker; middle and hind coxae

pale, much lighter than fore coxae or

rest of legs; scapes with 1-4 suberect

macrochaetae; in full face view eyes

reaching sides of head or failing to do

so by 1 or 2 facet-widths; small (HL

usually 0.51-0.57); (Florida) ................ wojciki

6b. Body typically uniform dark brown (at

most weakly bicolored); appendages

somewhat lighter or even yellowish;

scapes never with standing macrochaetae;

in full face view outer edge of eyes

failing to reach sides of head by 1/4

of eye width; larger (HL>0.57, usually

>0.60) (eastern U.S.) ........................ parvula

7a. Thorax and gaster mostly covered with

pubescence or lacking it only on pleura

and disk of pronotum; dull ......................... 8

7b. Thorax and gaster with greatly reduced

pubescence; shiny ................................. 10

8a. Body light reddish-brown with slender

flexuous light brown pilosity (Fig. 33) ....... pubens

8b. Body dull brown to nearly black or if

faded, pilosity shorter, stouter,

straighter, and dark (Figs. 31 and 34) ............. 9

9a. Uniform dark brown or black, thoracic

macrochaetae thick, often nearly

straight, relatively short and abundant

(PM 5-15); entire body pubescent; large

(HL usually 0.70-0.81); Florida, dis-

turbed habitats, mangrove islands,

occasional along Gulf Coast, in green-

houses, etc., elsewhere .................. bourbonica

9b. Yellowish-brown to dark brown with

middle and hind coxae distinctly

lighter; thoracic macrochaetae more

slender, weakly flexuous and tapering,

longer (at least relatively) and less

abundant (PM 3-7); disk and sometimes

sides of pronotum and pleura glabrous

and shining; smaller (HL usually

0.56-0.66); Homestead, Florida ......... guatemalensis

10a. Uniform dark brown with appendages

somewhat lighter; propodeum with a

dense row of longitudinally aligned

pubescence along anterior edge;

mesonotum and front of pronotum with

at least some dilute pubescence; head

with shallow pubigerous punctae and

dense pubescence which is mostly aligned

with long axis of head; marshes, ditches,

damp pastures, swamp edges; in rotten

wood, cow dung or tussocks; Florida,

Alabama, Georgia, Carolinas ................ concinna

10b. Middle and hind coxae and/or thorax and

legs lighter than gaster and head;

pubescence very sparse or absent from

promesonotum and often from propodeum;

head smooth and shining or irregularly

and weakly punctate beneath pubes-

cence; cephalic pubescence may be very

dilute with some setae markedly not

parallel to long axis of head; not

typically inhabiting marshes or

poorly-drained pastures ........................... 11

lla. Cephalic pubescence dilute; preoccip-

ital area with most spaces between

setae as wide as length of setae or

wider, anterior 1/2 of head (except,

perhaps, frons) lacking pubescence ............... 12

llb. Cephalic pubescence denser; preoccip-

ital area with most spaces between

setae no wider than length of setae,

usually less ................... .................. 13

12a. Eye about 1/4 HL or slightly larger

(OI usually 24-27) inhabiting old fields,

cultivated areas, gardens, disturbed

habitats across southern half of U.S.

or greenhouses, etc., further north;

also in native scrub vegetation in

southern Texas, New Mexico, northern

Mexico ..................................... vividula

12b. Eye smaller (OI 20-24), under stones,

moss clumps or bark in forest openings

and other open or disturbed habitats

in Texas and Plains states, and at

lower elevations in Southwest mountains,

Arkansas and Tennessee ..................... terricola

NOTE: The workers of terricola and
vividula cannot always be
reliably separated on morpho-
logical grounds. See male key
for separatory characters. See
also descriptions of these two

13a. Thorax, legs, antennae yellow; head

averaging broader (CI usually 85-90)

and with rounded sides (Fig. 16);

wooded areas, Pittsburg, Philadelphia,

Long Island ................ ................. flavipes

13b. Uniform brown or with slightly lighter

head and thorax; middle and hind coxae

usually markedly lighter than fore

coxae; head narrower (CI usually 83-87)

and less convex-sided (Fig. 15); mesic

woodlands, southeastern U.S., north

along the Atlantic coastal plain to

southern New Jersey ...................... faisonensis

14a. Scapes and legs very long and thin

(SI>165, FI>115), Gulf Coast, spora-

dic elsewhere ............................ longicornis

14b. Scapes and legs of more usual length

(SI<130, FI<105) ................................. 15

15a. Cephalic pubescence dense, partially

obscuring the sheen of the integument

beneath; the pubescence arising from

fine punctae; weak bluish reflections

often present on head and pronotum;

pronotum angular, with short, steep

anterior face and longer, flattened

or concave dorsal face (Fig. 17);

mid-elevation woodlands, mountains of

Southwest, Mexico ....................... austroccidua

15b. Cephalic pubescence very short, dilute

or absent, the integument strongly

shining and easily seen between the

setae; pronotum convex in profile, or

if angular only weakly so and otherwise

not fitting above description ..................... 16

16a. SM<10, PM<7, HL<0.65

........... .... terricola or vividula, see couplet 12

16b. SM>13, PM>7, HL>0.65 ............................. 17

17a. Brown to dark brown, or if partly yel-

lowish then with notably darker pleura,

propodeum and gaster; SM 13-19,

PM 7-11; desert washes and riparian

woodlands, Mexico and southern parts of

adjacent Texas, New Mexico and Arizona ....... bruesii

17b. Uniform yellow to yellowish brown with

abundant black pilosity; SM 21-29;

PM usually 12-16; deserts of east-

central California, Nevada and Utah .......... hystrix

Key to Males

la. Parameres broad margined (Figs. 42

and 43) .............................................. 2

lb. Parameres triangular or digitiform

(Figs. 39-41, 44) ................................... 3

2a. Scapes long (SI 168-174), 3/5 or more

of their length protruding beyond

posterior margin of head and com-

pletely lacking standing macrochaetae;

parameres entire-margined (Fig. 42);

urban areas, sporadic elsewhere .......... longicornis

2b. Scapes shorter (SI 128-135) a little

over 1/2 of their length protruding

beyond posterior margin of head, bear-

ing 7-10 suberect macrochaetae; para-

meres emarginate (Fig. 43); Florida,

disturbed habitats, mangrove islands,

occasional in greenhouses, etc.,

elsewhere ................................. bourbonica

3a. Scapes with 0-1 standing macrochaetae;

parameres relatively short, subtrian-

gular (Fig. 48); eastern U.S. ...................... 4

3b. Scapes with 4 or more standing macro-

chaetae; or if with less, then either

a western species (see lug 14b), or

an eastern one with elongate, triangu-

lar parameres and with prominent

cuspides (Fig. 47) .................................. 6

4a. Digiti straight, lying close to

aedeagus ............................................ 5

4b. In posterior view digiti curved

laterad distally (Fig. 48); scrubland

and dunes of Florida ....................... phantasma

5a. Small (TL<1.9); usually with 1-2

standing macrochaetae on scape; middle

hind coxae distinctly paler than fore

coxae; peninsular Florida .................... wojciki

5b. Larger (TL>1.9); scape lacking standing

macrochaetae, middle and hind coxae

usually same color as fore coxae;

Midwest and eastern U.S (except south

Florida) .................................... parvula

6a. Parameres elongate, triangular, cuspides

long and prominent, reaching to about

3/4 or more of the length of the aedea-

gus; sandy areas, Midwest, Gulf and

East Coast states .......................... arenivaga

6b. Parameres variously formed; cuspides

not reaching to 3/4 the length of the

aedeagus, smaller and not prominent ................ 7

7a. Antennal segment IV crooked or curved

(Fig. 6); aedeagus in side view rela-

tively slender and tapering but

spatulate (Figs. 7 and 8) .......................... 8

7b. Antennal segment IV straight, aedeagus

in side view tapering to a point, which

may be rounded but is never spatulate .............. 9

8a. Parameres viewed from the side tapering,

triangular, rounded truncate to subacum-

inate, viewed from behind distinctly

curved mesad, aedeagus in the form of a

weakly spatulate narrow triangle which is

notably shorter than the parameres (Fig.

