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Revision of the Ant Genus Liometopum (Hymenoptera:

Formicidae)

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296 Sociobiology Vol. 52, No. 2A, 2009
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 297
298 Sociobiology Vol. 52, No. 2A, 2009
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 299

Revision of the Ant Genus Liometopum Mayr

(Hymenoptera: Formicidae)

by
Israel Del Toro1, José A. Pacheco2, and William P. Mackay1

ABSTRACT

Liometopum Mayr is a Holarctic ant genus with representatives in

North America, Europe, Asia Minor, India and Asia. This genus has never
been completely revised. The most comprehensive taxonomic study was
completed in 1905, and only included the North American representatives.
In this work we address the taxonomy of the eleven extant taxa and
provide descriptions, illustrations, maps and identification keys.
Additionally, Discriminant and Canonical Correlation Analyses were used
to assess morphometric variation. We recognize the validity of seven
species, three are found in North America and the remaining four are
found Europe, Asia Minor, India and Asia. The valid species are:
Liometopum apiculatum Mayr (= Formica masonia Buckley New
Synonym), L. lindgreeni Forel, L. luctuosum Wheeler, L. microcephalum
Panzer, L. occidentale Emery, Liometopum orientale Karavaiev, and
Liometopum sinense Wheeler (= L. sinense var. sericatum Wheeler New
Synonym,= L. dentimandibulum Chang He New Synonym). Liometopum
minimum Zhou is shown to be a synonym of Chronoxenus myops (Forel)
(New Combination).

INTRODUCTION

Ants in the genus Liometopum Mayr, known by the common name

of “velvety tree ants”, are widely distributed in the Holarctic region.
Bolton et al. (2006) list nineteen valid species-group names in the genus:
nine fossil species, ten extant species, and one extant subspecies. Three
extant species are currently recognized in the New World, Liometopum
apiculatum, L. luctuosum, and L. occidentale, with a fourth name, L.
masonium, treated as incertae sedis (Bolton, 1995). Two species, L.
microcephalum and L. orientale, are found in the Palearctic region. In
300 Sociobiology Vol. 52, No. 2A, 2009

Asia this genus is represented by L. dentimandibulum, L. lindgreeni, L.

minimum, L. sinense and L. sinense sericatum.
This group is in major need of revision, as it has not had a
taxonomic treatment in over 100 years (i.e., since Wheeler 1905). Shattuck
(1992) addressed the taxonomy of the three New World species in his
generic revision of the Dolichoderinae but did not review the Old World
species. Here we present a taxonomic revision that includes all eleven
extant representatives of the genus.
The nine fossil Liometopum species are L. antiquum Mayr, 1867,
L. eremicum Zhang, 1989, L. goepperti Mayr, 1868, L. imhoffi Heer, 1850,
L. lubricum Zhang et al. 1994, L. miocenicum Carpenter, 1930, L.
oligocenicum Wheeler, W.M. 1915, L. potamophilum Zhang, 1989, and L.
scudderi Carpenter, 1930. Of these species L. miocenicum and L. scudderi
are reported in the New World, while the remaining species are found in
the Palearctic region and one (L. oligocenicum) is found in Baltic Amber.
We do not address the taxonomy of these species in this work.
Our primary objective is to organize the taxonomy of the extant
members of this group and provide keys to the workers, males and gynes.
Additionally, illustrations are presented with complete morphometric data.
Liometopum is an extremely polymorphic genus; therefore, ranges as well
as averages of morphometric characters are included.

METHODS AND MATERIALS

Specimens Examined

A total of 1697 specimens were analyzed and identified to species.

Of the specimens analyzed 852 were dried and mounted, 845 were in
alcohol, 58 specimens were males, and 63 were gynes. In addition to
examining older museum material, new field collection of the three North
American species was conducted from October 2004 to May 2007. The
majority of the material collected consisted of foraging workers. Field
sites were distributed throughout the southwestern United States (Texas,
New Mexico, Arizona, and California) and northern and central Mexico
(Chihuahua, Michoacán, Tamaulipas, and Nuevo Leon).
Type specimens of L. luctuosum, L. sinense, L. sinense var.
sericatum, L. lindgreeni, and L. minimum were examined. Males and
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 301

gynes of L. apiculatum, L. luctuosum, L. occidentale, and L.

microcephalum were also studied. The following collections were used:
California Academy of Sciences Entomology Collection (CASC)
Museum of Natural History in Geneva, Switzerland (MHNG)
Shan-Yi Zhou Collection at Guangxi Normal University (GNUC)
Los Angeles County Museum of Natural History (LACM)
Museum of Comparative Zoology at Cambridge (MCZC)
The Smithsonian National Museum of Natural History (USNM)
Universidad Autónoma de Guadalajara (UAGC)
Universidad Autónoma de Querétaro (UAQC)
William and Emma Mackay Collection at the University of Texas
at El Paso (CWEM)
Morphological/Morphometric Assessment
Specimens were measured and described using a Wild stereoscope
equipped with a micrometer at magnifications of 6X, 12X or 24X.
Measurements were recorded with an accuracy of ± 0.01 mm. Cephalic
characters were taken in full face view; mesosomal characters were
measured in lateral view. An ocular grid was also used to produce
drawings. The following measurements and indices were used:
Total Length (TL)- Measured in lateral view from the most anterior
portion of the face to the most posterior point of the pygidium, when the
head is anteriorly directed, and the mesosoma and gaster are on the same
horizontal axis. This measurement can be inconsistent due to specimen
shrinkage and is influenced by how the head and gaster are flexed. This
character is included for general comparison of overall size, especially in
the reproductive castes. Head length and width are more accurate
measurements.
Head Length (HL)- In full face view, the midline length of the head
proper, from the anterior clypeal margin to the midpoint of a line drawn
across the posterior margin.
Head Width (HW)- In full face view, the maximum width of the
head, including the eyes.
Eye Length (EL)- The maximum length of the compound eye.
Eye Width (EW)- The maximum width of the compound eye
perpendicular to EL.
302 Sociobiology Vol. 52, No. 2A, 2009

Scape Length (SL)- The maximum length of the scape exclusive

of the basal condyle.
Petiole Height (PH)- In side view, the maximum distance from
the base of the petiolar sternite to the petiolar node apex.
Petiole Length (PL)- In side view, maximum length of the petiole
from the most anterior region of the peduncle to the most posterior
region of the petiole.
Wing Length (WL)- In lateral view, maximum length from the
base to the apex of the forewing.
Cephalic Index (CI)- HW/HL X 100.
Scape Index (SI)- SL/HW X 100.
Petiolar Index (PI)- PL/PH X 100.
Measurement ranges followed by means (in parentheses) are
reported for each species. Additional morphological characters that
were analyzed include:
Number of ommatidia- Numbers of ommatidia are reported in the
species description but not accurately illustrated, due to their small size.
Antennal structure- The degree to which the scape surpasses the
posterior cephalic margin is included. Length and width of individual
funicular segments (especially the first funicular segment) are of
taxonomic importance.
Clypeal margin- The form of the anterior clypeal margin was
considered. Species vary in whether the anterior clypeal margin is
straight or weakly concave.
Mandibular Structure- The mandible is finely punctate.
Mandibular teeth along the basal and masticatory margin as well as
relative sizes of the apical, subapical and basal teeth were considered.
Mesosoma Shape and Structure- The degree of convexity along
the dorsal mesosomal margin is of taxonomic importance, varying
from straight to extremely convex. Additional mesosomal characters
considered are propodeal shape along the dorsal and posterior margins,
the definition (angulate, or round) of the metapleural angle and the
degree of indentation of the metanotal suture.
Petiolar shape- Observed in lateral view, the shape of the petiolar
apex varies among species (angulate or round). The petiole may be
inclined anteriorly or completely vertical.
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 303

Coloration- This is an extremely variable character, of little

taxonomic importance. However coloration ranges are noted in the
corresponding species descriptions.
Pilosity- The degree of pilosity is variable among species,
particularly between L. luctuosum and L. apiculatum. All species are
covered in appressed pubescence but pilosity in certain areas (posterior
cephalic margin, dorsal surface of the pronotum and the dorsal petiolar
surface) is of taxonomic importance.
Wings- Wing characters were considered for the males and the
gynes of L. apiculatum, L. luctuosum, and L. occidentale. There is
much variation that occurs in venation patterns among species.
Forewing cells of taxonomic importance are the radial, submarginal
and discoidal cells, and the hind wing medial cell. Hind wing hamuli
varied between L. apiculatum and L. luctuosum (Figures 62-68).
Male Genitalia- In treating males, genitalia were dissected. The
genital capsule was extracted and placed in 10% KOH solution for a
period of 24 hours. When the capsule was soft and easily
disarticulated, components were dissected, and illustrated (Figs. 36-
54). Dissected components were placed in genital capsule vials and
preserved in glycerin. The vial was pinned with the corresponding
specimen and returned to the source collection. Characters of
taxonomic importance include the shape of the posterior ventral
paramere margin, posterior volsella margin, posterior subgenital plate
margin, and the number of aedeagal denticles along the aedeagus
ventral margin.
Species delineation
We employ the Biological Species Concept as defined by Mayr
(1942, modified in 1995), which states that species are groups of
interbreeding natural populations that are reproductively isolated from
other such groups. Our primary criteria for assessing the Biological
Species Concept are similarities and differences in morphology, discrete
variation in zones of sympatry, and ecological differences.
Statistical Analysis
Discriminant analysis was conducted on the data matrix to identify
statistically significant characters (p<0.05), using SYSTAT 12 (Table 1
and Table 2). For taxonomic and determination processes, Discriminant
304 Sociobiology Vol. 52, No. 2A, 2009

Analysis is used to determine which characters best explain an

individual’s species membership (Moder et al., 2007). The data matrix
was organized with species as columns and morphometric variables as
rows. Canonical Correlation Analysis can be seen as the problem of
finding vectors for two sets of variables such that the correlation between
the projections of the variables onto these basis vectors are mutually
maximized (Hotelling 1936). Following a forward stepwise variable
selection using discriminant analysis (F to enter >4.0, F to exclude <3.9),
characters were plotted using canonical correlation plots (Fig. 106, 107).
Digital imaging and Distributional Mapping
Representative specimens of each of the species were
photographed using a Leica MZ16 microscope equipped with Auto-
Montage software at the MCZC.
Distribution data were mapped using ArcView 9.1. Only material
examined was plotted on distribution maps.

RESULTS/ TAXONOMY
Generic Level Diagnosis:
Workers: Polymorphic, 3.3-5.5 mm in total length. Mandibles with
7-10 teeth along masticatory margin, 3-5 teeth along basal margin, apical
tooth larger. Moderate concavity along anterior clypeal border, lacking
teeth. Strong concavity along posterior margin of head. Antenna 12
segmented, scape may or may not surpass posterior margin of head.
Majors usually possess ocelli. Compound eyes with 90-140 ommatidia. 6
segmented maxillary palp, 4 segmented labial palp. All surfaces covered
in dense, pubescence, largest hairs on dorsal surface of pronotum. Cuticle
smooth. Mesosoma broadly convex (similar to Camponotus spp.);
metanotal groove reduced to suture. Petiole scale-like, usually leaning
anteriorly, subpetiolar process not developed. Legs long, slender, 1 tibial
spur present on each tibia. First gastral tergite largest, usually not
concealing petiole. Coloration variable, usually reddish to dark brown.

Gynes: Large, reaching up to 16.5 mm in total length. Mandibles

punctate, 7-10 masticatory teeth, 2-3 teeth on basal margin. Clypeus with
broad, shallow concavity, lacking teeth. Posterior margin of head
moderately concave. 12 segmented antenna, scape reaching posterior
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 305

margin of head. Compound eyes with 100-145 ommatidia. Three ocelli

present and equal in size. 6 segmented maxillary palp, 4 segmented labial
palp.Cuticle smooth. Long erect hairs on dorsal surface of head. Highest
hair density on dorsal surface of propodeum. Petiolar scale-like, bidentate,
ventral margin keel-shaped. Subpetiolar process not developed. Gaster
extremely large, up to half total length. First gastral tergite may conceal
petiole, tergites 1-3 of approximately equal width when seen in lateral
view, tergites 4-5 smaller. Legs, long and slender, single tibial spur
present on each tibia. Forewing, radial, submarginal and discoidal cells
closed. Hind wing with two closed cells, 15-24 hamuli. Concolorous, light
brown to black.
Males: Variable in size, 8.8-13.5 mm in total length. Head small,
quadrate, posterior margin weakly concave, ocelli present, covered in
long, decumbent hairs. Mandibles punctate, masticatory margin with 8-12
teeth, basal margin with 0-4 small teeth. Clypeus straight, lacking teeth.
Antenna 13 segmented, scape never surpassing posterior margin of head.
Eyes large, about 1/3 of total head length. Palpal formula 6,4. Mesosoma
smooth, metanotal suture weakly indented. Petiole scale-like, bidentate at
apex. Genitalia large, genital capsule 2/3 exposed. Parameres large,
covered in long, decumbent hairs, sometimes concave along ventral
margin. Volsella with posterior margin straight or weakly concave.
Aedeagus with 15 to 25 denticles on ventral margin, posterior margin
rounded or triangular. Subgenital plate notched along posterior margin.
Forewing with submarginal and discodial cells closed. Hind wing with
medial cells closed, all other cells open, 13-22 hamuli. Concolorous dark
brown to black.
Larvae: (From Wheeler & Wheeler 1951) “Body and head
practically naked, body hairs exceedingly few; short and widely scattered.
Head hairs very few, exceeding minute, widely scattered. Antennae rather
large. Posterior surface of labrum with two subventrical rows of sensilla
near the middle and near the ventral border, short transverse rows of
minute spinules. Mandible small. Maxillae with a lateral patch of isolated
spinules between palp and galea; spinules with stout base and needle-like
apex; palp a low convexity bearing one large and three small sensilla.
Labium with spinules and a few sensilla on the anterior surface, palp a low
convexity bearing one large and three small sensilla.”
306 Sociobiology Vol. 52, No. 2A, 2009

Synopsis of Liometopum species

L. antiquum Mayr, 1867: 60 *
L. apiculatum Mayr, 1870: 961 Nomen Protectum
=L. masonium Buckley, 1866: 165 Nomen Oblitum
L. eremicum Zhang, 1989: 281 *
L. goepperti Mayr, 1868: 56 *
L. imhoffii Heer, 1850: 138 *
= Formica schmidtii Heer, 1850: 138
L. lindgreeni Forel, 1902: 293
L. lubricum Zhang, Sun & Zhang, 1994: 165 *
L. luctuosum Wheeler, 1905: 325
L. microcephalum Panzer, 1798: 2
= Formica austriaca Mayr, 1853:144
L. miocenicum Carpenter, 1930:46 *
L. occidentale Emery, 1895: 330
L. oligocenicum Wheeler, 1915: 95 *
L. orientale Karavaiev, 1927: 339
L. potamophilum Zhang, 1989: 280 *
L. scudderi Carpenter, 1930: 47 *
L. sinense Wheeler, 1921: 541
= L. sinense sericatum Wheeler, 1921: 541 New Synonymy
= L. dentimandibulum Chang & He, 2002: 110 New Synonymy
Name excluded from Liometopum
Chronoxenus myops (Forel, 1895:471)
= L. minimum (Zhou, 2001:557) New Combination, New
Synonymy
* indicates species known only from fossils

KEYS TO THE SPECIES OF LIOMETOPUM BASED ON WORKERS,

MALES, AND GYNES

Key to Liometopum Workers

1. Long erect hairs (0.30 mm) present on dorsum of gaster (Fig. 2) mixed
with shorter erect hairs (0.12 mm), some hairs nearly as long as those on
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 307

pronotum, pilosity may or may not cover entire gastral surface; mesosoma
evenly convex, not depressed at metanotal suture (Fig. 2)………………..2
-Erect hairs on dorsum of gaster (Fig. 12) short (0.10 mm), or absent and
of approximately equal length or usually shorter than erect hairs on pro-
notum; mesosoma convex but strongly depressed at metanotal suture (Fig.
12); Washington State to West Texas and Central Mexico……. luctuosum
2(1). Antennal scape surpassing posterior margin of head by at least twice
maximum thickness of scape (Fig. 1, Fig. 7)…………………………...…3
-Antennal scape of largest workers surpassing posterior margin of head by
amount that does not exceed maximum thickness of scape (Fig. 11)…….4
3(2). When seen from dorsal view, petiole coming to sharp angular apex;
in lateral view metapleural angle straight, posterior propodeal face vertical
(Fig. 2); entire gastral surface covered in pilosity; common from
Southwestern United States, west to West Texas, south to Southeastern
Mexico (Quintana Roo)……………………………………..… apiculatum
-When seen from dorsal view, petiole flat and straight at node; in lateral
view metapleural angle rounded, dorsal propodeal face rounded (Fig. 8);
gaster shiny with long appressed pilosity, not covering entire gastral
surface; known only from type locality, India, Assam
Province……………………………………………………….....lindgreeni
4(2). Bicolored, head and gaster darker than mesosoma; from dorsal view
petiolar apex sharply angular or rounded; USA Pacific Coast south to
northern Mexican Pacific Coast or throughout European continent……...5
-Concolorous dark brown to light brown; from dorsal view petiolar apex
always sharply angular; common throughout central China…….…sinense
5(4). Suberect pilosity present along posterior cephalic margin (Fig. 17,
Fig. 23); scapes with short erect hairs at junction of scape and funiculus;
first funicular segment equally long as wide; in posterior view, petiolar
node sharply angular (Fig. 31); in New World common along Pacific coast
from northwestern USA to northwestern Mexico, in Old World common
throughout southern Europe, east to Asia Minor, and south to Israel….....6
-Without decumbent pilosity along posterior margin of head (Fig. 29);
scapes with two notable long erect hairs at junction of scape and funiculus;
first funicular segment two times longer than wide; petiolar node rounded
when seen from posterior view (Fig. 30); Russia and northeastern
Europe…………………………………………………………..…orientale
308 Sociobiology Vol. 52, No. 2A, 2009

6(5). Larger, Eye Length (EL) 0.27-0.31 mm on average ~0.28 mm; Scape
Length (SL) 1.10-1.25 mm on average ~1.16 mm; southern European
continent, west to Asia Minor, south to Israel…………......microcephalum
-Smaller, EL 0.20-0.25 mm on average ~0.24 mm; SL 0.75-1.15 mm on
average 1.03 mm; Pacific West Coast from Washington (USA) south to
Baja California (Mexico)……………………………………....occidentale

Key to Liometopum Gynes (L. orientale, L. lindgreeni, and L. sinense

not included)

1. Head length (HL) greater than 2.20 mm; with multiple long (0.20 mm),
erect, suberect, subdecumbent or decumbent hairs along the posterior
cephalic margin (Fig. 3)……………………………………......apiculatum
-HL less than 2.00 mm; hairs on posterior cephalic margin short (0.08
mm), erect and suberect (Fig. 13)………………………………………... 2
2(1). Glossy dark brown appearance; medial posterior cephalic concavity
well defined, HL usually less than 1.65 mm; well defined metanotal suture
(Fig. 14); 7-8 masticatory teeth; California to West Texas, south to central
Mexico………………………………………………………...... luctuosum
-Dull, dark yellow to light brown appearance; HL 1.63-1.85 mm;
metanotal suture not clearly depressed (Fig. 20, Fig. 26); 8-10 masticatory
teeth; in New World commonly collected along Pacific coast from
northeastern USA to northeastern Mexico, in Old World commonly
collected throughout southern Europe, east to Asia Minor, and south to
Israel…………………………………………………………………….....3
3(2). Medial cephalic margin straight, not concave (Fig. 25); collected
only in New World from Washington State (USA), south to Baja
California (Mexico)………………………………………….... occidentale
-Medial cephalic margin clearly concave (Fig. 19); collected only in the
Old World, throughout southern Europe west to Asia Minor, south to
Israel…………………………………………………….... microcephalum

Key to Liometopum Males (L. lindgreeni and L. sinense not included)

1. 20 or more hamuli on anterior edge of posterior wing (Fig 62 B),
anterior wing length greater than 12.25 mm (Fig 61); dark brown to black;
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 309

TL greater than 11.3 mm (consistently larger than all other species);

Arizona to West Texas to Southeastern Mexico…………….....apiculatum
- 14-18 hamuli (Fig. 66), anterior wing length less than 12.25 mm (Fig.
65); light brown to dark brown; TL does not exceed 11.1 mm; in New
World, California, to West Texas to Central Mexico, in Old World
throughout European Continent west to Asia Minor, south to Israel……..2
2(1). Ventral margin of paramere straight (Fig. 34), not concave; collected
in Russia and northeastern Europe………………………………..orientale
-Ventral margin of paramere concave (Fig. 46); in New world collected
from Northwestern United States west to West Texas south to Central
Mexico, in Old World collected throughout southern Europe, east to Asia
Minor, and south to Israel………………………………………………....3
3. Aedeagus ventral margin emarginate posteriorally to aedeagal denticles
(Fig. 48); volsella dorsal margin straight (Fig. 49); collected throughout
southern Europe, east to Asia Minor, and south to Israel.....microcephalum
-Aedeagus ventral margin rounded posteriorally to aedeagal denticles (Fig.
53), volsella dorsal margin concave or straight (Fig. 54), collected in New
World only (Northwestern United States southeast to West Texas, south to
Central Mexico)…………………………………………………………...4
4. Volsella dorsal margin concave (Fig. 54), 15-18 aedeagal denticles
covering 1/2 of ventral aedeagal margin, posterior aedeagal margin
rounded (Fig. 53)........................................................................ occidentale
- Volsella dorsal margin straight, more than 20 aedeagal denticles
covering 2/3 of ventral aedeagal margin, posterior aedeagal margin
triangular (Fig. 43)…………………………………………….... luctuosum

SPECIES SYNOPSES

Liometopum apiculatum Mayr

Figs. 1-6, 35-39, 61-64, 69, 75-80

Formica masonium Buckley 1866: 165, worker, USA: Texas, Fort

Mason, New Synonymy, Nomen Oblitum

Liometopum apiculatum Mayr 1870: 961, worker, MEXICO; Emery

1895: 331, queen, USA: Texas; Wheeler 1905: 324, male, USA: Arizona;
Wheeler & Wheeler 1951: 181, larva, USA: California, Nomen Protectum
310 Sociobiology Vol. 52, No. 2A, 2009

Diagnosis

Worker: This species is concolorous light brown to dark brown. The

antennal scape is long (1.2 mm) and slender, with short, erect and appressed
hairs, and surpasses the posterior margin of the head by at least twice the
maximum thickness of the scape. The dorsum of the mesosoma is convex
with appressed hairs on all surfaces; the longest hairs are on the dorsal surface
of the pronotum.

Gyne: Gynes are concolorous dark brown to black. They are

extremely large at over 15.0 mm in total length. The mesosomal dorsum is
covered with many long (0.24 mm), erect hairs and with dense appressed
pubescence. The metanotal suture is distinct but does not break the smooth
dorsal mesosomal margin.

Male: Males of this species are concolorous black and smaller in total
length (11.3 mm) than gynes, but are larger than males of other Liometopum
species. The mesosoma is convex and smooth. The hind wings of males have
20 or more hamuli; the anterior wing length is greater than 12.25 mm. The
volsella has dense erect pilosity along the ventral margin and is finely
denticulate along the posterior margin (Fig. 39). The aedeagus has more than
20 denticles along the ventral margin (Fig. 38).

Comments: The worker of this species can be separated from that of

L. luctuosum based on greater hair density and scape morphology. In L.
luctuosum, the scape is wider (0.22 mm) and does not extend past the
posterior margin of the head by at least twice the maximum thickness of the
scape. The L. luctuosum worker is smaller in total length (3.6 mm), with the
mesosoma depressed at the metanotal suture. Liometopum apiculatum
workers can be separated from those of L. occidentale based on color, being
concolorous dark brown to black (L. occidentale is bicolored). Liometopum
occidentale is found along the Pacific coast of United States and Northern
Mexico. Gynes of this species are easily separated from those of L. luctuosum
and L. occidentale based on their larger size, (greater than 14 mm in total
length); the other species’ queens do not exceed 12 mm in total length. Males
of L. apiculatum are also consistently larger than those of the other two
species. Additionally males can be separated by having more than 20 hamuli
on the hind wing (fewer than 15 on the other two species) and an anterior
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 311

wing length that is greater than 12.25 mm (less than 11.10 mm in the other
two species).

Caste Descriptions

Workers (n=10)

TL 4.15-5.25 (4.64), HL 0.93-1.23 (1.19), HW 1.20-1.56 (1.42), EL 0.21-0.25

(0.23), EW 0.14-0.20 (0.15), SL 1.06-1.37 (1.21), PH 0.31-0.55 (0.45), PL
0.22-0.32 (0.26), CI 103-135 (119), SI 93-106 (102), PI 46-72 (58)

HEAD: Wider than long, posterior medial concavity well defined; largest
workers with ocelli. Eyes large, medial rather than lateral, 110 to 125
ommatidia. Scapes long and thin, surpassing posterior margin of head by at
least two times maximum thickness of scape. Funiculus 11 segmented, first
and last segments largest. Anterior clypeal margin straight. Mandibles
punctate, 8 teeth along masticatory margin, 4 to 5 smaller teeth on basal
margin. MESOSOMA: Slightly convex. Pronotum convex, from dorsal view
widest point narrower than maximum head width. Metanotal suture weakly
defined, not breaking dorsal margin of mesosoma. Propodeal spiracle small,
round. Petiole scale-like, sharp angular apex. Legs long, slender. GASTER:
First gastral tergite partially overlapping petiole. COLORATION AND
PILOSITY: Concolorous light brown to dark brown. Coloration similar in
members of same series. Cuticle covered in dense appressed pubescence to
erect pilosity. Subdecumbent to erect pilosity along posterior margin of head.
Antennae with short, appressed pubescence. More than 20 long erect hairs
along anterior clypeal margin. Mesosoma completely covered in pubescence.
Longest erect hairs on pronotal dorsum (12 or more hairs). Suberect
pubescence covering petiolar anterior surface. Pilosity coloration light grey to
golden brown.

Gynes (n=5)

TL 14.50-16.25 (15.23), HL 2.30-2.43 (2.37), HW 2.63-3.00 (2.76), EL 0.45-

0.50 (0.49), EW 0.38-0.50 (0.43), SL 1.63-1.80 (1.73), PH 1.15-1.65 (1.37),
PL 0.80-1.00 (0.90), CI 111-130 (117), SI 70-77 (73), PI 49-70 (67), WL
13.25-17.25 (15.30)
312 Sociobiology Vol. 52, No. 2A, 2009

HEAD: Wider than long, strong posterior medial concavity apparent. Ocelli
present, medial ocellus circular, lateral ocelli oval-shaped. Eyes located more
laterally than in workers, large oval-shaped with over 150 ommatidia. Scapes
not surpassing posterior margin of head. Anterior clypeal margin weakly
concave. Mandibles punctate, 8-10 teeth on masticatory margin, 3-5 teeth on
basal margin. MESOSOMA: Strongly convex, smooth, lacking sculpturing,
pronotum dorsal face straight. Mesonotum strongly arched, from dorsal view
mesonotum globular, metanotal suture distinct. Legs long, slender. Forewing,
radial, submarginal and discodial cells closed. Hind wing medial cells closed
with more than 20 hamuli. Propodeal dorsal margin round, spiracle large,
circular. Petiole scale-like, from lateral view with sharp angular node, from
dorsal view apex bidentate, leaning anteriorly. GASTER: Gaster large, ½ of
total length, overhanging petiole. Gastral tergites 1-3 equal width in lateral
view, 4 and 5 decreasing in total size. COLORATION AND PILOSITY:
Concolorous, dark brown to black. Dense pilosity covering entire cuticular
surface. Cephalic margins covered in subdecumbent pubescence. Antennae
with dense, appressed pubescence. More than 25 long, erect hairs surpassing
anterior clypeal margin. Mesosoma covered in erect pilosity. Dorsal
mesonotal surface with lower hair density. Dense pilosity covering propodeal
spiracle. Lacking pubescence on posterior petiolar surface. Dense pubescence
covering entire gastral surface. Pubescence light yellow to dark brown.