8); southern half of U.S., disturbed

areas, in greenhouses further north ......... vividula

8b. Parameres viewed from the side tapering

rapidly over their basal 1/2, then dig-

itiform over the distal 1/2, from behind

only weakly curved mesad, or straight;

aedeagus with distal portion drawn out

into a thin blade with a broadened up-

curved terminus, extending well beyond

parameres (Fig. 7); Arizona, New Mexico,

Texas, Louisiana, Arkansas, Tennessee,

Central Plains states ...................... terricola

9a. Dark species with at least gaster

nearly black, head and thorax piceous

brown or darker, and appendages dark

brown; larger spp. (HL>0.60) ...................... 10

9b. Lighter in color; or if nearly black,

then with brownish-yellow antennae and

legs, or pale middle and hind coxae,

and overall size is smaller (HL<0.57) ............ 11

10a. Propodeum and gaster elongate (Fig.

39); scapes usually with 7-10 macro-

chaetae; parameres strongly tapering

basally, then slender and digitiform

over most of their length, notably

longer than volsellae and aedeagus but

not concealing the latter entirely in

side view; Mexican border region, south

to Hidalgo, Nayarit, also Baja Cali-

fornia Sur .................................. bruesii

10b. Body of more usual conformation,gaster

short (Fig. 41); scapes with 5-7 macro-

chaetae, parameres triangular, as long

or a little longer than volsellae and

aedeagus and concealing the latter in

side view; Florida, Georgia, Carolinas ...... concinna

lla. Seen from behind, digiti straight and

lying parallel to aedeagus ........................ 12

llb. Seen from behind, digiti notably

curved laterad distally (Fig. 46);

Homestead, Florida (also Antilles,

Central and South America, Cocos

Islands) ............................... uatemalensis

12a. Scapes with 6 or more (usually 8-10)

standing macrochaetae ............................. 10

12b. Scapes with 6 or less (usually 1-5)

standing macrochaetae ............................. 14

13a. Body densely pubescent; pale reddish-

brown; eye large (I040); parameres

with more than 30 long blondish

macrochaetae forming a dense fringe

(Fig. 44); Miami, Florida, in green-

houses further north (also Antilles) .......... pubens

13b. Body nearly free of pubescence; shiny

yellowish-brown; eye smaller (OI<32);

parameres with about 15 brown, slender

macrochaetae of varying lengths which

do not form a dense fringe (Fig. 40) ......... hystrix

14a. Scapes with 4-7 standing macrochaetae;

eastern states ..................................... 15

14b. Scapes with 2-3 standing macrochaetae;

Southwest and Mexican mountains ......... austroccidua

15a. Uniform dark brown to nearly black (less

often) gaster slightly darker than the

blackish-yellow thorax and head;


appendages yellowish-brown; wooded

areas near Philadelphia and Pittsburg,

Pennsylvania, and Long Island .............. flavipes

15b. Thorax and usually the head notably

lighter brown than the dark brown

gaster; middle and hind coxae pale

relative to fore coxae and remainder

of leg segments; woodlands of south-

eastern U.S. north to New Jersey pine

barrens ................................. faisonensis


In the following chapters, I describe the 11 nearctic

Paratrechina species and the five introduced into the region.

I have seen a few series I cannot place, in every case con-

sisting of only a few workers. Thus, it has been possible

for me to identify virtually all nearctic Paratrechina mater-

ial in good condition as belonging to one of the 16 species

treated or, at the very worst, into one of the species com-

plexes (see below). Only workers and males are formally

described. A diagnosis of the queen follows that of the

conspecific male. The queens of most of the native eastern

species are best distinguished by association with workers

or males.

It has become increasingly common in ant systematics

to describe holotypes individually, followed by a section

on variation, as is standard in taxonomic works on non-

social insects. This procedure is followed here although

I believe members of a caste from a single colony are

essentially equivalent as name-bearers for ant species in

view of their close genetic relationship and obvious con-

specificity. There is no known case of a mixed colony of

ants comprised of species which are anything less than

clearly distinct from each other. On the other hand, it

often happens that not all syntypes of earlier authors are

concolonial nor even conspecific. I have designated and

described lectotype workers (and allolectotype males

wherever possible) for species described by earlier authors.

Holotypes and allotypes of the 5 species described as new

are in every case progeny of a single queen to the best of

my ability to determine so. Males and workers of the 3 new

eastern species have been deposited at MCZ, AMNH, USMN, LACM

and FSCA. Material of the 2 new western species is deposited

only at MCZ, USNM and LACM.

It has been stated repeatedly that the male genitalia

offer the best characters for separation of Paratrechina

species (Creighton, 1950; and included references), and

indeed, they are useful. The generally accepted corollary

to this is that Paratrechina workers are barely or not at

all distinguishable. In fact, within species complexes,

some of our species are quite similar in the structure of

the male genitalia while the workers are readily distin-

guished. The males of these species are best distinguished

by non-genitalic characters.

All but 2 of our native species form a rather homo-

geneous group. On a world-wide basis, they appear to belong

to the vividula group of Emery (1925). This group includes

all native eastern U.S. species and 2 western species, plus

a number of tropical forms not treated here. Also in the

group are the introduced flavipes and a few other Asian

forms. The workers of these species share reduced thoracic

and abdominal pubescence (but, except in the Vividula Com-

plex itself, fully pubescent heads); reduced thoracic pil-

osity; somewhat shortened legs and scapes with reduced pil-

osity; and fairly uniform size in the medium range for the

genus. The external male genital apparatus typically in-

cludes isoceles-triangular parameres which are curved mesad

and relatively simply constructed volsellae approximating

the structure of those of faisonensis (Fig. 42).

Since only a small part of Paratrechina is treated here,

no attempt was made to reconstruct a phylogeny for the

species. Even a thorough knowledge of the all extant species

would most likely yield only an approximation of the events

leading to the present taxonomic structure of the genus, in

view of the extinctions that have undoubtedly occurred.

All species are grouped in complexes containing groups

of what I believe are ecological or geographical cognate

species. The names of the complexes are simply the names of

their oldest described species. The term "complex" is used

here to avoid confusion with the "species groups" of Emery

(1925) which are larger and more heterogeneous assemblages

within the genus.

Only when I have seen specimens studied by a previous

author or where his description is sufficiently thorough

and clear to make certain the identification have I included

any reference to a Paratrechina species in the synonymies

presented here. Such a conservative approach undoubtedly

will result in omissions but is necessitated by the follow-

ing conditions: (1) Certain taxa, especially vividula and

fulva have been used as catch-alls for new forms which may

not even be closely related and (2) later authors have

rarely bothered to study types or reliably type-compared

material in making identifications. Under such circum-

stances, synonymies based purely on nomenclatural similarity,

without reference to specimens, were considered unreliable.

I have dispensed with the usual practice of listing

the collection data of all specimens studied. Instead, I

have listed states from which specimens studied originated

in the sections on specimens examined, have described the

preferred altitudes and habitats based on my collecting

experience and information gleaned from labels and the

literature in the sections on natural history, and have

provided range maps. In my opinion, these present as much

of the truly useful information as a complete listing of

localities but in a far more readily assimilable form.

Those wishing to collect Paratrechina would do better to

find them in previously uncollected localities than to re-

duplicate past efforts.

The descriptions follow a format that is standard in

systematic literature on ants. Except where otherwise in-

dicated, the descriptions of the head refer to it in full

face view, as in the figures. Descriptions of the thorax,

on the other hand, refer to it in lateral view. Descrip-

tions of the male genitalia, in general, refer to them as

they appear, intact, on the preserved specimen. The vesti-

ture of vividula is described in greatest detail; vestiture

of other species is briefly described in as much as it dif-

fers from that of vividula.

Measurements of the type speciemns are presented in

the following order: TL, HL, HW, SL, EL, PW, MCL, WL,

FL, GL, SM, PM, MM, CI, OI, SI and FL. These are listed

at the start of each worker description. All measurements,

counts, and indices taken are likewise listed at the start

of the description of the type male, but note that PW,

MCL, FL, PM and MM were not recorded for the males. In

the sections on variation and in composite descriptions,

the ranges of all measurements are reported at the head

of the section. For a complete listing of the measurements

and summary statistics, refer to Appendices A and B,


The section on natural history of the species is nec-

essarily rather short. There is little published on any of

these species, including the tramps, and the observations

reported are primarily my own made in Florida. The term

"disturbed" has been used to characterize habitats of some

species. Mild or light disturbance includes grazing activ-

ity, human foot traffic, or selective clearing of vegetation.

Heavy, disturbance refers to more thorough clearing of vege-

tation, tilling of soil, construction of buildings, and

temporary flooding. It is my hope that this revision will

help other investigators to more thoroughly characterize the

lifeways of Paratrechina species.


Diagnosis of Complex

Worker weakly bicolored, very shiny; cephalic pubes-

cence sparse, limited mostly to rear 1/2 of head; scape

pilosity usually subdecumbent to suberect, sometimes incon-

spicuous, SM usually 7 or 8. Nests in open, often disturbed

habitats, under stones, logs, trash, moss clumps, etc.

Male shiny except thoracic dorsum; antennal segment IV

distinctly bent or curved; parameres triangular; digitus

boomerang-shaped; cuspis short; aedeagal lobes triangular

with a spatulate or otherwise broadened tip.

Paratrechina vividula

(Figs. 4, 8, 10; Map-Fig. 49)

Formica vividula Nylander, 1846, Acta Soc. Fenn. 2:

900. Type loc., Botanical Garden, Helsinfors, Finland.

Tapinoma vividula (sic): F. Smith, 1858, Cat. Hym.

Brit. Mus. 6: 56 (in part ?)

Prenolepis vividula: Mayr, 1861, Europ. Formicid.,

p. 52.

Prenolepis kincaidi Wheeler, 1906, Bull. Amer. Mus.

Nat. Hist. 22: 350. Fig. 1, e, 2, z. NEW SYNONOMY. Type

loc., Bermuda.

Prenolepis vividula: Emery, 1906, Am. Soc. Ent. Belg.

50: 130-134. Figs. 1-4, e, 9, d; Emery, 1910, Deutsch Ent.

Z., p. 131. Figs. 6, 7, e, .

Paratrechina vividula: Emery, 1925, Gen. Insectorum,

Faxc. 183: 223, Creighton, 1950, Bull. Mus. Comp. Zool.

Harvard 104: 409 (in part).


Worker. TL usually 2.0-2.5, OI usually 24-27. A

Southern species of disturbed habitats. Weakly bicolored;

head and gaster yellowish-brown to piceous, thorax and

appendages yellow to dark reddish-brown. Head subquadrate,

sides weakly convex, subparallel. Cephalic pubescence,

mostly limited to rear half of head and often rather varied

in length and spacing.

Male. TL about 1.9-2.25, OI usually 38-40. Body color

dingy yellow to dark brown, thorax usually lighter.

Antennal segment IV curved or bent. Parameres, viewed from

the rear, strongly curved mesad. Aedeagus as short or

shorter than parameres, spatulate, hyaline.

Queen. Not readily distinguishable from those of

terricola, faisonensis or concinna. In contrast to the

latter two species, often bicolored as in worker. Normally

has larger eyes than queen of terricola.


Allolectotype worker. TL 2.15, HL, 0.62, HW 0.53,

SL 0.71, EL 0.15, PW 0.38, MCL 0.16, WL 0.77, FL 0.57,

GL 0.77, SM 8, PM 4, MM 4, CI 85, OI 25, SI 115, FI 92.

In full face view, clypeus arcuate and entire. Dorsum

of clypeus with sides meeting in a rounded, median angle.

Head, exclusive of clypeus, subquadrate, the apparent

squareness accentuated by the anterior 3/4 of the sides

being virtually parallel except where they curve inward to

meet the mandibular insertions. Eyes in full face view

appearing about 1/4 as long as distance from mandibular

insertions to rear border, though in fact their greatest

diameter is longer. Distance between front of eye and

mandibular insertion slightly greater than EL, eyes sep-

arated from sides of head by less than 1/3X eye width.

Scapes curved over basal 1/4 of their length and over short

distal section, otherwise nearly straight. Scapes of about

average length for vividula group. No ocelli visible.

Pronotum in profile weakly angular; the anterior and

dorsal faces flat, the dorsal a little longer than the

anterior. Promesonotal suture clearly visible, though

barely impressed. Mesonotum flat-topped with short anterior

and posterior faces rising steeply to meet the dorsal face

in rounded angles, the posterior face a little longer and

steeper than the anterior. In profile, the propodeum nearly

evenly rounded, its highest point distinctly lower than that

of pronotum, the angle between its anterior and descending


faces weakly obtuse. Legs of about average length for

vividula group.

Petiole sharp-crested, cuneate in profile. In dorsal

view, with straight sides weakly divergent dorsad; crest

comprised of two nearly straight sides meeting in a blunt


Clypeus with 12 forward-projecting macrochaetae, those

closer to the front and center as long as clypeus in dorsal

view, and with a few appressed hairs just to the sides of

the median ridge. Front half of head with a few macro-

chaetae, those on frons somewhat shorter than longest

clypeal macrochaetae and those on infraocular region much

shorter and subdecumbent. Rear half of head with longer

more abundant macrochaetae and pubescence composed of 4 or

5 dozen unevenly spaced appressed simple hairs of varying

length. Scape macrochaetae suberect, less than half as long

as greatest width of scape and of the same color as the

scape; most easily seen in posterolateral view. The eight

major thoracic macrochaetae weakly curved; the remaining

ones half or less as long and straight. Forelegs with 3-6

macrochaetae on flexor surfaces of femora and tibiae and 1-2

on extensor surfaces. Middle and hindlegs with the reverse

pattern, namely several hairs on extensor surfaces and 1 or

2 on flexors. Propodeum with a narrow patch of longitudi-

nally oriented pubescence, along its front edge. Gaster

with sparse, very fine, appressed hairs and the usual com-

plement of macrochaetae.

Smooth and shining from head to first gastral tergite.

Weakly shagreened posterior to this.

Head and gaster light yellowish-brown. Thorax, brown-

ish-yellow. Mandibles light brownish-yellow with dark

reddish-brown teeth. Legs and scapes yellow.

Lectotype male. TL 1.89, HL 0.52, HW 0.48, SL 0.59,

EL 0.19, WL 0.81, GL 0.57, SM 4, CI 94, OI 37, SI 114.

Mandibles apically unidentate, without subapical den-

ticles in this specimen. Clypeus, in dorsal view with a

very shallow median emargination and median raised portion

narrower than in worker, evenly rounded, subumbonate. Sides

of head weakly convex, convergent anteriad, meeting the rear

border through slightly obtuse rounded corners. Limits of

weakly convex rear border lie approximately behind inner

border of eyes. Eyes convex, apparently about 1/3 as long

as head. Median and lateral ocelli separated by about 2X

their diameter. Scapes slenderer than worker's. Antennal

segment IV bent as in Fig. 6.

Petiolar crest more blunt than worker's. Petiole

broader than worker's in posterior view.

Parameres about 2X as long as broad in side view,

nearly triangular, but slightly recurved over distal 1/3

of their length. In rear view, parameres sickle-shaped,

curving mesad distally. Digitus shaped like a short

boomerang, i.e., narrow throughout, bent downward at the

middle through a broadly obtuse angle and tapering gently

to a rounded terminus. Cuspides of the usual shape for

vividula group, i.e., like an inverted bowl of a round

spoon. Aedeagal lobes shorter than parameres, slender,

triangular, weakly spatulate and splayed (as is normal in

preserved males of this species).

Vestiture as in worker but cephalic pubescence more

abundant and evenly distributed over rear half of head and

on frons. Scapes less pubescent than worker's and with a

row of 3, plus 1 offset, macrochaetae; these suberect, short

and pale, inconspicuous as in worker. Parameres with about

25 simple, decurved, yellow macrochaetae on their outer


Head, thorax, front of gaster and appendages uniform

yellowish-brown. Gaster increasingly darkening brown from

middle of first gastral tergite posteriad. Genitalia

yellowish-brown, except aedeagal lobes, which are yellowish-



Workers. TL 1.90-2.64, HL 0.55-0.69, HW 0.44-0.61,

SL 0.61-0.77, EL 0.13-0.17, PW 0.32-0.44, MCL 0.12-0.19,

WL 0.66-0.90, FL 0.46-0.62, GL 0.56-1.17, SM 6-9, PM 3-7,

MM 2-5, CI 80-91, OI 21-27, SI 105-118, FI 83-93, (n=58).