Males (n=5)

TL 10.13-13.50 (11.56), HL 1.28-1.50 (1.38), HW 1.58-1.67 (1.67), EL 0.45-

0.50 (0.48), EW 0.35-0.43 (0.39), SL 0.60-0.75 (0.69), PH 1.00-1.25 (1.08),
PL 0.50-0.65 (0.54), CI 105-132 (122), SI 43-55 (50), PI 44-55 (50), WL
12.25-13.75 (13.13)

HEAD: Wider than long, lacking concavity along posterior cephalic margin.
Ocelli present, equal in size, oval-shaped. Eyes lateral with more than 140
ommatidia. Scape never surpassing posterior margin of head. Funicular
segments equal in length. Anterior clypeal margin straight. Mandibles
punctate, 9-11 teeth on masticatory margin, 1-2 teeth on basal margin.
MESOSOMA: Strongly convex, mesonotum arched. Legs long and slender,
femur 3/4 length of tibia and tarsus combined. Submarginal and discodial
forewing cells closed. Discodial cell quadrate. Hind wing with more than 20
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 313

hamuli. Dorsal surface of propodeum forming angle. Petiole scale-like with

sharp, angular node, straight, not leaning anteriorly. GASTER: First gastral
tergite larger than 2-5. Genital capsule large, parameres large, ventral margin
of paramere weakly concave. Volsella finely denticulate along posterior
margin. Aedeagus with more than 20 triangular denticles along ventral
margin, rounded at ventral margin apex. Posterior margin of subgenital plate
medially emarginate. COLORATION AND PILOSITY: Concolorous dark
brown to black. Dense appressed to erect pilosity covering all surfaces.
Lateral surfaces of head with dense suberect and decumbent pilosity. Dorsal
surface of mesonotum with most dense pubescence. Appressed and suberect
pubescence covering gastral surface. Genital capsule lacking pubescence,
except on posterior margin. Parameres with long, suberect hairs. Volsella
with erect pilosity along ventral margin. Subgenital plate covered with
decumbent to erect pilosity.

Distribution
Southwestern United States & Northwestern to Southeastern Mexico,
Quintana Roo.

Type Material: No type material was examined

Material Examined

MEXICO- Chihuahua: 100 m above Rio Urique, 84 S of Creel, 28.ii.66, J.

Reddell, W. Bell. Coahuila: Arteaga, Cañon San Lorenzo, 25°23'28.0"N
100°42'49.8"W, WY 57 nr. Lirios, 10.vii.1973, R. Snelling. Colima: Volcan
de Colima. Durango: Durango, Revolcaderos, 21.vii.1965, E.M. Fisher.
Guanajuato: Camino 57, Km. 306, Rancho Jardin, 10.viii.1965, Cornell
University Mexico Field Party. Hidalgo: Grutas de Xoxa, 19.vii.1965, J.
Reddell; Guerrero Mill, W.M. Mann. Mexico: Piñón del Marquis, near
Mexico City, 27.iii.1933; Teozulcan, 9.vi.1929, W.M. Wheeler. Michoacán:
South of Zitacuaro, 9.vii.1965, R. Snelling; 18.2 km. South Nupecuato; 12.3
miles E. Morelia, R. Snelling; Zacapu, vi.45, A.J. Sharp. Nuevo Leon: Near
Monterrey, Mesa de Chipinque, 16-18.vii.1965, Cornell University Mexico
Field Party; Galeana, Cerro El Potosí, 24°53'19.9"N 100°11'58.0"W; 30.5 km
east of Arroyo San Pedro Garza Garcia,W.P. Mackay; Parque Chipinque,
Camino al Pinar, 25°36'05.5"N 100°19'57.4"W; 5 mi W Iturbide, 9.xi.46,
314 Sociobiology Vol. 52, No. 2A, 2009

Ross Collection. Querétaro: Municipio Queretaro, La Barreta, 20°49.35'N

100°30.939'W, M. Rocha. Quintana Roo: Lazaro Cardenas, El Eden
Ecological Reserve, 25 km NNE Leona Vicario Det. W.P. Mackay 2002. San
Luis Potosí: 10 miles N.E San Luis Potosí, vii-22-1954, J. Bequaert; Rio
Verde, Valle de los Fantasmas, 22°03'76.9"N 100°36'83.6"W. Sinaloa: 6.5
mi E. Potrillos, 21.viii.1964, Hwy. 40, R. Schlinger. Tamaulipas:
Miquihuana, Camino a San Jose del Llano, 23°32'27.7"N 99°48'02.8"W, M.
Bolaños. Zacatecas: Pinos Los Alpes, Nopalera, M. Bolaños. USA-
Arizona: Oracle, 16.iii.1919, W.M.Wheeler; Ash Spring, 6 miles SW Portal,
31.iii.1966, B. Vogel; Hilltop, Portal, 20-27.vii.1967, G. Alpert; Mt.
Lemmon, 36 miles N. of Tucson, 30.vii.1965, Freitag & Gibson; Montezuma
Pass, Coronado Nat. Mem., 9.ix.1965; Coronado Nat. Memorial, 31.viii.1972;
Madera Canyon, Santa Rita Mts., L. Martin; Ramsey Canyon, Huachuca
Mts., Mann; Huachuca Mts. 21.vii.1937, Knull; Cooper Canyon, 8.1 Miles
SE Sunnyside, 26.viii.1972, R. Snelling; Chiricahua Mts., Sycamore Spring;
1 mi. N. Paradise; 12 miles NW Portal, Carr Canyon, Huachuca Mts.,
23.viii.1972, R. Snelling; 5 miles W Portal, 30.vi.1970; Stewart Camp, 2
miles W Portal, Chiricahua Mts., 28.viii.1965; Dragoon Mtns., Cochise
Stronghold, 21.vii.1993, S.P. Cover; Black Hills, 16.9 mi. ENE RT. 89 on Rt.
89 A 34, 41.47”N, 112, 10.13”W, 20.viii.1998, S.P. Cover. Colorado:
Manitou, 17.vii.1903; Garden of the Gods, 1.viii.1941; Canon City, 2.vi.1900,
W.M. Wheeler; Boulder, 7.vii.1927, W.S. Creighton; Boulder, Gregory
Canyon, 27.iii.1932, L.F. Bayns; Colorado Springs and vicinity, W.M.
Wheeler. New Mexico: Alamogordo, G. Krockow; Cimarron Canyon,
26.vii.1962, A.C. Cole; Silver City, 15.viii.1952; Bernardo,10.ix.1951, A.C.
Cole; Bandelier 45 km NE Las Cruces, W.P. Mackay; Bandelier Natl. Mon,
30.vii.1952, A.C. Cole; Silver City, on state 61, 14.viii.1952; Cherry Creek
Recreation Area, near Silver City, 9.vii.1963, P.J. Spangler; Stratton,
27.vii.1917, W.M. Wheeler; Hwy. 78 at Arizona State Line, Gila National
Forest, 7.ix.1972, R. Snelling; Catwalk, W.P Mackay; 15 k NW Datil, W.&E.
Mackay; Ox Spring Canyon, W.&E. Mackay; Snow Lake, W.&E. Mackay;
Gila Mts., W.&E. Mackay; Iron Creek, W.&E. Mackay; Wright’s Cabin,
W.&E. Mackay, 77 k E Silver City, W.&E. Mackay; 88 k E. Silver City,
W.&E. Mackay; 9 k NW White Signal, W.&E. Mackay; Los Alamos, W.&E.
Mackay; Rio Grande, W.&E. Mackay; Abiquiu Dam, W.&E. Mackay; 26 k S
Cuba, W.&E. Mackay; Magdalena Mts., Water Canyon, W.&E. Mackay;
Sacramento Mts., Ruthven, W.&E. Mackay; Highrolls, 12.vi.1902. Texas:
Franklin Mts., R. Worthington; Ft. Davis Mts., 19.vii.1933, Creighton; Ft.
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 315

Davis, 10.vi.1902; McKitrick Canyon, 3.iv.69; 30 mi S. Ft. Stockton, J.M.

Tenorio; Big Bend Nat. Park, Juniper Cyn., 3.vi.1970; Green Gulch, Big
Bend Nat. Park., 5.vii.1970, O’Brien; Davis Mts., 3.vi.1970, C.W. O’Brien;
Chisos Basin, Big Bend Nat. Park, 18.vii.1966, A.E. Lewis; Chisos Mts.,
13.vii.1933, W.S. Creighton; Paisano Pass, 11.xii.1901.

Habitat
This species nests in a wide variety of substrates in oak forests, at
elevation ranges of 1000 m to 2500 m. The prime habitat is oak forests
around an elevation of 2000 m, but can be found in higher elevations in areas
of pinyon pine, up to ponderosa pine and riparian sites. It also nests in
creosote bush scrub and grasslands. At higher elevations, this species
abundance decreases and is replaced by L. luctuosum.

Biology
This species is commonly found nesting in dead logs, under stones,
and in decaying stalks of Yucca spp. (Miller, 2007). Nests have even been
collected in glass containers and rubber tires. It is polydomous with segments
of nests scattered over the landscape. Nests from Colorado Springs, Colorado,
are carton nests, but are usually difficult to reach, as they are usually deep
under heavy boulders or large trees (Wheeler 1905; Gregg 1963). It is the
dominant ant in most of the oak forests in the southwestern United States and
can be easily found foraging on the sides of oak trees. Colonies are variable in
size ranging from a few hundred workers to 85,000 workers per colony
(Ramoselorduy & Levieux, 1992). Behavioral dominance has also been
observed in this group (Andersen, 1997). This species is opportunistic, but
larger colonies are predaceous, while other colonies have been observed
foraging for dead insects and tending Homoptera (Shapley, 1920).
The primary mode of defense is to emit a strong odor and attack
aggressively in large numbers. Liometopum apiculatum has a mutualistic
relationship with the cholla cactus, Opuntia imbricata, and uses its aggressive
behavior to protect the cactus from herbivores and seed predators and collects
the extrafloral nectars produced by the plant (Miller, 2007). Workers actively
forage from March to September, and cover a range up to 580 m2 (Mackay &
Mackay, 2002; Ramoselorduy & Levieux, 1992). Sexuals occur in nests from
May to August. Males and gynes were collected on the ground in June to
August; queens were commonly collected in July and August under stones,
cow manure or logs.
316 Sociobiology Vol. 52, No. 2A, 2009

Inquilines appear to be especially common in the nests of this species.

The small cricket, Myrmecophila spp. occurs in the nests throughout its
range. Staphylinids including Sceptobius dispar Sharp (Fig. 103) and
Dinardilla liometopi Wasmann (Fig. 104) are also found in nests, often taking
food from the ants. Navarette et al., (2006) provide the most comprehensive
analysis of these inquiline guests. Other known myrmecophiles include
Liometophilus manni (Fig. 105). Dinardilla species often directly interact
with L. apiculatum while Sceptobius beetles tend to avoid direct contact with
their host (Danoffburg, 1994).

Species Summary
We consider Liometopum masonium (Buckley, 1866) to be a synonym
of L. apiculatum. Although L. masonium is the senior name, L. apiculatum
has been in wide usage for the past 141 years. For L. apiculatum to qualify for
nomen protectum status, L. masonium must not have been used as a valid
name after 1899 (ICZN 23.9.1.1) and L. apiculatum must be mentioned in at
least 25 works, published by at least 10 authors since 1957 encompassing a
span of not less than 10 years (ICZN 23.9.1.2). Liometopum masonium has
only been mentioned as insertis sedis since 1899 (Shattuck, 1994). More than
25 works, by more than 10 authors in the preceding 50 years refer to this
species as L. apiculatum not L. masonium. Examples of such works include:
(Gregg, 1963; Gulmahamad, 1995; Mackay et al. 1988, Mackay & Mackay,
2002; Merickel & Clark, 1994; Moreno-Garcia et. al 2003; Shattuck, 1992,
1994; Kupyanskaya, 1988; Wheeler & Wheeler 1973, 1986; Bolton 1995,
Bolton et al., 2006; Miller, 2007; Navarette et al., 2007; and Dannoffburg,
1994). Therefore, L. masonium is considered as nomen oblitum and L.
apiculatum as nomen protectum as ICZN criteria proposed by Article 23.9.1.1
and 23.9.1.2 are both met.
Even though the types for L. masonium and L. apiculatum could not be
located, we are confident (based on the original descriptions) that these two
taxa are identical. The type description of L. apiculatum is somewhat short
and vague. However, Wheeler, (1905) provides extremely accurate
descriptions of the worker, gyne and male. These descriptions consider
multiple specimens from different localities and provide an accurate
definition of the morphology used to distinguish L. apiculatum. Based on this
description we were able to easily identify L. apiculatum.
The vague description presented by Buckley is indicative of L.
apiculatum, which we have collected several times in the same region of
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 317

Texas (Fort Mason) where the L. masonium holotype specimen was collected.
Many of the characters mentioned by Buckley are characteristic of L.
apiculatum (cuticular coloration, number of mandibular teeth, cephalic
shape). Additionally, most of these characters provide a generalized
description of all species within the genus.
We recognize the uniqueness of the specimen from Southeastern
Mexico (Quintana Roo), which represents a disjunction in distribution. We
have extensively collected throughout the region of the Isthmus of
Tehuantepec and the Yucatan Peninsula and have not found any other
Liometopum specimens. Dolichoderus bispinosus is the dominant
Dolichoderine species in this geographic region. The specimen from Quintana
Roo may represent an isolated or relic population.

Liometopum lindgreeni Forel

Figs. 7-8, 70, 99-100

Liometopum lindgreeni Forel 1902: 293, worker, INDIA: Assam, Debrugarh.

[holotype examined]

Diagnosis
Worker: This species is only known from the holotype collected in
Assam, India. It is a concolorous light brown specimen with a dull luster on
the cuticle. The cuticle is covered in short pubescence with a low pilosity
density on the dorsal surface of the gaster. The antennal scapes surpass the
posterior margin of the head by at least two times the maximum thickness of
the scape. The first funicular segment is three times longer than wide. The
anterior clypeal margin is straight with 10-12 hairs surpassing the margin.
The pronotum is weakly convex, the metanotal suture is weakly concave. The
petiole is scale-like, and flat at the apex when seen from posterior view. The
longest decumbent hairs are found on the anterior petiolar surface.