The clypeus may have a shallow concave or flat median

emargination in dorsal view. The sides of the head are

usually subparallel (in eastern specimens) or are wekaly

convex (especially in material from Texas westward). OI is

almost always greater than 24, but two series from separate

localities in the Guadalupe Mountains, Texas, have unusually

small eyes. These may represent a hybrid swarm of vividula

x terricola or simply are aberrant. Mean OI for the species,

including six specimens from Guadalupe Mountains is 25. The

rear face of petiole in lateral view is often convex or sub-

angular, rather than straight as in type, and the crest may

be rounded or with a flat top and sloping sides in rear view.

The pilosity of the scapes is often longer (almost

equal to width of scape) and brown, thus, much more con-

spicuous than in type; this is especially true in Gulf Coast

and Florida material. The latter specimens have more

abundant cephalic pubescence distributed further forward,

especially on frons, and may have some long sparse appressed

hairs on the pronotum and mesonotum. Specimens from Arizona

and California typically have the cephalic pubescence short,

very fine and more evenly spaced but still limited to rear

portion of head.

Western material is generally colored about as type,

and most eastern material is darker. However, I have seen

series from Arizona and Mississippi which are nearly uniform

very dark brown, thus, recognizable only by head shape, eye

size, and pattern of pubescence.

The types and one additional series of kincaidi from

Jamaica, here synonymized with vividula, have color and

vestiture like that of peninsular Florida specimens.

European greenhouse material tends to look more like vivid-

ula from Texas or Northern Mexico. It should be recalled

that Emery (1906) suggested Mexico as the provenience of


Males. TL 1.89-2.25, HL 0.46-0.55, HW 0.45-0.52,

SL 0.54-0.62, EL 0.18-0.21, WL 0.69-0.85, GL 0.57-0.98,

SM 4-6, CI 91-98, OI 33-42, SI 107-119, (n=ll).

The mandibles of the males of vividula vary from simple

apically unidentate to having an apical tooth, a small cleft,

then a subapical denticle or two. The crook in antennal

segment IV is sometimes manifest only as an even curve. Very

few specimens lack the splaying of the aedeagal lobes typi-

cal of preserved males in this species. Most specimens are

significantly darker than the type ranging from a dingy yel-

low to a dark reddish-brown, often with the thorax somewhat


Material Studied

The type material (HM) was collected in a greenhouse;

Helsinki, Finland; 2-25/X/1845. Included are the lecto-

type, allolectotype, three paralectotype workers and a

paralectotype queen (minus gaster). I have deposited

conspecific material from Texas and Florida at HM. In

addition, I have studied many hundreds of specimens from

across the southern U.S. and Europe. The material was

loaned or donated from LACM, USNM, TPN, DPW, and TTU. JCT

contains Florida material. European material and types of

kincaidi (here synonymized with vividula) are at MCZ. The

material from TTU was particularly valuable (in addition

to the suggestions of Emery, 1906), in helping me make a

correct guess about the identity of vividula even before I

saw the types, because of the large number and size of the

series, many of them containing7 associated males. TTU also

contained a similar large amount of terricola, thus helping

to preserve the latter from being synonymized, as was my

initial inclination.


This species is very closely related to terricola;

isolated workers cannot always be assigned to one or the

other species with certainty. The larger OI and more

straight-sided head will suffice to separate the great

majority of vividula workers. The ranges of the two species

are different (Fig. 49). I collected one series of concinna,

normally uniform dark brown, which I mistook in the field

for vividula because they were bicolored. The specimens

were in all other characters typical concinna. Males of

vividula are distinguished from terricola by genital char-

acters as described above. Most of the vividula material I

have seen in collections has been identified as melanderii,"

while most of the material identified as vividula previous

to this study does not belong to vividula (see discussion

of faisonensis). I believe this is due to the long standing

dogma stated in Creighton's key (1950, p. 404), "erect hairs

on the antennal scapes [are] abundant and occurring on the

sides as well as the front of the scape" in vividula. In

fact, the macrochaetae on the scape are almost always lim-

ited to two rows on the anterodorsal part of the scape com-

prised of 3 or 4 hairs each. The macrochaetae themselves

are often inconspicuous, giving the casual observer the

impression that there are less of them than, in fact, occur.

It should be noted that the scape pilosity of terricola is

of the same character as that of vividula and cannot be used

as a separatory feature for the two species.

The type series of kincaidi (MCZ) unfortunately lacks

the male that originally belonged to it, and he could not

be found in the unit tray in which the series had been

stored by MCZ staff who searched for it upon my request.

Nevertheless, it seems unquestionable to me, based on the

features of the workers and queens in the series and upon

Wheeler's (1906) description and figure of the genitalia

of male kincaidi, that kincaidi can be none other than a

synonym of vividula.

Natural History

Under conditions of sufficient warmth, alates may be

present at any time of year and may fly any warm day when

humidity is high. In most of the range in the U.S., flights

are restricted to the months May-October. Females are

occasionally attracted to lights at this time of year,

though flights, from the little available evidence, occur

during the day. I observed a group of six males patrolling

a fence post at 1300 hr in Lake Placid, Florida, in January,

and I have alates of both sexes collected from the surface

of a pool in a Juniper-oak woodland in Texas on 7 Septem-

ber (PSW).

This ant is characteristic of open, usually rather dis-

turbed habitats including beaches, parks and other land-

scaped areas, crop fields, fallow fields, vacant lots,

parking lots, etc. Nuhn and Wright (1979) reported finding

vividula (as Paratrechina sp.) in fields and other open,

disturbed habitats in North Carolina. It commonly estab-

lishes colonies in plant pots or mulch piles and is trans-

ported in these materials to greenhouses and nurseries far

from the areas where it survives outside. It has apparently

been established as a greenhouse ant in Europe for nearly

150 years. This ant occurs in less disturbed rangeland and

other open habitat in southern Texas and northern Mexico,

where it overlaps with and may occur in a mosaic distribu-

tion with respect to its sibling species, terricola.

Paratrechina terricola

(Figs. 5, 6, 7; Map-Fig. 49)

Formica (Tapinoma) terricola Buckley, 1966, Proc. Ent.

Soc. Phila. 6: 168. ,, 9, d. Type loc., Austin, TX.

Prenolepis melanderi Wheeler, 1903, Psyche 10: 104.

Fig. 8, t, 9, -. NEW SYNONYMY. Type loc., New Braunfels,

TX (designated by Creighton, 1950).

Prenolepis (Nylanderia) vividula melanderi: Emery,

1906, Ann. Ent. Soc. Belg. 50: 132. Fig. 5, d; Emery, 1925,

Gen. Insectorum, Fasc. 183: 223.

Paratrechina (Nylanderia) melanderi: Creighton, 1950,

Bull. Mus. Comp. Zool. 104: 407; G.C. and J. Wheeler, 1953,

Ann. Ent. Soc. Amer. 46: 144. Pl. II, Figs. 16-20, larva.


Worker. TL usually 1.9-2.4 (southern part of range),

2.2-2.7 (north), OI 20-24. A vividula-like species of open

habitats and disturbed areas of northern Mexico, Arizona

and Louisiana, north to Illinois and South Dakota and in the

foothills of the Smoky Mountains. Particularly abundant

in Texas. Yellowish-brown with darker head and gaster (more

common south) to piceous with yellowish-brown appendages

(more common north). Cephalic pubescence very sparse, head

shiny. Eye small.

Male. TL 2.1-2.4, OI usually 35-37. Piceous brown to

nearly black. Eye relatively small (compared to vividula).

Parameres only weakly curved mesad. Distal portion of

digitus about 2X as broad as that of aedeagus. Aedeagus

longer than parameres, curved upward distally. Crook in

antennal segment IV conspicuous.

Queen. Averages smaller and smaller-eyed than

vividula, often not distinguishable.


Alloneotype worker. TL 1.98, HL 0.61, HW 0.52,

SL 0.68, EL 0.13, PW 0.36, MCL 0.16, WL 0.77, FL 0.54,

GL 0.61, SM 8, PM 4, MM 3, CI 86, OI 22, SI 112, FI 88.

Clypeus lentiform; anterior clypeal border arcuate

with a faint, narrow median concavity; rear border less

strongly arched than front border, also with a small median

concave emargination. Sides of head weakly convex along

most of their length and slightly convergent anteriad.

Rear border of head rounded with a shallow median notch a

little wider than the distance between the frontal carinae.

Head of about average breadth for the species. Eyes smaller

than in vividula, approximately their rear 1/5 overlapping

the midpoint of the postmandibular head length; separated

from sides of head by about 1/2X eye width and from mandibu-

lar insertions by about 1 1/3X EL. Scapes weakly curved

near the ends, straight over most of their length; of about

the mean relative length for the species.

Pronotum broadly angular, the somewhat shorter anter-

ior face flat and the dorsal face weakly convex. Mesonotum

flat on top, sloped downward to the rear, its short anter-

ior face meeting pronotal dorsum face at about the same

angle as that between 2 faces of the pronotum. The poster-

ior mesonotal face shorter and much steeper than the anter-

ior mesonotal face. Propodeum low and arched, its highest

point only about as high as the rear edge of the mesonotal

dorsum. Legs of about average length for the species.

Rear face of petiole obscured by anterior face of

gaster, but petiole apparently cuneate with a blunt rounded

crest and flat front face in profile. From above, petiolar

crest is flat across the middle with rounded corners, and

the sides are straight and weakly divergent dorsad.

Cephalic vestiture resembling that of small vividula

workers but sparser than in most individuals of the latter

species. Cephalic pilosity is rather short, weakly curved,

and sparse, especially on the sides of the head. Cephalic

pubescence sparse, widely spaced, the spaces between the

hairs wider than the length of the hairs in most cases.

The appressed hairs themselves are longer than those of

other U.S. species. Thoracic macrochaetae about half as

long as the greatest width of the scape, of the same color

as the scape, protruding above the finer subdecumbent pubes-

cence which covers the scape. Ancillary thoracic macro-

chaetae about half as long as the shortest major macrochaetae

and of the same dark brown color (pronotum) or somewhat

lighter (mesonotum). Gaster covered with thick, dark brown

rearward-curved macrochaetae of about even length and

spacing throughout; and with some very sparse, long pubes-


Body very shiny and smooth, except gaster which has

some very fine almost indetectable (at 40X) shagreening.

Head piceous brown becoming a little lighter on the

sides and clypeus. Scapes and mandibles brownish-yellow,

the latter with dark brown teeth. Thorax, except mesonotum,

colored like sides of head, mesonotum lighter and yellower.

Legs brownish-yellow with the middle and hind coxae paler.

Gaster piceous brown.

Neotype male. TL 2.27, HL 0.53, HW 0.48, SL 0.59,

EL 0.18, WL 0.74, GL 1.01, SM 6, CI 92, OI 35, SI 112.

Mandible with an apical tooth, a sharp subapical

denticle and a crenulate masticatory border. Anterior

border of clypeus arcuate, rear border with straight sides

and a narrow median concave emargination. Sides of head

convex from posterior corners to just before the eyes,

straight from there to mandibular insertions. Head narrower

than that of vividula. Rear border of head less convex

than sides, about as broad as the distance between the

mandibular insertions. Eyes apparently only about 1/3X HL

(in fact a little longer) but strongly convex, protruding

well beyond sides of head. Ocelli not large but fairly

close set, median and laterals separated by about 2X their

width. Scape weakly sigmoid, a little shorter than average

for the species. Antennal segment IV longer than adjacent

segments and distinctly bent at the middle.

Petiole cuneate with a blunt rounded crest, thicker

than that of worker. In dorsal view, the crest is flat

across the top and about twice as wide as that of the worker

with straight sides weakly divergent dorsad; the anterior

face is concave.

Genitalia distinctive, even from those of closely

related vividula. In side view, paramere is about twice

as long as its basal breadth, but it tapers quickly in the

basal 1/3 of its length and is digitiform over the distal

2/3. Cuspis curved mesad to overlie the digitus in pos-

terior view and is a little larger and more prominent than

in vividula, faisonensis, etc. but much shorter and less

prominent than in arenivaga. Digitus not decurved as in

vividula but with basal edge straight. Distal portion of

digitus about 2X as broad as narrowest portion of aedeagus;

the former subtriangular with angles rounded. Outer face

of digitus convex. Aedeagus distinctly longer than para-

meres; the lobes closely appressed (not splayed, as in

vividula); conspicuously broadened dorsad and truncate

distally, resembling a miniature inverted tomahawk (Fig. 7).

Pilosity finer than worker's, brown to dark brown, of

approximately equal spacing and length over entire dorsal

surface, except on anterior half of gaster. Pilosity

virtually lacking on sides of head and thorax. Parameres

with more than 20 yellowish, weakly decurved hairs. Pubes-

cence on head sparse not obscuring sheen of integument.

Pubescence of thoracic dorsum denser.

Sculpture much as on workers, but shagreening on

gaster consisting of larger, more conspicuous integumental


Eyes grey. Body and appendages uniform piceous,

except articulations and tarsi of legs and post-mandibular

mouthparts, all of which are light yellowish-brown.


Worker. TL 1.87-3.03, HL 0.54-0.73, HW 0.44-0.63,

SL 0.58-0.84, EL 0.11-0.16, PW 0.30-0.44, MCL 0.12-0.19,

WL 0.65-0.93, FL 0.45-0.67, GL 0.61-1.37, SM 5-10, PM 3-6,

MM 2-4, CI 80-92, OI 20-24, SI 106-118, FI 83-93, (n=42).

This species is sympatric (or more precisely, mosa-

ically parapatric) with arenivaga and parvula in the western

half of the latter two species' ranges. Interestingly,

terricola exhibits the same tendency as these other species

in having larger overall body size and a broader head in

the northern and eastern parts of its range. TL of Arizona

specimens averages about 1.9, that of Texas and Oklahoma

specimens about 2.2, that of Kansas and South Dakota speci-

mens about 2.4, and that of Tennessee specimens almost 2.6.

CI of specimens from southern states ranges 80-87, while

CI of Kansas and South Dakota specimens ranges 84-92. Cor-

related with size trends is a greater degree of investiture

of northern material. Four of the five workers with SM 10

and five of the eight workers with SM 9 are from northern

states, as are all individuals with PM>5 and MM>3. Greater

density of cephalic pubescence is normal on larger specimens

wherever they occur. Specimens from Arkansas (UARK), pre-

dictably, span the gap between the large, hairy, Tennessee

material and the smaller, nearly glabrous Texas material.

Male. TL 2.12-2.40, HL 0.49-0.59, HW 0.46-0.54,

SL 0.57-0.69, EL 0.17-0.21, WL 0.73-0.93, GL 0.85-1.05,

SM 4-6, CI 90-94, OI 31-37, SI 110-117, (n=9).

These numbers describe fairly well the metric varia-

tion of terricola males. Most males have CI 35-37. The

two specimens with CI 31 and 33 are unusually small-eyed.

I have seen no terricola males from Arizona but would

expect them to have small eyes as do workers from that


The paramere is generally narrower and straighter

(i.e., more digitiform) than in vividula males. While it

is always less curved, the shape may closely resemble the

weakly recurved triangular form of the vividula paramere.

The aedeagus of terricola males is normally considerably

longer than the parameres, and the lobes of the aedeagus

are closely approximated in preserved material. In side

view, in the intact animals, the aedeagus normally projects

below the parameres. Occasionally, especially in Tennessee

material, the aedeagus more closely resembles that of vivid-

ula in being shorter than the parameres and having its halves

splayed and projecting above the parameres. In such cases,

the structure of the volsella is that typical of terricola,

and the specimens have in each case belonged to series in

which other males have the aedeagal structure more typical

of terricola.

Material Studied

The neotype male and alloneotype worker were desig-

nated from a nest series (TTU) labeled Texas, Real Co. 12

mi NE Leakey. Among small rocks. 19 Mar 1979. O. Francke

et al. #3725. The pins bearing the neotype and alloneotype

are deposited at LACM, and the remaining three pins from

the series are deposited at MCZ. Other series, containing

at least males and workers, from nearby Texas localities

(all from TTU) are deposited at these and the other usual


Material from North Dakota and Oklahoma was from GJW.

Kansas material was from JCT and SBSK. A large amount of

Texas material and smaller samples from other states is at

LACM and USNM. Wheeler's types of melanderi (here synony-

mized with terricola) are at MCZ and AMNH. A large sample

of workers from Arkansas (UARK) seems to fill in the middle

portion of the size and hairiness dines between Texas and

Tennessee and upper midwestern specimens.