Comments: This species can be separated from other Old World species by
having a narrow head (1.20 mm wide). Liometopum lindgreeni is unique in
that dense pilosity does not cover the entire cuticular surface. The petiole is
also unique when seen from posterior view, in that it is flat and emarginate at
the apex, in the other species the petiolar node forms a sharp angle.
318 Sociobiology Vol. 52, No. 2A, 2009

Gyne and Male: Unknown

Caste Description

Worker (n=1)

TL 5.35, HL 1.06, HW 1.20, EL 0.24, EW 0.15, SL 1.03, PH 0.35, PL 0.23,

CI 129, SI 96, PI 64
HEAD: Posterior medial concavity well defined. Lacking ocelli. Eyes oval-
shaped, medial rather than lateral, with 115 ommatidia. Antennal scapes
slender (0.10 mm) surpassing posterior cephalic margin by more than two
times maximum scape width. First and last funicular segments longest (0.18
mm), segments 1-5 and 11 longer than wide, segments 6-10 equally wide as
long or wider than long. Anterior clypeal margin straight. Mandibles finely
punctate, 10 teeth on masticatory margin, 2 reduced denticles on basal
margin. MESOSOMA: Anteriorly convex, posteriorly straight, in dorsal view
widest at posterior edge of pronotum. Pronotum, arched. Promesonotum
strongly convex. Metanotum straight, weakly depressed at metanotal suture.
Legs long, slender, tibia 2/3 length of maximum femur length. Propodeal
dorsal margin weakly convex. Posterior metapleural angle absent, dorsal
posterior face of propodeum rounded, spiracle small, oval-shaped on posterior
dorsal lateral surface. Petiole scale-like from posterior view, flat at apex,
leaning anteriorly. GASTER: Smooth, lacking sculpturing, tergites 1-3 equal
in size, 4-5 decreasing in size. COLORATION AND PILOSITY:
Concolorous light brown species. Dorsal lateral cephalic margins with short,
appressed pubescence. Antennal scapes with short, suberect hairs, funiculus
with suberect to decumbent hairs. Up to 20 erect to suberect hairs surpassing
anterior clypeal margin. Dorsal mesosomal surface covered in appressed
pubescence, longest hairs on anterior dorsal mesonotum. Lateral mesosomal
surface with sparse pubescence. Posterior petiolar surface lacking hair. Gaster
entirely covered in appressed to suberect pubescence. Pubescence light brown
to yellow.

Distribution

Known only from type locality in the Assam Province of India.

Type Material: Holotype worker: India (MHNG).

Del Toro, I. et al.- Revision of the Ant Genus Liometopum 319

Material Examined

INDIA: Assam: Debrugarh, O. Lindgreeni, Holotype.

Habitat and Biology: Unknown

Species Summary
This is the least collected species of the genus. A suite of
morphological differences in cephalic characters (having a narrower head),
mesosomal characters (rounded dorsal face of propodeum), and petiolar
characters (flat at petiolar apex), indicate that this is a valid species and
should retain its current status. To support our hypothesis more material
needs to be analyzed, however no reports or additional collections of this
species were encountered. We believe that the sample size constraints are not
a major issue in retaining a valid species status for this taxon.

Liometopum luctuosum Wheeler

Figs. 11-16, 40-44, 65-68, 71, 81-86

Liometopum apiculatum subsp. luctuosum Wheeler 1905: 325, worker,

USA: Colorado Springs, Colorado; Forel 1914: 619, male; USA: California,
Lake Tahoe; Liometopum occidentale subsp. luctuosum [2 Examined]:
Creighton 1950: 339; Liometopum luctuosum: Wheeler & Wheeler 1986: 55,
larva, USA: Nevada.

Diagnosis

Worker: Liometopum luctuosum is a concolorous dark brown species

with a glossy cuticle covered in dense, appressed pubescence. The scapes do
not surpass the posterior margin of the head by two times the maximum
thickness of the scape. The metanotal suture is slightly depressed and breaks
the continuity of the posterior mesosomal margin. Liometopum luctuosum is
less commonly encountered than L. apiculatum or L. occidentale. This
species is usually collected at elevations higher than 1600 m.

Gyne: Queens are four times larger than the workers, slightly larger
than the males and are concolorous light to dark brown. The dorsum of the
320 Sociobiology Vol. 52, No. 2A, 2009

mesosoma is covered with short erect hairs of equal length (0.18 mm). The
metanotal suture is clearly defined and strongly depressed, breaking the even
dorsal mesosomal margin.

Male: Males are concolorous black and smaller than the gynes. The
hind wing has 14-18 hamuli. The volsella lacks pilosity along the ventral
margin and has a straight posterior margin. The posterior ventral margin of
the aedeagus has more than 20 clearly defined triangular denticles with the
posterior vertex of the aedeagus coming to a triangular apex. Males of this
species are usually collected at elevations higher than 1900 m and range from
Pacific West Coast east to West Texas and south to Central Mexico.

Comments: Workers of this small concolorous dark brown species are

easily separated from L. occidentale workers, which are bicolored.
Liometopum luctuosum workers can be separated from those of L. apiculatum
by having a smaller total length, a metanotal suture that breaks the straight
dorsal outline, and a shorter antennal scape (~0.88 mm). Gynes of
Liometopum luctuosum can be separated from those of L. apiculatum and L.
occidentale by having a smaller total size and by having a lower pilosity
density than those of the other Nearctic species. Liometopum luctuosum males
separate from those of L. occidentale and L. microcephalum by having more
than 20 clearly defined triangular denticles along the posterior ventral margin
of the aedeagus and having a triangular posterior aedeagal apex.

Caste Descriptions
Workers (n=10)
TL 3.31-4.31 (3.68), HL 0.75-1.05 (0.95), HW 0.78-1.14 (1.06), EL
0.19-0.23 (0.20), EW 0.11-0.15 (0.13), SL 0.88-0.93 (0.89), PH 0.25-0.37
(0.29), PL 0.13-0.25 (0.16), CI 105-121 (111), SI 90-107 (87), PI 50-75 (54)
HEAD: Wider than long, posterior medial concavity well defined.
Majors without ocelli. Eyes medial rather than lateral, 90 to 100 ommatidia.
Scape surpasses posterior margin of head by less than two times maximum
width of scape. Anterior clypeal margin straight. Mandibles finely punctate, 8
teeth along masticatory margin, 3-4 teeth along basal margin. Apical teeth
largest, teeth decreasing in size along masticatory margin. MESOSOMA:
Mesosoma convex, metanotal suture breaking dorsal outline. In dorsal view
widest at middle of pronotum, as wide as head. Metanotal suture well defined,
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 321

depressed. Legs long, slender, femur as long as tibia. Propodeal dorsal margin
rounded into posterior face, propodeal spiracle small and round. Petiole small,
scale-like with sharp apex, leaning anteriorly, higher than long (0.29 mm
high, 0.16 mm long). GASTER: First gastral tergite largest, tergites
decreasing in size posteriorly. In dorsal view gaster rounded at pygidium.
COLORATION AND PILOSITY: Concolorous light golden brown to dark
brown. Cuticle covered in short, appressed pubescence. Head covered in
appressed pubescence, with short, erect hairs on antennal scapes and posterior
cephalic margin. Fewer than 10 short, erect hairs surpassing anterior clypeal
margin. Longest erect pilosity on pronotal dorsum (fewer than 15 erect hairs).
Pilosity on petiolar lateral surfaces, few hairs on anterior petiolar surface.
Pilosity dull gray to silver.

Gynes (n=10)
TL 9.50-10.63 (10.16), HL 1.48-1.65 (1.57), HW 1.75-2.06 (1.93),
EL 0.35- 0.40 (0.38), EW 0.25-0.35 (0.29), SL 1.13-1.38 (1.22), PH 0.63-0.79
(0.73), PL 0.29-0.44 (0.34), WL 11.00-12.25 (11.61), CI 112-131 (120), SI
76- 86 (79), PI 38-58 (50)
HEAD: Wider than long, posterior cephalic margin weakly concave.
Lateral ocelli longer than wide, medial ocellus circular. Eyes medial rather
than lateral, oval-shaped with 140-150 ommatidia. Scapes never surpassing
posterior margin of head. Anterior clypeal margin concave. Mandibles
coarsely punctate, with suberect hairs originating at punctures, 7-8 teeth along
masticatory margin, 3 smaller teeth along basal margin, apical and subapical
teeth largest. MESOSOMA: Dorsum convex. From dorsal view widest at
pronotum, lateral dorsal corner of pronotum touching axillary sclerite.
Metanotal suture clearly defined. Legs long and slender, femur and tibia
subequal in length. Propodeum arched, ventrally emarginate, spiracle round.
Petiole scale-like, bidentate at apex, petiolar apex rounded rather than sharp,
when seen in posterior view, large lateral spiracle on lower lateral half.
GASTER: Gaster large, first gastral tergite wider than second through fifth,
continuously decreasing in size. COLORATION AND PILOSITY:
Concolorous light brown. Short, appressed pubescence on all cuticular
surfaces. Dorsal and lateral cephalic margins with decumbent hairs, 15 to 20
suberect hairs surpassing anterior clypeal margin. Antennae covered with
short suberect hairs. Mesosoma covered in short appressed hairs, short erect
hairs of equal length along dorsal surface. Suberect hairs on anterior and
322 Sociobiology Vol. 52, No. 2A, 2009

lateral petiolar surfaces. Gaster mostly with appressed pubescence, short erect
hairs along ventral and dorsal surfaces. Pilosity golden yellow.
Males (n=8)
TL 7.13-10.50 (8.86), HL 1.00-1.10 (1.03), HW 1.10-1.25 (1.22), EL
0.38-0.44 (0.41), EW 0.31-0.38 (0.34), SL 0.48- 0.56 (0.51), PH 0.80- 0.85
(0.83), PL 0.29-0.38 (0.33), WL 9.21-11.25 (10.05), CI 125-127 (126), SI 51-
53 (52), PI 50-51 (50)
HEAD: Wider than long. Ocelli present, lateral ocelli longer than
wide, medial ocellus round. Eyes lateral, extending past lateral cephalic
margin, with more than 140 ommatidia. Scapes never surpassing posterior
cephalic margin. All funicular segments longer than wide. Anterior clypeal
margin straight (sometimes weakly convex). Mandibles finely punctate, 10
teeth along masticatory margin, 3-4 teeth along basal margin. MESOSOMA:
Strongly convex. Anterior pronotal margin straight, vertical. Mesonotum
convex. Widest at anterior edge of scutellum. Legs long and slender, femur
1/4 longer than tibia. Dorsal posterior surface of propodeum rounded.
Propodeal spiracle round and very small. Petiole scale-like, bidentate and
upright, apex sharp. GASTER: First gastral tergite largest, others
continuously decreasing in size. Genital capsule smooth and glossy.
Parameres large, strongly concave dorsally, ventrally weakly concave.
Volsella large and lobose, posterior margin straight. Aedeagus large, with 20-
25 triangular denticles along posterior ventral margin. Posterior aedeagal
apex triangular. Subgenital plate with rounded notch along posterior margin.
COLORATION AND PILOSITY: Concolorous light to dark brown.
Appressed pubescence on most cuticular surfaces. Head with long, suberect
pubescence. Mandibles with decumbent pubescence. Lateral surfaces of
mesosoma lacking appressed pubescence, resulting in glossy appearance,
dorsal surface covered in dense, appressed pubescence. Gaster covered with
appressed pubescence. Genital capsule without pilosity. Parameres with long,
suberect hairs on ventral surface. Pilosity golden yellow.

Distribution
Northwestern United States (Washington State), south and west to
Western Texas, south to Central Mexico.
Type Material: 2 cotype workers examined: USA (MCZC); 2 syntype
males examined: USA (MHNG).
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 323

Material Examined
MEXICO- Coahuila: Arteaga, El Paraíso, 30.x.2005, 25°23’28.0” N
100°42’49.8” W, J.L. Navarrete; 53 km E. Arteaga, 24.ix.1987, W.P.
Mackay. Guanajuato: Parque Nacional Jose Morelos, 22.v.1988, W.P.
Mackay. Michoacán: Aguililla, Paracho, 19°36’05.4” N 102°05’06.1” W, F.
Núñez, L.A. Navarrete, J. Cortes, M. Vásquez. Nuevo Leon: San Pedro
Garza Garcia, Parque Chipinque, Camino al Piñar, 11.x.2005, M. Vásquez.
Zacatecas: Teul de González, Ortega camino a Millipillas, 1-6.ii.2004,
21°19’ 43.8” N, 103°35’21.0” W, M. Vásquez. USA- Arizona: Chiricahua
Mts., 11.7 Km. W Portal, 31’56’ N 109’15.8”W, 5-14.viii.1976; Stoneman
Lake, 28.viii.1933, R. Lingenberry; San Francisco Mts., 0.6 miles SW Jct. Rt.
89 on FSR 420, 8.vii.1992, S.P. Cover; 48 miles NW Nixon Spring,
5.viii.1969, R. Snelling; Chiricahua Mts., Herb Martyr Dam, 6.viii.1963;
5.xii.1998, N. Cobb; Grand Canyon, 20.vii.1931, A.C. Cole; Nixon Spring,
5.vii.1969, R. Snelling; Overguard, 30.vii.1950, R.G. Robinson; Granite
Dells, 4 mi W. Prescott, 15.vii.1971, L. Martin; Sunny Flat Campground,
31°88’ N 109° 17’ W, 22-24.viii.1993, B.V. Brown. California: San Gabriel
Mts., Charlton Flat, 19.vii.1963; Blue Ridge, San Gabriel Mts.; Figueroa Mt.
Campground; Mirror Lake, Yosemite Park 23.v.1930; Carrol Cr., 6 mi N
Bishop, 20.viii.1960, E. Schlinger; San Gabriel Mts., 13.vii.1966, R. Snelling;
3 mi W Cachuma Lake, 06.vii.1959; Mill Portero, Arroyo Seco, Santa Lucia
Mts., 16.v.1978, J.P. Donahue; Eagle Lake, 11.vii.1970, H. Jacobson; Sierra
N.F., 4.4 mi. SW Big Creek, 16.v.1976, A. Newton; Caruthers Canyon,
35’15’N 115’18’W, 22.iii.1996, P.S. Ward; Cold Canyon, 15.iii.1983, J.
Longino; Sta. Rosa Mountains, 4.ii.1984, J. Longino. Colorado: Decker,
29.vii.1941, Buren; Cheyenne Canyon, Near Colorado Springs, 20.vii.1903,
MCZ Cotype, W.M. Wheeler; Colorado Springs, 18.vii.1903. New Mexico:
Jemez Springs, 25.v.1988, R. S. Peigler. Nevada: Sparks, 20.x.1979; Geiger
Grande, 6 mi. SW Reno, C&J Wheeler; Elk Point, La Rivers, Hiko,
7.viii.1974; White Canyon on Mt. Rose, 19.vi.1977; Verde Wash, 9.vi.1951,
W.M. Wheeler; Reno, 12.vii.1949, R.C. Bretcher; Sparks, 7.viii.1974, R.C.
Bechtel; Elk Point, 9.v.1970, G.&J. Wheeler; Charleston Park, Lake Tahoe,
29.vi.1954, A.C. Cole; 12.3 Km. SSE Miden; Reno, 12.vii.1978; Charleston
Park, 9.v.1970. Texas: McCutchens Ranch, 7.v.1912, J.D. Mitchel. Utah:
Zion National Park, 22.vii.1932, W.S. Creighton; Hach Canyon, 21.vi.1935.
Washington: Chelan, 14.v.1983, J. Longino; Eightmile Campground, near
Leavenworth, 23.v.2007, J. Longino, 3 km NE Lyle; Klickitat 18.iv.2003, J.
Longino.
324 Sociobiology Vol. 52, No. 2A, 2009

Habitat
This species has a strong association with pine trees. However, it has
also been reported in sagebrush, oak forests, ponderosa pine-riparian,
Douglas fir, and riparian habitats at high elevations. Liometopum luctuosum
has recently been reported from Idaho nesting in rock crevices in juniper
woodlands (Clark & Bloom, 2007). It usually occurs at elevations higher than
2000 m., however in coastal environments, this species may be collected at
lower elevations.