The resurrection of Buckley's (1866) name terricola

for the ant which has been known for 80 years as melanderi

is based on several considerations. The first is that

despite an adequate description and fine drawings accompany-

ing Wheeler's (1903) description, the name melanderi has

been applied recklessly to almost every native species of

nearctic Paratrechina. Most references to melanderii" in

the literature apply either to other species or to a mix

of species which may or may not include the melanderi of

Wheeler's description. In its modern usage, the name has

become at worst meaningless and at best highly ambiguous.

Of course, this is true for many names in this genus and

would not alone constitute sufficient reason for bringing

into use a name of apparently uncertain identity in place

of melanderi.

Another problem concerning melanderi is Creighton's

designation of New Braunfels, Texas, as its type locality.

Among the syntypes of melanderi at AMNH are seven workers

and parts of two others from New Braunfels, but no males.

MCZ contains no syntype material from New Braunfels. The

only male in the entire syntype series, as presently con-

strued, is a rubbed, headless male from Austin.

The importance of the lack of suitable male specimens

among the melanderi syntypes becomes clear when we consider

that vividula and terricola are separable with certainty

only in the males. It is for this reason that I have desig-

nated males as lectotype and neotype, respectively, for

these two species.

There is considerable overlap in the phenology of

workers of terricola and vividula, so much so that lacking

associated males, worker specimens often cannot be identi-

fied. The differences in the males, contained in the de-

scriptions and in Figs. 7 and 8, need no elaboration. What

follows is a listing of hints for identifying workers of

these two species in the absence of males. The list is, of

course, based on the assumption that all other possibilities

besides terricola or vividula have been eliminated.

1. Specimens collected outdoors from north of the

Mason-Dixon line, from undisturbed or moderately disturbed

habitats are almost certainly terricola. Specimens col-

lected in Gulf or Atlantic coast states east of Louisiana

are almost certainly vividula.

2. In Mexico and Arizona, terricola averages about

2.0 mm in length while vividula from the same area averages

a little larger, about 2.25 mm.

3. Specimens with OI 24 or less are usually terricola.

Workers with OI 25 or more are certainly vividula.

4. Specimens with very sparse cephalic pubescence are

usually terricola, though small vividula specimens may have

the pubescence on the head quite dilute. Specimens with

denser (though very fine) pubescence on the rear portion

of the head are probably vividula.

Finally, we consider Buckley's description itself and

enumerate the reasons why it must concern the ant Wheeler

redescribed as melanderi.

1. The name Formica (Tapinoma) terricola--Creighton

(1950, p. 24) points out that Buckley was strongly influ-

enced by the British hymenopterist Frederick Smith. Smith

called the Paratrechinas he described Tapinoma (e.g.,

flavipes). It is possible that Buckley would have used the

name in the same way.

2. The size of the castes--The size reported for the

queen is certainly too large for a Paratrechina (6.86 mm).

The sizes for the worker and male (2.0 mm) are appropriate.

(Buckley gave the queen measurement as 0.27 inch which may

be a misprint for 0.17 inch which equals about 4.32 mm,

approximately the size of a Paratrechina queen).

3. Coloration--The color of the queen is described as

piceous, and the males are said to be "like the female."

The worker is described as dark brown above, lighter below

and on the petiole and legs. Fully sclerotized Texas ma-

terial of this species fits these descriptions.

4. Head shape--The head of the queen is described as

"small, triangular," i.e., strongly convergent anteriad.

The worker's head is said to be "little wider than the

thorax." While the latter is an understatement of the true

width, it does suggest that he observed the relative narrow-

ness of the head of many workers from Texas.

5. Vestiture--The queen is described as "thickly

sprinkled with short grey hairs," probably referring to her

dense pubescence. No such pubescence is mentioned for

workers or males, as is appropriate.

6. Venation--Buckley says the wings have one marginal

and two submarginal cells (i.e., a submarginal and a median

cell) and that the discoidal cells are obsolete--as in


7. Petiole--In Buckley's words, pediclee short, in-

serted in the base of the abdomen . inclined forward,

wedge-shaped." This describes the petiole of Paratrechina


8. Ecology--Terricola is described as "very active"

(i.e., fast-moving) "with shallow nests in soil. Winged

males and females captured in March" and as "rare," which

reflects the great difficulty in encountering nests except

in early spring. Among ants that could fit the rest of

Buckley's description, only Paratrechina has alates in the

nest in March. The color is inappropriate for arenivaga,

the locality is inappropriate for bruesii, and the alate

season is inappropriate for vividula. Melanderi is the only

suitable Texas Paratrechina for the description.

Any one or a few of these points, except perhaps number

8, would leave us in doubt, but considered together, I

believe they describe, however poorly, Wheeler's melanderi

which must, therefore, be relegated to synonym. It is

probable that Buckley's Formica picea (1866, p. 163) is in

fact a description of terricola minims or other unusually

small workers. Lacking sufficient evidence, I do not include

picea in the synonymy. While I realize that resurrecting

terricola from the ranks of Buckley's numerous nomina nuda

constitutes a virtual myrmecological apostasy, I trust that

the evidence presented shows this to be the best course for

eliminating confusion and stabilizing the taxonomy of the

nearctic Paratrechina.

Natural History

Alates of terricola overwinter in the nest and, as

indicated in both Buckley's and Wheeler's descriptions, are

commonly found in March in Texas. Texas alate records range

from January through April. A Kansas record is from April,

and I collected a colony with male pupae from Lawrence,

Kansas, on 2 August. Series from Tennessee and Missouri

with alates are dated March and April, respectively. Col-

lections of alates in May from Tennessee and Arkansas are

both from mountainous areas.

Mitchell and Pierce (1912) reported "large numbers of

winged forms were collected running, flying and mating,

March 2, 1909." This description recalls to me my observa-

tion of a mating swarm of vividula in which males ran about

on the substrate, periodically flying up then quickly cir-

cling back into the swarm and running about on the substrate

with wings vibrating.

Buckley (1866), Wheeler (1903), Mitchell and Pierce

(1912) and Van Pelt (1983) state that this species nests

in soil, usually beneath a stone, log or cow dung. Records

from the TTU collection indicate they are found in most

soil types, except very sandy soils, and in vegetation

varying from open disturbed areas and pasture to mesquite

and post-oak woodland and occasionally in denser more mesic

woods. Specimens from South Dakota (GJW) were from Juniper--

Rhus trilobata--Chrysothamnus woodland. Kansas collections

are from woodland edges and openings, pastures and meadows.

Male terricola, lateral view.

Male vividula, lateral view.

Worker of terricola, lateral view.

Worker of vividula, lateral view.

Fig. 7.

Fig. 8.

Fig. 9.

Fig. 10.


Diagnosis of Complex

Worker brown or bicolored, shiny; head with dense pubes-

cence, this arising from very fine to rather conspicuous

punctae; scape pilosity reduced in some species, suberect

to erect. Nests in a variety of undisturbed or lightly dis-

turbed, moist habitats in leaf litter, rotten wood, clump-

ing plants or in soil.

Male shiny, but thoracic and cephalic dorsum usually

dulled by pubescence; parameres triangular; digitus boomer-

anged-shaped; cuspis short; aedeagus triangular, acuminate.

Paratrechina concinna, New Species

(Figs. 11, 14, 38; Map-Fig. 50)


Worker. TL usually 2.3-2.9, CI usually 84-89,

OI 24-28. A species found from Florida to North Carolina,

usually found in marshes, wet grasslands and pastures, and

swamp edges. Uniform brown or with thorax and legs slightly

lighter than gaster and head. Scapes and middle and hind

coxae brown (never tan or whitish). Overall size large

among native eastern U.S. species. Cephalic pubescence

arising from small foveolae giving punctate appearance.

Pubescence of pronotum and proprodeum usually conspicuous,

that on front of pronotum subdecumbent.

Male. TL 2.25-2.5, OI usually 34-37. Closely resem-

bling faisonensis but darker in color, has larger body and

eyes and cephalic sculpture as in workers.

Queen. Resembles vividula and faisonensis. Differs

in having uniform dark color, cephalic punctation, and

slightly larger size.