Biology
This species is less common and a morphologically smaller species
than L. apiculatum. Nests are commonly located under rocks, under decaying
logs or at the base of large trees. This species also constructs carton nests
much like L. apiculatum, which are often located in very inaccessible places.
Liometopum luctuosum has been reported as being a competitor with
Camponotus species in Idaho, since both of these genera compete for similar
nesting sites (Merickel & Clark, 1994). The staphylinid genus Liometoxenus
was first described from specimens found foraging next to colonies of L.
luctuosum and L. occidentale (Kistner et al., 2002). Reproductive castes are
usually observed during the months of June and July and can easily be
collected the day after the flight by looking in large bodies of water or using a
backlight trap.
This species has been collected in or around rural housing areas and
can be considered a minor pest. No severe damages to housing or property
have been reported, but the uncomfortable bite and smell of this ant makes it
a nuisance to affected residents. Additionally, L. luctuosum is of particular
interest in the Idaho National Laboratories, where it can potentially uncover
buried waste materials (Clark & Blom, 2007).
This species tends to be very easily disturbed and aggravated. When
this occurs a strong odor is released from the anal glands and the ants begin to
attack and bite any intruder, making this species just as aggressive as L.
apiculatum.

Species Summary
We recognize this species as valid. Syntypes of the males to this
species were examined, illustrated, described and photographed. Multiple
works (Mackay and Mackay, 2002; Shattuck, 1992; Cook, 1953) make
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 325

reference to this species and provide additional morphological characters,

which were helpful in identifying L. luctuosum. Gynes of this species were
not described in detail prior to the work presented here. There are multiple
morphological differences identified in the gyne caste (pilosity, total size,
suture definition, wing venation), which aid in the identification of L.
luctuosum gynes. This species has been considered to be a subspecies of both
L. apiculatum and L. occidentale. However, the lack of intermediates and
species clines suggests that this species is reproductively isolated. Even
though, the ranges of L. luctuosum apparently do overlap with those of L.
apiculatum and L. occidentale, collecting experience suggests that this
species typically occurs at elevations above 1600 m while L. apiculatum and
L. occidentale are normally lower elevation species typically inhabiting
regions with a maximum elevation of 1500 m. There is no evidence of
hybridization in zones of sympatry.

Liometopum microcephalum Panzer

Figs. 17-22, 31, 45-49, 72, 93-98

Formica microcephala Panzer 1798: 1652, male, GERMANY; Liometopum

microcephalum: Mayr 1861: 39, worker and queen, AUSTRIA.

Formica austriaca Mayr 1853: 144, worker, AUSTRIA (Junior Synonym of

microcephalum Mayr 1861: 39)

Diagnosis

Worker: Liometopum microcephalum is bicolored, with a dark brown

head, gaster, and legs, the mesosoma is lighter colored. The cuticle is covered
in dense pubescence, interspersed with erect and suberect hairs. The longest
erect hairs are on the pronotum (0.40 mm long). The scapes do not surpass
the posterior margin of the head.

Gyne: Queens are slightly larger than males and similar in

morphology to queens of L. occidentale. The mandibular apical tooth is
distinctly larger than the remaining masticatory teeth. The medial cephalic
concavity is clearly defined and not as straight as in L. occidentale.
326 Sociobiology Vol. 52, No. 2A, 2009

Male: Males are concolorous dark brown to black. The anterior

clypeal margin is concave medially; it is straight in males of other species that
were seen. The dorsal mesosomal margin is convex and notably depressed at
the metanotal suture. The parameres are large with the ventral margin
concave and dorsal margin convex. The posterior lobe of the volsella has a
straight posterior margin. The aedeagus has 15-18 small triangular denticles
along the ventral margin with a triangular notch posterior to the denticles.

Comments: Liometopum microcephalum can be separated from other

Old World species based on the consistency of the bicoloration, not seen in L.
lindgreeni or L. sinense. Having a moderate concavity on the anterior clypeal
margin can also be used to separate this species from L. lindgreeni, which has
a straight anterior clypeal margin. Liometopum sinense only occurs in China,
and L. microcephalum has a more western distribution. This species is
morphologically similar to L. occidentale, but geographic isolation and
morphometric statistical analysis of the two taxa suggest that the two species
are different (Fig. 107). Males of L. microcephalum can be separated from
those of L. orientale by having a paramere with a concave ventral margin,
which is straight in L. orientale. Males of this species also separate from
males found in the New World by having fewer denticles along the posterior
ventral aedeagal margin than L. apiculatum and L. luctuosum. Additionally
the aedeagal ventral margin is notched whereas in L. orientale the aedeagal
ventral margin is concave but not notched.
Caste Descriptions
Workers (n=10)
TL 4.19-6.00 (5.13), HL 1.28-1.41 (1.35), HW 1.54-1.63 (1.55), EL
0.27-0.31 (0.28), EW 0.18-0.21 (0.19), SL 1.10-1.25 (1.16), PH 0.46-0.58
(0.53), PL 0.20-0.31 (0.26), CI 110-120 (115), SI 78-92 (86), PI 42-54 (48)
HEAD: Wider than long, posterior medial concavity well defined. Majors
with ocelli. Eyes large, medial rather than lateral, with 115-120
ommatidia. Scape slender (diameter =0.22 mm), does not surpass posterior
cephalic margin by more than two times maximum thickness. Funiculus
11 segmented, segments longer than wide, first and last segment longest.
Clypeal anterior margin medially concave. Mandibles punctate, 8-10 teeth
along masticatory margin, 2-3 smaller teeth along basal margin.
Apicalmost teeth largest. MESOSOMA: Strongly convex (similar to
Camponotus spp.). Pronotum convex, widest at dorsal medial point of
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 327

pronotum, as wide as maximum head width. Mesonotum arched,

metanotal suture distinct, but not depressed. Legs long and slender, femur
and tibia of equal length. Propodeal dorsal margin round, vertical at
posterior face, spiracle, round. Petiole scale-like, with sharp angular apex
in dorsal view, rounded at apex, inclined anteriorly. GASTER: First
gastral tergite largest, others decreasing continuously in size. Rounded at
pygidium. COLORATION AND PILOSITY: Bicolored, head, gaster and
legs darker brown than golden yellow mesosoma. Dense appressed
pubescence covering entire cuticular surface. Lateral and posterior
cephalic margins with appressed pilosity, few short decumbent hairs along
posterior cephalic margin. No more than 20 short erect hairs surpassing
anterior clypeal margin. Dorsal mesosomal surface covered in dense
appressed pubescence. Longest hairs on posterior dorsal edge of pronotum
(0.25 mm). Lateral surfaces of mesosoma with fewer erect and suberect
hairs. Short erect to appressed pilosity on anterior surface of petiole.
Dense appressed pubescence covering entire gastral surface. Pilosity
golden yellow.
Gynes (n=2)
TL 11.5-12.0, HL 1.81-1.88, HW 2.23-2.37, EL 0.44, EW 0.38-0.40,
SL 1.35-1.50, PH 0.79-1.00, PL 0.25-0.35, CI 124-126, SI 75-80, PI 32-35,
WL 13.00
HEAD: Wider than long, posterior cephalic margin concave. Ocelli
present, lateral ocelli longer than wide, medial ocellus circular. Eyes medial
rather than lateral, large, oval-shaped, with more than 150 ommatidia. Scapes
not surpassing posterior cephalic margin, funicular segments longer than
wide, first and second segments, longest. Anterior clypeal margin medially
concave. Mandibles punctate, 8-9 teeth along masticatory margin, 2-3
reduced teeth along basal margin. MESOSOMA: Strongly convex, weakly
depressed at metanotal suture. Dorsal pronotal surface vertical, finely
punctate. Anterior mesonotum convex, rounded posteriorly. Metanotal suture
weakly depressed, touching axillary sclerite. Wing discodial and submarginal
cells closed. Second submarginal cell triangular, first discodial cell
rectangular-shaped. Legs long, and slender, tibia 2/3 length of femur. Dorsal
propodeal margin rounded, posterior surface vertical. Propodeal spiracle
lateral, medially located, small and round. Petiolar scale with sharp apex,
from posterior view, bidentate at apex. GASTER: Gaster large half of total
length, first tergum widest, tergites 2-5 decreasing in width. COLORATION
328 Sociobiology Vol. 52, No. 2A, 2009

AND PILOSITY: Concolorous brown. Dense pilosity covering most cuticular

surfaces. Erect to sub-decumbent pilosity along posterior and lateral cephalic
margins. Antennae covered with short, appressed pubescence. 15-20 hairs
surpassing anterior clypeal margin. Mesosomal dorsal surface covered in
appressed to erect hairs, longest hairs on propodeum 0.20 mm. Dorsal face of
propodeum with 30-40 short erect hairs of equal length. Suberect pilosity on
anterior petiolar surface. Appressed pubescence covering entire gastral
surface, erect and suberect hairs on dorsal and ventral surfaces. Pubescence
golden yellow to silver.

Males (n=1)

TL 9.13, HL 1.10, HW 1.65, EL 0.40, EW 0.31, SL 0.50, PH 0.69, PL 0.31,

CI 150.0, SI 45.4, PI 45.4, WL 11.00
HEAD: Wider than long, posterior cephalic margin straight, lacking
concavity. Ocelli present, medial ocellus wider than long, lateral ocelli longer
than wide. Eyes lateral rather than medial, extending past lateral cephalic
margins, large, with more than 145 ommatidia. Antennal scapes reduced,
never surpassing posterior cephalic margin. Funiculus 12 segmented,
segments longer than wide. Anterior clypeal margin convex medially.
Mandibles with 8-9 teeth on masticatory margin, no teeth on basal margin,
two apical teeth largest. MESOSOMA: Strongly convex. Dorsal margin of
mesonotum anteriorly vertical, medially convex. Widest at posterior
mesonotum from dorsal view. Metanotal suture well defined, depressed.
Discoidal and submarginal wing cells closed. Second submarginal cell
triangular, first discoidal cell rectangular. Posterior wing with 14 hamuli.
Legs long, slender, tibia 3/4 length of femur. Propodeum arched, dorsal
margin rounded. Propodeal spiracle, round. Petiole scale-like, bidentate at
petiolar apex, vertical. GASTER: First gastral segment largest, overhanging
petiole. Genital capsule large, parameres large, ventral margin concave,
dorsal margins convex. Volsella large, posterior margin straight. Aedeagus
with 15-18 denticles covering 1/2 of ventral margin, posterior apex rounded
and overhanging. Subgenital plate medially notched in U-shape along
posterior margin. COLORATION AND PILOSITY: Concolorous dark
brown. Dense appressed pubescence covering all body surfaces. Erect to
decumbent hairs along posterior and lateral cephalic margins. 22 hairs
surpassing anterior clypeal margin. Dorsal and ventral mesosomal surfaces
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 329

with appressed pilosity, lateral surfaces with short, appressed pubescence.

Gaster covered in appressed pubescence, genital capsule lacking pilosity.
Suberect to decumbent pilosity on parameres. Pilosity golden yellow.

Distribution

Southern Europe, Italy east to Asia Minor, south to Israel.

Type Material: No type material was examined.

Material Examined
ALBANIA- Corovade: Karkrake, 40°32’17”N 20°10’32”E. AUSTRIA-
Neiderosterrrieich: 30 km W. Veienna. Laxenburg: Scholsspark, 16.35° E,
48.07° N, 13. ix. 2002, B.C. Schlicksteiner. BOSNIA- 25km. W. Banja Luka,
44°42’08” N 16°52’30”; 32 km. S. Mostar 43°03’06” N 17°51’44” E.
BULGARIA- Oblast Lovech: Lukowit, 01.33 43°11’ N 24°15” E. Rila: Pilo
Selo, Forel Collection. CROATIA- Brgat: 42°39’15”N 18°08’37” E.
CZECHOSLOVAKIA- Moravia: mer. Lednice, 30.v.1982, P. Bezdeck, 1gt.
P. Werner. GREECE- Lefkada: Karva, 38°45’ N 20°40’ E. HUNGARY-
Szekesfeharvar: 47°09’N 18°20’ E, 25 km, 1981, A. Hefetz. ITALY-
Portici: A. Forel. LEBANON- Christilansen: Hasbani River Source,
27.ii.1953. ROMANIA- Juderul: Siba. SERBIA- Frusva Gora Mt.,
14.v.1989, leg. I. Petrov; Kosovo, Doljani 42°27’42”N 19°18’30”E.
Vojvodina: Vrdnik, Lediniko Jezero. UKRAINE- Piatyhatky.

Habitat
This is a wide ranging species, collected throughout southern and
Eastern Europe (Atanassov & Dulusski, 1992; Baroni Urbani, 1971; Brako,
2003; Kutter, 1977; Weist, 1967). This species builds carton nests in the
crevices of trees, particularly oaks. Habitats may range from coastal
environments, to floodplain and oak forests.