Holotype worker. TL 2.72, HL 0.72, HW 0.65, SL 0.81,

EL 0.19, PW 0.46, MCL 0.20, WL 0.91, FL 0.65, GL 1.09,

SM 12, PM 7, MM 4, CI 90, OI 27, SI 113, FI 90.

In full face view, clypeus lentiform, anterior border

arcuate with a shallow concave emargination across the mid-

dle 1/4; rear border also arcuate. Dorsum of clypeus with

sides meeting in a rounded median angle. Sides of head

rounded, convergent anteriad; widest just behind the eyes.

Rear border of head straight with rounded corners; the

corners about as far apart as the width of the clypeus.

Head subquadrate, broader than usual for the species (CI 90

versus mean CI 86 for the species). Eyes longer and some-

what broader than in faisonensis, slightly more convex.

Eyes separated from sides of head by less than 1/3 their

width; from mandibular insertions by slightly over IX EL.

Scapes a little shorter than average for the species, which

has SI slightly but probably insignificantly longer than

average for vividula group. Scape very slightly but evenly

curved over its entire length, distal portion tapering giv-

ing the illusion of greater curvature. Small round markings

at the positions of the ocelli probably represent vestiges

of those organs.

Pronotal profile evenly curved, somewhat more convex

than in faisonensis. Anterior face of mesonotum rising from

promesonotal suture to dorsal face through about a 50% slope;

posterior declivity steeper and longer than usual in the

faisonensis; mesonotal dorsum slightly sloped to the rear.

Propodeum low and evenly rounded, only a little higher in

profile than metathoracic spiracles, the angle between its

anterior and declivious faces slightly obtuse.

Petiole sharp-crested, create in profile; in dorsal

view broader and with more rounded sides and crest than


Cephalic pilosity as in faisonensis. Cephalic pubes-

cence very dense, reminiscent of queens of other species,

and neatly aligned, giving a combed appearance. Thoracic

macrochaetae conspicuous like those of faisonensis. Thorax

appears quite bristly because the macrochaetae are less

curved than those of faisonensis, and the eight major hairs

are subtended by ancillary hairs about 3/4X as long and

these in turn (on the pronotum) by smaller hairs about 1/2

length of the latter. Pubescence of thorax is diagnostic:

subdecumbent and unusually abundant (for the vividula

group) on the anterior portion of the pronotum, scattered

appressed hairs on the metanotum and metapleuron, and

abundant suppressed hairs on the anterior portion of the

propodeum. Gaster with a few appressed hairs, unlike fais-

onensis. Petiole with two small macrochaetae on the crest.

Thorax and gaster strongly shining, the former very

smooth, the latter delicately shagreened. Head and

appendages punctate, feebly shining, the sheen reduced by

the denser pubescence which arises from tiny foveolae.

Uniform castaneous brown with only the articulations

of the appendages and the blade of the mandibles a little


Allotype male. TL 2.46, HL 0.65, HW 0.61, SL 0.73,

EL 0.22, WL 0.97, GL 0.85, SM 6, CI 94, OI 34, SI 112.

Head large compared to all other males in vividula

group. Head shape, including that of clypeus, strikingly

like that of worker. Though CI indicates head is propor-

tionately wider than worker, this is because the head width

is taken across the eyes which are much larger than workers'

and protrude beyond the sides of the head. Head is, in

fact, a little narrower than workers' minus the eyes.

Mandibles strongly sclerotized, dark colored with an apical

tooth, cleft and a subapical denticle, the identical larger

than in any specimen of vividula having it. Scape a little

shorter and with fewer macrochaetae than workers but pilose

for a vividula-group species. Eyes larger and more

prominent than faisonensis. Median ocellus notably larger

than laterals.

Upper portion rear face of petiole concave. Petiole

otherwise as in worker but broader. Gaster contracted,

shorter than thorax. Genitalia closely resembling those

of faisonensis but with cuspis a little more convex and

generally heavier and darker colored.

Vestiture as in workers with the usual differences.

Parameres with decurved pilosity as in faisonensis but

this less abundant, about 20 hairs.

Not as shiny as workers, due to occurrence of pubi-

gerous punctation on thoracic dorsum and coxae. Gaster also

appearing somewhat roughened, this due to a slightly coarser

version of the usual shagreening.

Darker in color than all other vividula group species,

expect flavines, from which concinna differs by larger size

and having dark colored appendages. Head, thorax, and ap-

pendages dark brown, gaster piceous.


Worker. TL 2.13-3.10, HL 0.61-0.76, HW 0.52-0.65,

SL 0.71-0.85, EL 0.15-0.20, PW 0.37-0.49, MCL 0.16-0.20,

WL 0.77-0.93, FL 0.57-0.71, GL 0.65-1.41, SM 7-16, PM 4-8,

MM 2-6, CI 83-90, OI 24-28, SI 110-125, FI 88-97, (n=41).

This species seems to be quite uniform in general

appearance throughout its range. Head shape somewhat vari-

able as indicated by the large range of CI. This seems to

have no relationship to overall size, as even large workers

may have heads in the narrow end of the range. The abun-

dance and degree of appression of the pronotal pubescence,

which occasionally consists of only a few appressed hairs

is also somewhat variable. The macrochaetae on the petiolar

crest vary from 0-2. The color of the thorax is not uncom-

monly a little lighter and yellower than that of the head

and gaster. This is probably the normal condition of

younger workers.

Male. TL 2.24-2.46, HL 0.59-0.67, HW 0.55-0.63,

SL 0.67-0.74, EL 0.20-0.24, WL 0.87-1.01, GL 0.69-0.85,

SM 5-7, CI 92-97, OI 33-38, SI 108-117, (n=8).

The large size, dark color, distinctly short gaster

and worker-like aspect of the head of concinna males show

little variation in the specimens from Florida and North

Carolina I have seen. Most have the ocelli more equal in

size than those of type.

Material Studied

Holotype and allotype and a large series of paratypes,

including queens, were collected 15 April 1980, 1.5 miles

ESE of Gainesville, Florida, under a plant pot on a damp

lawn at the edge of a sweetgum swamp. Holotype and allo-

type are deposited at FSCA. Paratypes are deposited in the

usual depositories.

Most of the material studied was collected by myself

in Alachua County and Highlands County, Florida. A number

of collections made by Van Pelt (1957) are stored at FSCA.

Additional small series were found in LACM, USNM, AMHN,

DPW (USDA) and TPN, these from Florida, Georgia, and North



The name concinna (Latin--neat, orderly) refers to the

neatly aligned appearance of the cephalic pubescence.

This species falls into Creighton's concept of vivid-

ula, but I do not believe he ever saw it and, thus, have not

included his vividula treatment in the synonomy. There are

only a few specimens of concinna in the major collections

due to its preference for habitats rarely collected by ant

hunters. My collection contains many more concinna than

all other collections together except FSCA, which reflects

both the poor representation of Florida ants in these col-

lections and the unlikelihood of collectors'entering marshy

areas to find ants.

Concinna could be confused in the field with vividula,

faisonensis or bourbonica. Fiasonensis lacks thoracic

pubescence and is smaller, while bourbonica is more uni-

formly covered with dense grey pubescence and is larger.

Large, dark specimens of vividula can be recognized by

their smooth heads, less abundant cephalic and thoracic

pubescence and lower SM.

Based on the overall similarity of the workers and

of the genitalia of the males, I believe concinna is most

closely related to faisonensis.

Natural History

Like vividula, this ant may, under appropriate condi-

tions, produce alates at any time of year. I have records

of alates and alate pupae in nest series from south Florida

in December (Highlands County) and May (Dade County), and

I have alates from North Carolina collected in April and

September. In the Gainesville, Florida, area, these ants

seem not to raise alates during the cool months; the peak

rearing of sexual brood occurs March-July.

I believe, based on the collections of Van Pelt (FSCA)

and myself in relatively pristine habitats, that concinna

was originally a species of marshy areas and of openings

in flatwoods and swamp. In such habitats, it nests in

grass tussocks and in pieces of rotten wood that have

fallen from nearby wooded areas. Today, it probably is

more readily found in pastures and lawns of high water-table

areas and along ditches and drainage canals where it nests

in tussocks, cow dung and discarded wood. I found one

colony at a laboratory complex in Gainesville, nesting in

leaf litter beneath shrubbery. Vividula is abundant at

this site, which is adjacent to a low pasture. A collec-

tion from xericc mixed woods" is Seminole County, Georgia

(LACM), is unusual and possibly mislabeled.