Biology
A great deal is known concerning the biology of this species. In
recent years the potential for this species to be used as a method of biological
control of Cameraria ohridella has been investigated (Grabenerger et al.,
2005). It is believed that populations of this species may be endangered due
to habitat loss (Schlick-Steiner et al., 2003). Colonies are extremely large and
330 Sociobiology Vol. 52, No. 2A, 2009

may be composed of several thousand individuals (Zettel et al., 2004).

Foraging trails are large and cover up to 80 m in length. This species has been
observed tending homopterous insects; however predaceous behavior has
been documented in larger colonies (Brako, 2003). This species is extremely
aggressive and will defend the colony by standing on its hind legs and biting
the intruder (Martinez & Tinaut, 2001). Natural predators that specialize in
preying on L. microcephalum include Tracheliodes curvitarrsus and T. varus
(Hymenoptera: Crabronidae) (Zettel et al., 2004).

Species Summary
Liometopum microcephalum is extremely similar to L. occidentale
morphologically. However, we retain its current status as a valid species due
to its separation based on morphological differences in the reproductive
castes, unique morphometric characteristics (Fig. 107) and geographic
isolation. Workers of this species were analyzed using the descriptions
provided by Mayr 1861. For a more detailed account of the morphology of L.
microcephalum we used the descriptions and drawings provided by
Kupyanskaya (1988). Liometopum orientale was considered a subspecies of
L. microcephalum (Karavaiev, 1927). Kupyanskaya (1988) later raised L.
orientale to valid species status based on a suite of morphological characters,
most importantly significant differences on exterior male genitalia
morphology. No new synonymies are presented for this species.

Liometopum occidentale Emery

Figs. 23-28, 50-54, 73, 87-92

Liometopum microcephalum var. occidentale Emery 1895: 330, worker and

male, USA:
California, San Jacinto; Liometopum apiculatum var. occidentale:
Wheeler 1905:
324, USA: California, San Jacinto; Liometopum occidentale Wheeler
1917: 521, queen, USA: California, San Jacinto

Diagnosis
Worker: This is a common bicolored species, having a dark brown
head and gaster with a lighter colored mesosoma, usually golden yellow, but
some may be light brown. The antennal scapes are short (1.02 mm) and
surpass the posterior margin of the head by less than twice the maximum
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 331

thickness of the scape. The scape has few short, decumbent hairs (no more
than 20). The longest erect hairs are on the dorsal surface of the pronotum
(0.26 mm in length). The mesosoma is convex with the highest point at the
mesonotum.

Gyne: Queens of this species are concolorous dark brown to dull

yellow. Liometopum occidentale queens are slightly larger (TL ~11 mm) than
queens of L. luctuosum but not as large as the queens of L. apiculatum. The
metanotal suture is distinct, but weakly depressed and does not break the
continuity of the mesosomal margin.

Male: Males of this species are concolorous dark brown to black. The
first discodial cell of the forewing is rectangular, the hind wings have only
15-17 hamuli. The posterior lobe of the volsella has a strong concavity along
the dorsal margin. The aedeagal ventral margin has 15-18 small triangular
denticles covering 1/2 of the ventral aedeagal margin, with an even concavity
posterior to the denticles.

Comments: The worker of Liometopum occidentale can be separated

from other species by cuticle coloration, which is bicolored in this species and
concolorous in L. apiculatum and L. luctuosum. Additionally, it differs from
L. apiculatum by having antennal scapes that do not surpass the posterior
margin of head by twice the maximum thickness of the scape. Also it differs
from L. luctuosum by having a continuous mesosomal outline whereas in L.
luctuosum the mesosoma is depressed at the metanotal suture. Gynes are
similar in total size as the gynes of L. luctuosum, but are consistently smaller
than the gynes of L. apiculatum. Males of this species separate from L.
apiculatum males by being smaller in total length, and having fewer hamuli
on the anterior margin of the posterior wing. Liometopum occidentale males
separate from males of L. luctuosum and L. apiculatum males by having
fewer denticles along the ventral aedeagal margin, which is rounded at the
aedeagal apex (notched in L. microcephalum). Additionally males of this
species have a strong concavity along the posterior margin of the posterior
lobe of the volsella (straight in all other species). This species is generally
collected at elevations below 1800 m, whereas L. luctuosum is normally
collected at elevations above 1900 m.
332 Sociobiology Vol. 52, No. 2A, 2009

Caste Descriptions
Workers (n=10)
TL 2.88- 5.06 (4.44), HL 0.81-1.23 (1.09), HW 0.89-1.38 (1.23), EL
0.20-0.25 (0.24), EW 0.15-0.19 (0.16), SL 0.75-1.15 (1.03), PH 0.35- 0.59
(0.49), PL 0.16-0.31 (0.21), CI 96- 125 (112), SI 85- 105 (93), PI 29- 50 (43)
HEAD: Wider than long, posterior medial concavity well defined.
Majors rarely with ocelli. Eyes large, longer than wide, medial rather than
lateral with 110-125 ommatidia. Antennal scapes short (1.03 mm), surpassing
posterior margin of head by less than two times maximum thickness of scape.
Funicular segments 1, 2, and 11 longest, all segments longer than wide.
Clypeal anterior margin moderately concave. Mandibles punctate, 8-9 teeth
on masticatory margin, 2-4 teeth on basal margin. MESOSOMA: convex,
most arched at promesonotum, from dorsal view, widest at pronotum 2/3 as
wide as maximum head width. Pronotum smooth, convex. Mesonotum
straight, weakly indented at sutures. Metanotal suture not depressed. Legs
long, slender, femur and tibia equal in length. Propodeum concave, spiracle
small, not well defined. Petiole scale-like, sharp petiolar apex, from posterior
view, petiole rounded at apex, leaning slightly anteriorly. GASTER: First
gastral tergite sometimes covering petiole, other tergites smaller,
continuously decreasing in maximum width, rounded at pygidium.
COLORATION AND PILOSITY: Bicolored, darker head and gaster, usually
dark brown, lighter colored mesosoma usually golden yellow to golden
brown. Dense pilosity covering entire cuticular surface. Suberect hairs on
dorsal and lateral cephalic margins. Appressed pubescence on lateral cephalic
margin. Scape with few short, erect hairs, funicular segments with appressed
pubescence. 8-12 erect hairs surpassing anterior clypeal margin. Dorsal
surface of mesosoma covered in appressed pubescence, lateral surfaces with
few or no hairs. Longest erect hairs (0.26 mm long) on dorsal surface of
pronotum. Highest hair density on dorsal surface of gaster. Gaster covered
entirely with appressed pubescence and short suberect hairs. Pilosity golden
yellow.

Gynes (n=7)
TL 10.5-12.0 (11.03), HL 1.63-1.85 (1.77), HW 2.00-2.38 (2.20), EL
0.38-0.44 (0.39), EW 0.31-0.44 (0.39), SL 1.10-1.40 (1.23), PH 0.83-1.10
(0.95), PL 0.35-0.56 (0.42), CI 112-139 (124), SI 61-79 (70), PI 36-46 (43)
HEAD: Wider than long, posterior cephalic margin weakly concave.
Ocelli present, all circular and equal in size. Eyes medial rather than lateral,
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 333

large, oval-shaped, with 150-170 ommatidia. Scape short, not surpassing

posterior cephalic margin. Frontal lobes protruding anteriorly. First and last
funicular segments longest, others continuously decreasing in length. Anterior
clypeal margin slightly concave. Mandibles punctate, 8 teeth along
masticatory margin, 2-3 reduced teeth along basal margin. MESOSOMA:
Strongly convex. Dorsal pronotal surface rounded. Anterior mesonotum
convex, posterior mesonotum straight. Metanotal suture touching axillary
sclerite. Legs long, slender, femur and tibia equal length. Dorsal posterior
propodeal margin rounded. Propodeal spiracle round, medial, not clearly
defined. Petiole scale-like, leaning anteriorly, bidentate at apex when seen
from posterior view. GASTER: First tergite 1/2 total length, widest, tergites
3-5 continuously decreasing in width. COLORATION AND PILOSITY:
Generally concolorous light brown to dark brown, some with lighter colored
mesonotum, propodeum, and legs. Dense pilosity covering most cuticular
surfaces. Erect to decumbent pilosity along lateral and posterior cephalic
margins. Antennae densely covered with short, appressed pubescence. 12 to
18 erect hairs surpassing anterior clypeal margin. Dorsal surface of mesosoma
covered in appressed to erect hairs, longest erect hairs on promesonotum.
Metanotum and propodeum with few or no hairs. Lacking pubescence on
posterior petiolar surface, suberect to decumbent hairs on anterior face of
petiole. Appressed pubescence covering entire gastral surface, short suberect
and erect hairs on dorsal and ventral gastral surfaces. Pilosity golden yellow
to silver.

Males (n=9)
TL 8.50-11.25 (9.58), HL 1.00-2.20 (1.20), HW 1.30-2.85 (1.55), EL
0.39-0.44 (0.41), EW 0.31-0.38 (0.34), SL 0.50-0.63 (0.55), PH 0.76-0.94
(0.85), PL 0.23-0.30 (0.22), CI 123-130 (128), SI 26-57 (47), PI 27-32 (31)
HEAD: Wider than long, posterior cephalic margin straight. Ocelli
present, medial ocellus wider than long, lateral ocelli longer than wide. Eyes
lateral rather than medial, extending past lateral cephalic margins, large with
more than 130 ommatidia. Antennal scapes reduced, never surpassing
posterior margin of head, reaching only lateral ocular margin. Funicular
segments longer than wide, funiculus 4 times longer than scape, 12
segmented. Anterior border of clypeus convex medially. Mandibles with 8-9
teeth on masticatory margin, 3-4 teeth on basal margin, apical teeth largest.
MESOSOMA: Strongly convex. Anterior edge of pronotal margin broadly
convex. Mesonotum arched, widest (dorsal view) at posterior mesonotum.
334 Sociobiology Vol. 52, No. 2A, 2009

Metanotal suture well defined, but not breaking continuous mesosomal

margin. Submarginal and discodial cells closed, first discodial cell
rectangular. Hind wing with 15 or fewer hamuli. Legs long and slender, tibia
3/4 length of femur. Propodeum arched. In lateral view, propodeal spiracle
centered, small, oval-shaped. Petiole scale-like, petiolar sclerite keel-shaped,
petiole bidentate at apex, upright. GASTER: First gastral segment largest,
overhanging petiole. Genital capsule large. Parameres large, ventral margin
straight, dorsal margin weakly concave. Volsella large, posterior lobe with
concave posterior margin. Aedeagus with 15-18 small, triangular denticles
reduced to bumps along posterior ventral margin, posterior apex rounded.
Subgenital plate medially notched in U shape along posterior margin.
COLORATION AND PILOSITY: Concolorous light brown to dark brown.
Dense appressed to erect hairs covering all body surfaces. Suberect to
decumbent hairs along posterior and lateral cephalic margin. 18-20 hairs
surpassing anterior clypeal margin. Short, erect pilosity on antennal scape.
Funiculus covered with dense, appressed pubescence. Dorsal and ventral
mesosomal surfaces with erect hairs, lateral surfaces with scarce pilosity.
Gaster covered in appressed pubescence, appressed pubescence covering
dorsal and ventral surfaces of gaster, lateral surfaces with lower appressed
pubescence density, little or no pilosity on genital capsule. Erect to
decumbent hairs on parameres. Pilosity golden yellow to silver.

Distribution

Northwestern United States (Washington, Oregon, and California)

south to Northwestern Mexico (Baja California).

Type Material: No type material was examined

Material Examined
MEXICO- Baja California: Tecate, Km 105 Carretera Tecate-
Ensenada, Rancho San Ignacio Calamar, 32°30’10.9” N, 116°35’05”W, J.
Cortes Aguilar, V. Flores; Tecate, Km 65 Carretera Tecate-Ensenada,
21.vi.2005, 32°11’01” N, 116°29’34.3”W, J. Navarrete-Heredia, E. Lopez.
USAU- California: 06.v.1962, R. Snelling; China Flat, 15.vii.1948, C.D.
McNeil; Riverton, 28.ii.1958; Chico 4.ix.1958, Schuster; Chico, 5.iii.1967;
Mendocino National Forest, Brittan Ranch, 6 mi WNW of Stonyford,
24.ii.1997, 39°23’ 35” N 122° 39’41” W; Tohouse Spring, Westgard Pass,
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 335

12.iv.1970; Sequoia National Forest, W. Cedar Grove, 14.v.1976; 2 Km SW

Marshal Station, 5.v.1999, 37’N 119’36’W, P.S. Ward; Lakeport, 5.vi.1961;
Santa Monica Mts., 30.xi.1963, C.L. Hougue; Tehachapi Mts., 15.v.1976;
Santa Monica Mts., Stunt Canyon, 1964; HWY 198, 30 miles W of Coalingis,
10.vi.65; Toll Pence, Ribbonwood, San Jacinto Mts., 20.v.1937, J.G.
Shanafelt; Franklin Canyon, 24.iv.1966; Idyllwild, vii.5.1947, W.D. Pierce;
Santa Barbara, 28.iv.1944, W.S. Ross; 15 miles N. Bear Lake, 8.x.1963, G.R.
Ferguson; Arroyo Seco 10.v.1958; 7 mi S Squaw Tank, 20.iii.1965; 0.5 mi N.
Crestline, 7. viii.1967, G.R. Norman; 2. mi E. Cedar Springs; San Gabriel
Mts., 7.3 mi West Mt. Brudy Village, 4.ii.1965; 15 mi SE Big Bend,
20.iv.1923; San Juan Canyon, 11.v.68; DeVol, 15.vi.1963, R. Snelling;
Pinyon Flat, Bear Creek, 8.viii.1967, T. Bunch; Del Puerto Canyon,
15.ix.1956; Coffee Camp, 7.v.1970; Pine Mt. Top NE Hot Springs,
30.v.1984; Preston, 20.vii.1917; Sta. Lucia Mts., 5.vii.1984, Bryant; 12.7
Miles NNW Happy Camp, vi.1969; Indian Flats, 12.vi.1989; Topanga Co., D.
Sheppard; Santa Ynez, 31.x.1962; 5 mi. N. Descano, iv.1972, J.H. Hunt; 12
km ENE Pt. Conception 34°30’ N 120° 21’W, 26.iv.1986, J. Longino; Pauma
Valley, iv.11.1955; Bayou Rd.; 5.2 miles E. Isabel Cr. 10.viii.1965; Mix
Canyon, 11.xii.1960; Del Puerto Canyon7.v.1970, R. Snelling; Iron Mts., 24
miles E Placerville, 1.vii.1980, R. Mason, H. Paul; Matilija nr. Ojai, 1963 R.
Snelling; Borrego Springs, 11.vi.1965; Santa Rosa Plateu Reserve, 3.5 Air
Miles WSW Murrieta, 7.8 iv.1985, Donahue; Sespe Canyon, 25.iii.1967, J.A.
Honey; 4.5 Miles WNW Jct. Rt. 78, Grapevine Canyon rd., 2.vi.1997, S.P.
Cover; Yosemite Valley, 15.vii.1915, W.M. Wheeler; Happy Isles, 30.v.1930,
W.M. Wheeler; Stanford University, 1930, Watson; C. Ishida, Camp Ozona;
Upper Cuyana, 3 miles N. Rumsey McConnell 14.iii.1983, J. Longino;
Arroyo Burro, 19.vii.1987, J. Longino; San Rafael Mountain, 02.ix,1986, J.
Longino; Inyo Co, Big Pine, 10.viii.1953; Los Angeles Co., San Gabriel Mts.,
14.vii.1965; Riverside Co, San Bernardino Mts. Big Pine Flat. Oregon:
Takilma, 31.vii.1999, 42’03’N 123’37’W, P.S. Ward; Champoeg State Park,
22.vi.2002, J. Longino. Washington: 1 km up road on east bank White
Salmon River, 13.v.2001, J. Longino; 10 km E. White Salmon Burdoin
Mountain Area, 19.x.200, J. Longino.

Habitat
This species is commonly collected along coastal regions of southern
Oregon, California, south to Northern Mexico. Nesting sites range from
lowlands up to 1600 m. More recently this species has been reported in urban
336 Sociobiology Vol. 52, No. 2A, 2009

habitats, nesting in homes, underneath insulation or between walls

(Gulmahamad, 1995).
Biology
This species is often found nesting in soil and in crevices of trees and
underneath bark of dead trees. These ants have been observed on oaks, alders,
elms, cottonwoods, creosote, and under rotten logs (Cook, 1953).
These ants have large colonies (up to 60,000 workers) and may be
considered pests as they often forage in housing and may cause minor
damage to facilities. Foraging trails may reach up to 60 m in length and cover
areas of up to 740 m2. Liometopum occidentale has been observed tending
Homoptera, but is also predaceous, and forms massive foraging trails
(Gulmahamad, 1995; Ramoselorduy & Levieux, 1992). When they are
disturbed they emit pungent disturbing odors and resort to biting for defense.
Dinardilla and Sceptobius beetles are often observed alongside foragers or in
nests (Danoffburg, 1994). Flights have observed throughout May.

Species Summary
This species is difficult to evaluate, due to the morphological
similarities with L. microcephalum. Workers, gynes and males of the two
species all appear to have a very similar morphological structure. However
we are hesitant to synonymize it with L. microcephalum for three principal
reasons, one due to our limited sample size of L. microcephalum specimens,
secondly due to the differences in distributional ranges, L. occidentale only
occurring along the western coast of the United States south to the
Northwestern coast of Mexico and L. microcephalum having a wide
distribution in the European continent, Asia Minor and South into Israel. The
different distributions suggest two geographically isolated species. Finally,
discriminant function analysis and canonical correlation analysis separate
both of these species by a suite of morphometric differences (Fig. 106). Male
genitalia dissections were also completed for both species and notable
differences were found (Fig. 47-54). Unfortunately only one male of L.
microcephalum was available for dissection. Descriptions in Wheeler (1905 -
workers), (1917 - gyne) were used as the most reliable to identify this species.
The males described here were collected in association with workers;
therefore we are confident that the identification is correct. Genotypic
analyses should be conducted to better determine the relationship between
these two species. With the available information we believe they should be
recognized as two separate species.
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 337

Liometopum orientale Karavaiev

Figs. 29-30

Liometopum microcephalum var. orientalis Karavaiev 1927: 339, worker,

RUSSIA; Liometopum orientale: Kupyanskaya 1988: 29, raised to
species, male and female, RUSSIA.

Diagnosis

Comments: Though similar in morphology to L. microcephalum, this species

is different in that L. orientale lacks decumbent pilosity along the posterior
margin of the head, and the scapes of this species present two long, erect hairs
at the junction of the scape and the funiculus. The first funicular segment is
twice as long as wide in L. microcephalum; the first funicular segment is
three times longer than wide in L. orientale. Perhaps the most obvious
difference between the two species is petiolar shape, in L. microcephalum the
petiole (when seen from posterior view) is spade-like and comes to a sharp
apex, however in L. orientale, the petiole retains some of the spade-like
features, but does not come to a sharp apex (rounded at the apex). In the
males of this species, the parameres are triangular-shaped, the ventral margin
of the paramere is straight, and lacking the concavity seen in L.
microcephalum. Due to this suite of morphological character differences, we
consider this to be a valid species and retain its current status. For more
details on this species refer to Kupyanskaya (1988).

Type Material: No type material was examined

Material Examined: None

Species Summary
Even though type material was not seen, Kupyanskaya’s assessment
(1988) appears to be thorough and reliable. Cephalic and petiolar characters
as well as male genital capsule morphology are the primary reasons for the
recognition of this as a separate species from L. microcephalum. For future
reference, more samples of this species and of L. microcephalum should be
analyzed to solidify current taxonomic assessments.
338 Sociobiology Vol. 52, No. 2A, 2009

Liometopum sinense Wheeler

Figs. 9-10, 74, 101-102

Liometopum sinense Wheeler 1921: 541, worker, CHINA: Mokanshan

[examined];

Liometopum sinense var. sericatum Wheeler 1921: 542, worker, CHINA:

Mokanshan; New Synonymy [examined]

Liometopum dentimandibulum Chang & He 2002:110, worker, CHINA: New

Synonymy

Diagnosis

Worker: Liometopum sinense is concolorous dark brown to copper in

coloration, covered in dense appressed pubescence on all body surfaces, the
pubescence is finer on the mesosoma than on the gaster. The antennal scapes
reach, but do not surpass the posterior margin of the head, and are covered in
appressed pubescence. The anterior clypeal margin is weakly concave, the
mandibles are punctate, and the apical tooth is the largest. The majors have a
medial ocellus and 8-10 teeth along masticatory margin with 3-4 teeth on the
basal margin of the mandible. The pronotum has 8-12 long erect hairs on the
dorsal surface. Fine appressed pubescence covers the entire mesosomal
surface. The mesosoma is moderately convex, and the highest point is at the
mesonotum. The petiole is scale-like with a sharp apex.

Gyne and Male: Unknown

Comments: This species can be separated from the other Old World species
by the geographic distribution, as it has only been collected in central China.
Morphologically, the petiole differs from those of L. lindgreeni and L.
orientale in that it is scale-like and has a sharp apex; the other two species
have a rounded apex. It differs from L. microcephalum in that L. sinense has
fine appressed pubescence on the mesosoma and the L. microcephalum
mesosomal pubescence is coarse. Additionally, this species is concolorous
while L. microcephalum is bicolored.
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 339

Caste Description
Worker (n=20)
TL 3.25-5.75 (4.71), HL 0.88-1.36 (1.20), HW 1.00-1.53 (1.33), EL
0.21-0.29 (0.25), EW 0.14-0.21 (0.18), SL 0.90-1.38 (1.07), PH 0.38- 0.51
(0.46), PH 0.21-0.38 (0.29), CI 86- 125 (110), SI 80-104 (89), PI 42-73 (63)
HEAD: Wider than long, posterior medial concavity well defined.
Majors with one poorly defined medial ocellus. Eyes medial rather than
lateral, oval-shaped, 110-115 ommatidia. Scape slender (diameter ~0.14 mm),
does not surpass posterior cephalic margin by more than twice maximum
thickness of scape. First and last funicular segments longest (0.24 mm) all
segments longer than wide. Medial anterior clypeal margin moderately
concave. Mandibles punctate, 8-10 teeth on masticatory margin, majors and
larger workers with 3-4 teeth on basal margin. MESOSOMA: Strongly
convex, pronotal margin convex, from dorsal view widest medially. Dorsum
of mesonotum straight. Metanotal suture weakly depressed, not breaking
convex dorsal mesosomal margin. Legs long and slender, first tarsus with
comb-like pubescence on ventral surface. Propodeum arched, posterior dorsal
surface vertical. Propodeal spiracle on posterior lateral surface small, round.
Petiole scale-like with sharp angular apex, may or may not lean anteriorly,
from posterior view, apex sharp rather than rounded. GASTER: First gastral
segment widest, gaster decreasing continuously in size posteriorly. Gaster
oval-shaped, pygidium rounded. COLORATION AND PILOSITY:
Concolorous light brown to dark brown. Cuticle covered in dense, appressed
to erect pilosity. Posterior and lateral cephalic margins lacking erect pilosity,
with short subdecumbent to appressed pilosity. Antennae completely covered
in appressed pubescence. 24-30 long erect hairs surpassing anterior clypeal
margin. Mesosoma completely covered in appressed pubescence. Longest
pilosity on propodeal dorsal surface, majors with up to 12 long erect hairs on
dorsum (0.20 mm). 4-7 long erect hairs on dorsal mesonotal surface. Long
decumbent pubescence on anterior petiolar face. Pubescence on gaster clearly
defined, more so than pubescence on head and mesosoma. Pilosity golden
yellow.

Distribution

North Central to South Central China (105°E to 120° E; 20° N to 42°N).

340 Sociobiology Vol. 52, No. 2A, 2009

Type Material: 6 syntype workers; Lectotype worker (W.M. Wheeler); 1

Paralectotype worker examined: CHINA (MCZC).

Material Examined
CHINA- Foochow: N. Gist Gee. Guangdong: 23°25’50” N, 116°12’40”E.
Gangxi: 22°49’ N, 108°19’, E, Nanning, Zhou. Guiyang: Guiyang, 26°35’ N
106°43’E. Hebei: Schijashuang, 38°02’ 29” N, 114° 28’43”E. Hunan:
Hengshan, 8.ix.2003, J. Huang leg. Hupeh: Lichuan Distr. Suisapa,
31.vii.1948, Gressitt & Djou. Manking: G.P. Dung. Mokanshan: N. Gist
Gee, Paralectotye (MCZC). Soochow: N. Gist Gee, Syntype (MCZC).
Shannxi: Xi’an 34°15’44”N, 108°56’16”, E, Zhou. Szechwan: MTS 4600 ft.
above Kuanhsein City, 9.iii.45. Youluslan: Silvestri.

Habitat and Biology: Unknown

Species Summary
We consider L. sinense var. sericatum and L. dentimandibulum to be
synonyms of L. sinense. Discriminant Function analysis revealed significant
morphometric variation between L. sinense and L. sinense var. sericatum.
However, we believe that such differences are attributed to having a small
sample size, covering a limited geographic range. After analyzing the type
series of both L. sinense and L. sinense var. sericatum, along with fresh
material obtained from Dr. Zhou, we recognize the extreme degree of
polymorphism within this species; therefore we consider these two taxa to be
synonymous. The second synonymy is that of L. dentimandibulum. Even
though the type for this species was not seen, the diagnostic characters of L.
dentimandibulum are consistent with the majors of L. sinense. Again, the high
degree of polymorphism is to blame for the recognition of the two taxonomic
entities. Larger workers and majors of L. sinense present the same diagnostic
characteristics that were used to define L. dentimandibulum as a valid taxon.
The principal character is the presence of 3-4 teeth along the basal
mandibular margin; this character is present in the majors and larger workers
of L. sinense.

Liometopum minimum Zhou = Chronoxenus myops (Forel)

Figs 55-60
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 341

Liometopum minimum Zhou 2001: 557, worker; CHINA; Guangxi [10

paratypes Examined], New Synonymy

Bothriomyrmex myops Forel 1895: 471, workers; [6 Syntypes Examined,

MCZC]

Chronoxenus myops (Forel 1895):471; Transferred to Chronoxenus,

Dubovikoff 2005: 93

Diagnosis:

After reviewing paratype material sent from China by Dr. Zhou, we

have concluded that this is not a member of Liometopum, but rather a member
of the genus Chronoxenus. Prior to the new combination of Dubovikoff
(2005), members of the genus Chronoxenus were considered to be part of the
genus Bothriomyrmex. Syntypes of B. myops (now C. myops) were examined
at the MCZC. Two reported species of Chronoxenus have been recorded in
China, C. dalyi (Forel) and C. myops (Forel) (Bolton 1995). We have
concluded that Liometopum minimum is a synonym of Chronoxenus myops.
The diagnostic character used in identifying this specimen was the definition
of the apical teeth. Specifically, the apical tooth is extremely well defined and
large.

Type material: 10 paratype workers of L. minimum examined: CHINA: 2

deposited at CWEM, 3 deposited at MCZC, 5 deposited at GNUC. 6 syntypes
of Chronoxenus myops examined: INDIA: Deposited at MCZC.