Paratrechina faisonensis

(Figs. 1, 2, 12, 15, 42; Map-Fig. 50)

Prenolepis arenivaga var. faisonensis Forel, 1922.

Rev. Suisse Zool. 30: 98. >. Type loc. Faison, NC.

Paratrechina vividula: Creighton, 1950. Bull. Mus.

Comp. Zool. Harvard 104: 408 (in part?); Nuhn and Wright,

1979, Amer. Mid. Nat. 102: 353-362; Thompson et al., 1979,

Psyche 86: 321-325.

Paratrechina melanderi: Lynch, Balinsky and Vail,

1980, Ecol. Ent. 5: 353-371; Lynch, 1981, Oikos 37: 183-198.


Worker. TL usually 2.0-2.5, CI usually 83-87, OI

usually 20-23. A deciduous woodland species of the south-

east U.S. north to the New Jersey Pine Barrens. Brown with

tan to whitish middle and hind coxae. Head narrow, CI

usually 83-87. Eye small, weakly convex. Cephalic pubes-

cence dense, partly obscuring sheen of head. Erect macro-

chaetae on scape conspicuous, almost as long as width of

scape. Thorax and propodeum normally completely lacking

pubescence, very shiny. SM 8-13.

Male. TL usually 2.0-2.25, OI usually 31-33. Color

as in worker. Head narrow due to small, weakly convex eyes.

CI usually around 90 (94+ in sympatric species). Scape

macrochaetae conspicuous, MS 4-7.

Queen. Small eyes, light coxae and very shiny in-

tegument of head and thorax (visible even through dense

pubescence when specimens are clean) distinguish this

species from vividula, concinna and arenivaga.


Lectotype worker. TL 2.10, HL 0.59, HW 0.48, SL 0.69,

EL 0.13, PW 0.35, MCL 0.16, WL 0.71, FL 0.53, GL 0.81, SM 9,

PM 3, MM 2, CI 83, OI 22, SI 117, FI 90.

In full face view, clypeus subtrapezoidal, anterior

border truncate, its straight portion about half as wide as

its sinuate rear border. Sides of head rounded, slightly

convergent anteriad, widest just behind eyes. Middle half

of rear border of head nearly straight except for narrow

median concavity. Head, exclusive of clypeus, longer than

broad, narrower than that of related species. Eyes small,

weakly convex separated from the sides by more than 1/2 the

eyes' width, and separated from mandibular insertions by

about 1 1/2X EL. Scapes a little longer than average for

the species, evenly and very slightly curved for about the

basal 85% of their length, a little more strongly curved

distally. No ocelli visible.

Pronotal profile an even, very weakly convex upward

slope toward the rear. Promesonotal suture feebly impressed.

Mesonotal profile almost flat, parallel to long axis of

thorax and lacking an anterior face, thus yielding a hemi-

parabolic profile together with the pronotum.

Posterior portion of mesonotal profile bearing a small

setigerous swelling, followed by a posterior declivity.

Proprodeum partly crushed on the left side but in profile

appears evenly rounded like that of vividula. The angle

between its base and its rearmost portion near the petiole

weakly obtuse. Legs of usual length for vividula group.

Petiole partly imbedded in glue, appears cuneate, but

more rounded than in vividula. In posterior view, crest

is an even curve.

Cephalic pilosity as in vividula; the pubescence fine,

more densely and more uniformly distributed over entire head

but difficult to see in this callow specimen. Scape macro-

chaetae brown, suberect, a little longer and stouter than

those of vividula and quite visible. Thoracic macrochaetae

dark brown, gently and evenly curved mesad. Pilosity of

gaster and legs just as in vividula. Thorax and abdomen

lacking appressed pubescence.

Head and thorax shiny (in the former case even through

the pubescence). Gaster delicately shagreened.

Head and gaster light brown, thorax, legs and scapes

a shade lighter, middle and hind coxae very pale yellow.


Worker. TL 1.83-3.00, HL 0.53-0.69, HW 0.44-0.61,

SL 0.63-0.80, EL 0.12-0.16, PW 0.29-0.44, MCL 0.14-0.20,

WL 0.65-0.89, FL 0.47-0.65, GL 0.57-1.49, SM 8-13, PM 2-6,

MM 2-5, CI 81-90, OI 20-24, SI 112-119, FI 87-97, (n=40).

TL is typically less than 2.5; the 3.0 specimen's length

largely due to its very distended gaster. CI is usually less

than 87; only one specimen had CI 90.

The clypeal shape described in lectotype is common,

but the anterior border may range from arcuate and virtually

entire to having two straight sides with a narrow concave

emargination. The eyes of some specimens are larger and

more convex, approximating the condition of related species

(see discussion). A vestigial median ocellus is rarely

present. The rear border of the head of larger specimens

in full face view usually has a broader emargination than

the type (and most other smaller specimens), yielding a

weakly subcordate appearance. The widest portion of head

is directly across eyes in many larger specimens. The

promesonotal profile is usually hemiparabolic as described

in type, but often the parabola is broken by a short, steep

anterior mesonotal face. Rarely, the pronotum is very

obtusely subangular. The sides of the petiole appear to

diverge a little more toward the crest than in vividula

(the latter feature not visible in lectotype).

Pilosity is often a little more abundant than on the

type, particularly on the clypeus and promesonotum. In the

latter case, the eight major macrochaetae are 2X or more as

long as the six or eight adjacent ancillary hairs, and a

few still shorter hairs may be present. Some large speci-

mens may have a small amount of pubescence along the anter-

ior edge of the propodeum, or rarely on the anteromedian

portion of the pronotum.

The color of fully sclerotized individuals varies from

nearly uniform dark brown to dark yellowish-brown. The

middle and hind coxae are less contrastingly pale in the

lighter specimens. Occasionally, the thorax and, less often,

the head are a little lighter than the gaster. The color de-

scribed for the type is typical of allows. Many darker

specimens may have some faint bluish reflections on the


Description and Variation

Male. (Composite description, no males in syntype

series. Two males from my collection have been deposited

at MHNG. I have not formally designated them as allopara-

types, since they are not from the type locality.)

TL 1.96-2.24, HL 0.52-0.57, HW 0.46-0.51, SL 0.61-0.65,

EL 0.16-0.18, WL 0.73-0.83, GL 0.65-0.93, SM 4-7, CI 87-92,

01 31-35, SI 113-123, (n=8).

Anterior border of clypeus arcuate, often with a shal-

low median concavity or truncation. Rear border of clypeus

more arcuate than worker's, yielding a lenticular general

form to the clypeus in full face view. Median portion of

clypeus rounded, subumbonate. In full face view, rear bor-

der of head straight with rounded corners. Mandibles with

smooth masticatory border, apical tooth often only indis-

tinctly set off from this border. Eyes smaller and less con-

vex than any other sympatric Paratrechina in full face view.

Eyes less than 1/3 the length of the head, and in some spec-

imens, barely protruding beyond the sides of the head. Head

narrower than in sympatric species.

Petiole shaped as in worker but broader. Genitalia

(Fig. 45) have the form that I consider most generalized

among nearctic species. Parameres in side view triangular,

about 2X as long as broad. In rear view, parameres weakly

curved mesad over their distal half. Volsella weakly

sclerotized,of simple structure; the digitus resembling

the boomerang shape of that of vividula but shorter, less

strongly bent and less tapered distally; cuspis thumb-like

in appearance and not convex in dorsal aspect, appearing

more truncate than rounded distally. Aedeagus resembling

paramere in shape but narrower at the base and less taper-

ing, less sclerotized and slightly decurved near the tip

(Fig. 45 does not show this clearly because it is a more

dorsal view). In rear view, the aedeagal lobes are nor-

mally closely appressed in preserved specimens and are

about equal in length to the parameres.

Vestiture, color and sculpture almost like those of

the workers with the usual differences in the vestiture of

the scapes and thorax. In colonies with fully sclerotized

workers at the lighter end of the color range, the males

tend to be lighter, as well. Parameres with about 25 de-

curved brown hairs, like those of vividula, except for their

darker color.

Material Studied

Syntype material (MHNG) consists of nine workers on

three pins collected in a forest at Faison, North Carolina,

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