Material Examined

CHINAU- Guangxi: Shiwandashan Natural Reserve, 26-ix-2000, Zhou;

Shan-Yi; Shiwandashan Natural Reserve, 30.x.2001, Zhou. INDIAU-
Smythies.
DISCUSSION

Species Delineation and Additional Notes

Differences in morphology, geographic distribution and ecology of the
examined taxa infer that the genus Liometopum is divided into seven extant
species. Three species occur in North America. Using Mayr’s Biological
342 Sociobiology Vol. 52, No. 2A, 2009

Species Concept, we hypothesize that these three species are valid taxa
because of the significant morphological differences, the unique distribution
patterns observed, and the ecological differences that occur with each taxon.
The Old World species each follow different evolutionary lineages and
constitute reproductively isolated species. No hybrids or clines between
species were observed within the seven recognized taxa, which indicate that
the seven species are reproductively isolated in their corresponding
geographic ranges.
Our results are a hypothesis of the possible structure of the taxonomic
species arrangement within the genus Liometopum, and are not the final
taxonomic assessment of this genus. Our review should be simply viewed as
an attempt to arrange a group with a confusing taxonomic past.
Future work should include a deeper examination of the Old World
species, including detailed gyne and male caste descriptions for L. sinense
and L. lindgreeni when available to bring more clarity to the taxonomic
arrangement of species of the Old World.
The generic level phylogeny of Liometopum has been addressed by
Shattuck 1992, 1995, Chiotis et al. 2000, Brady et al. 2006, and Moreau et al.
2006. Shattuck’s analysis used morphological characters to derive a
phylogeny, placing Tapinoma, Technomyrmex, and Liometopum in the same
Dolichoderine clade. The molecular analyses presented by Moreau et al.
(2006) and Brady et al. (2006) coincide with Shattuck’s clades, but place
Liometopum as a sister genus to Tapinoma with Technomyrmex as a basal
genus (Chiotis, et al. 2000; Brady et al., 2006; Moreau et al., 2006). There are
no analyses that consider all Liometopum species.
Bolton (1995) and Bolton et al. (2006) indicate that this genus is well
represented in the fossil record (9 species of extinct representatives).
Taxonomic assessments of the extinct members of this genus remain a
possibility for further investigation. No works to date have analyzed the
Liometopum fossil record. From the available works, it is believed that
Liometopum is of Early Paleocene origin (Moreau et al. 2006). Such figures
are derived from fossil calibrations used in the Moreau et al. (2006)
phylogenetic analyses.

STATISTICAL ANALYSIS

Discriminant analysis and canonical correlation plots were used to

evaluate evidence for the separation of species based on morphometric
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 343

characters. In genera where multiple characters are needed to identify species,

discriminant analysis allows us to assign statistical values to morphometric
measurements (Seifert, 2003). Consequently, we can better understand
morphometric variation within a species and use these characters in species
delineation. Using the statistically significant morphometric characters
(derived from the discriminant analysis), L. apiculatum, L. luctuosum, L.
occidentale, L. microcephalum, L. sinense, and L. sinense var. sericatum were
separated. Discriminant analysis alone does not yield a clear understanding of
species limits, but can offer some resolution to certain species groups (Moder
et al. 2007; Schlick-Steiner et al. 2003), but we do not rely completely on
morphometric analysis for taxonomic validity. Additional morphological
characters yield the strongest evidence for our taxonomic conclusions. For
each taxon, a minimum of 10 specimens were measured (when available).
Only one specimen of L. lindgreeni (holotype) and 8 specimens of L. sinense
var. sericatum (syntypes and lectotypes) were obtained and measured. The
lack of data for L. lindgreeni yields unreliable results. To assess the species
validity of L. lindgreeni, meristic characters were used (see L. lindgreeni
Species Diagnosis).
Our first discriminant analysis and canonical correlation plot (Fig.
106) compared the three recognized species in the New World and one
morphologically similar species from the Old World, L. microcephalum.
Eleven morphometric characters were considered in the Discriminant
Analysis. Eye Length, Scape Length and Petiole Height yielded F values
greater than 4 and were selected for Canonical Correlation Analysis (Table
1). Ninety-five percent confidence intervals were calculated and plotted as
ellipses on the figures. All four species separate into individual taxa,
supporting our interpretation of the characteristics. Overlap does occur, but
we attribute this overlap to the consistent polymorphism of the species and
ranges of morphometric data. Liometopum microcephalum, of which L.
occidentale was considered a potential synonym, clearly separates from the
New World species and thus L. occidentale should retain a valid species
status. Overall species definitions based on morphometric character analysis
supports our taxonomic assessment.
Our second discriminant analysis and canonical correlation plot (Fig.
107) analyzed the four Old World taxa of Liometopum. Eleven morphometric
characters were considered by the discriminant analysis. Using forward
stepwise variable selection, four characters had F values greater than 4.0; the
three most informative characters were selected. These characters are:
344 Sociobiology Vol. 52, No. 2A, 2009

Petiolar Index, Scape Length, and Scape Index. The three factors were plotted
and 95 percent confidence ellipses were plotted as ellipses on the figures.
There is little overlap between L. microcephalum and the other Old World
species. Some overlap occurs between L. sinense and our proposed synonym
L. sinense var. sericatum. The lack of constant overlap between L. sinense
and L. sinense var. sericatum may be attributed to the limited sample size of
L. sinense var. sericatum, from the single type locality. In the future, if more
specimens are available we suggest a larger sample size to be analyzed. Such
results should clarify any taxonomic doubts regarding this proposed
synonymy. Liometopum lindgreeni groups with L. sinense, however, only one
specimen of L. lindgreeni was available. Liometopum lindgreeni appears to
be easily separated from L. sinense by meristic characters that were not part
of the statistical analysis (see L. lindgreeni discussion).
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 345

FIGURES AND TABLES

1 2

3 4

5 6
Figures 1-6 Liometopum apiculatum; (1,2) Worker head and mesosoma, [MEXICO, Queretaro,
Queretaro, La Barreta, 20˚ 49.35” N 100˚ 30.94” W (CWEM)]; (3,4) Gyne head and mesosoma,
[MEXICO, San Luis Potosi, 41 km SE Sto. Domingo (CWEM)]; (5,6) Male head and mesosoma,
[USA, California, San Juan Co. San Juan River mile 165, (CASC)].
346 Sociobiology Vol. 52, No. 2A, 2009

7 8

9 10

Figures 7-8 Liometopum lindgreeni; (7,8) Worker holotype head and mesosoma, [INDIA, Assam,
Debrugarh, (MHNG)]; Figures 9-10 Liometopum sinense; (9,10) Worker syntype head and
mesosoma, [CHINA, Soochow, (MCZC)].
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 347

11 12

13 14

15 16

Figures 11-16 Liometopum luctuosum; (11, 12) Worker head and mesosoma, [USA, California, San
Bernardino Co, Big Pine Flat, (CWEM)]; (13, 14) Gyne head and mesosoma [USA, Arizona,
Coconino Co., San Francisco Mts. (MCZC)]; (15, 16) Male syntype head and mesosoma [USA,
California, Lake Tahoe (MHNG)].
348 Sociobiology Vol. 52, No. 2A, 2009

17 18

20
19

21 22

Figures 17-22 Liometopum microcephalum; (17, 18) Worker head and mesosoma,
[CZECHOSLOVAKIAU- Moravia, mer. Lednice (CWEM)]; (19, 20) Gyne head and mesosoma,
[UROMANIAU, Juderul, Siba, (CASC)]; (21, 22) Male head and mesosoma, [SERBIA,
Vojvodina, Vrdnik, Lediniko Jezero (CASC)].
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 349

29 30

31 32

33 34

Figs. 29-34. Liometopum orientale and Liometopum microcephalum (modified from Kupyanskaya
1988).Liometopum orientale: (29) Worker head; (30) Worker petiole; (31) Liometopum
microcephalum:Worker petiole; (32) Gyne head; (33) Male head; (34) Male messosoma.
350 Sociobiology Vol. 52, No. 2A, 2009

35 36
37

39
38

40 41

42

43 44

Fig. 35-39 Liometopum apiculatum male genitalia, [USA, California, San Juan Co. San Juan River
mile 165, (CASC)] ; (35) genital capsule; (36) paramere; (37) subgenital plate; (38) aedeagus; (39)
volsella.
Fig. 40-44 Liometopum luctuosum male genitalia, [USA, Arizona, Yavapai Co, Granite Dells 4 mi.
N. Prescott, (CASC)]; (40) genital capsule; (41) paramere; (42) subgenital plate; (43) aedeagus;
(44) volsella.
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 351

45 46
47

48

49

51 52
50

53 54

Fig. 45-49 Liometopum microcephalum male genitalia, [SERBIA, Vojvodina, Vrdnik, Lediniko
Jezero (CASC)] ; (45) genital capsule; (46) paramere; (47) subgenital plate; (48) aedeagus; (49)
volsella
Fig. 50-54 Liometopum occidentale male genitalia, [USA, California, Riverside, San Jacinto Mts.
(CWEM); (50) genital capsule; (51) paramere; (52) subgenital plate; (53) aedeagus; (54) volsella.
352 Sociobiology Vol. 52, No. 2A, 2009

55
56

58
57

59
59

Figs. 55-60. Liometopum minimum= Chronoxenus myops (modified from Zhou 2001). (55, 56)
Worker head and mesosoma; (57, 58) Gyne head and mesosoma; (59, 60) Male head and
mesosoma.
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 353

61 62

63 64

65 66

67 68

Figs. 61-64. Liometopum apiculatum. (61) Gyne anterior wing, (A) Costal Cell, (B) Median Cell,
(C) Submedian Cell, (D) First Submarginal Cell (E) First Discodial Cell, (F) Second Discodial
Cell, (G) Second Submarginal Cell, (H) Marginal Cell; (62) Gyne hind wing, (A) Median Cell, (B)
Hamuli; (63) Male anterior wing; (64) Male hind wing.
Figs. 65-68. Liometopum luctuosum; (65) Gyne anterior wing; (66) Gyne hind wing; (67) Male
anterior wing; (68) Male hind wing.
354 Sociobiology Vol. 52, No. 2A, 2009

Fig. 69. Liometopum apiculatum distribution .

Fig. 70. Liometopum lindgreeni distribution.

Del Toro, I. et al.- Revision of the Ant Genus Liometopum 355

Fig. 71. Liometopum luctuosum distribution.

Fig. 72. Liometopum microcephalum distribution.

356 Sociobiology Vol. 52, No. 2A, 2009

Fig. 73. Liometopum occidentale distribution.

Fig. 74. Liometopum sinense distribution.

Del Toro, I. et al.- Revision of the Ant Genus Liometopum 357

Figs. 75-80. Digital Micrographs of Liometopum apiculatum. (75-76) Worker head and mesosoma
[MEXICO , Queretaro, Queretaro, La Barreta, 20˚ 49.35" N 100˚ 30.94" W (CWEM)]; (77-78)
Gyne head and mesosoma [MEXICO , San Luis Potosi, 41 km SE Sto. Domingo (CWEM)]; (79-
80) Male head and mesosoma [US A, Colorado, Colorado Springs, (MCZC)].
358 Sociobiology Vol. 52, No. 2A, 2009

Figs. 81-86. Digital Micrographs of Liometopum luctuosum. (81-82) Worker head and mesosoma
[MEXICO , Coahuila, 13 km E San Antonio, (CWEM)]; (83-84) Gyne head and mesosoma [US A,
Arizona, Coconino Co., San Francisco Mts. (MCZC)]; (85-86) Male head and mesosoma [US A,
Arizona, Cochise Co., Chiricahua Mts. (MCZC)].
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 359

Figs. 87-92. Digital Micrographs of Liometopum occidentale. (87-88) Worker head and mesosoma
[US A, California, Riverside, San Jacinto Mts. (CWEM)]; (89-90) Gyne head and mesosoma, [US
A, Calaveras, Riverside, 4.8 km S. West Point (CASC )]; (91-92) Male head and mesosoma, [US
A, California, Riverside, San Jacinto Mts. (CWEM)].
360 Sociobiology Vol. 52, No. 2A, 2009

Figs. 93-98. Digital Micrographs of Liometopum microcephalum. (93-94) Worker head and
mesosoma [CZECHOSLOVAKIA - Moravia, mer. Lednice (CWEM)]; (95- 96) Gyne head and
U

mesosoma [ RO MANIA , Juderul, Siba, (CASC )]; (97-98) Male head and mesosoma, [SERB IA,
U U

Vojvodina, Vrdnik, Lediniko Jezero (CASC )].

Del Toro, I. et al.- Revision of the Ant Genus Liometopum 361

Figs. 99-105. (99-100) Liometopum lindgreeni worker head and mesosoma [INDIA, Assam,
Debrugarh, (MHNG)]. (101-102) Liometopum sinense Worker head and mesosoma [CHINA,
Soochow, (MCZC)]; (103) Sceptobius dispar (MCZC); (104) Dinardilla liometopi, (MCZC); (105)
Liometophilus manni, (LACM).
362 Sociobiology Vol. 52, No. 2A, 2009

Fig. 106. Three Factor Canonical Analysis Plots: Factor 1= Eye Length; Factor 2= Scape Length;
Factor 3= Petiole Height.
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 363

Fig. 107. Three factor Canonical Analysis Plots: Factor 1= Petiolar Index; Factor 2= Scape Length;
Factor 3= Scape Index.
364 Sociobiology Vol. 52, No. 2A, 2009

ACKNOWLEDGEMENTS

We would like to thank Francisco Serna, Shawn T. Dash, Samuel Del

Toro, and Paul Lenhart from the University of Texas at El Paso for their
valuable input and review of the manuscript. We would also like to extend
our appreciation to Dr. Alex Wild, and Dr. John Longino for their review and
comments on this is manuscript. We thank Dr. Phil Ward for manuscript advice
and identification of specimens from California collections. Special thanks
are extended to Ms. Rebecca Marin for her help in preparation of distribution
maps. Additionally, we would like to thank the instructors Laura Cade and
John Wilson for their influence and direction that led to the creation of this
manuscript. We would like to thank the various curators of the museum
collections including, Stefan Cover and Dr. Gary Alpert (MCZC), Dr. Brian
Fisher
isher and April Nobile (CASC), Weiping Xie, Dr. Brian V. Brown and Roy
Snelling (LACM), Luis Quiroz (UAGC), Dr. Robert Jones and Maya Rocha
(UAQC), Dr. Ted Shultz (USNM), and Dr. Bernhard Merz (MNHG).
Additional specimens and distributional data were provided
provi by Dr. Jose-Luis
Navarrete-Heredia, and Miguel Vasquez-Bolaños
Bolaños and Georgina A. Quiroz-
Quiroz
Rocha. We also would like to thank Dr. Zhou for fresh material collected in
China and Dr. John Longino for specimen distribution data. We especially
appreciate the helplp of Dr. Gary Alpert, Stefan Cover, Dr. Brian Fisher and
April Nobile for allowing us to utilize imaging equipment. Travel for the
MCZC visit was provided by the Ernst Mayr Grant. Funding for this project
was provided by NSF Grant DBI 0405470, W. Mackay Principal
P Investigator,
the University of Texas at El Paso Honors Program and NSF Grant, Colorado
Del Toro, I. et al.- Revision of the Ant Genus Liometopum 365

Alliance for Graduate Education and the Professoriate; HRDB-639653,

NSF06-552.

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