Hindawi Publishing Corporation
Psyche
Volume 2012, Article ID 850893, 7 pages
doi:10.1155/2012/850893
Research Article
First Record of Pyramica epinotalis (Hymenoptera: Formicidae)
for the United States
Xuan Chen,1 Joe A. MacGown,2 Benjamin J. Adams,1 Katherine A. Parys,1
Rachel M. Strecker,1 and Linda Hooper-Bui1
1 Department
2 Mississippi
of Entomology, Louisiana State University Agricultural Center, Baton Rouge, LA 70803, USA
Entomological Museum, Mississippi State University, Mississippi State, MS 39762, USA
Correspondence should be addressed to Joe A. MacGown, jmacgown@entomology.msstate.edu
Received 29 March 2012; Accepted 10 April 2012
Academic Editor: David Roubik
Copyright © 2012 Xuan Chen et al. This is an open access article distributed under the Creative Commons Attribution License,
which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
Pyramica epinotalis is an arboreal dacetine ant previously known only from Brazil, Costa Rica, Ecuador, and southern Mexico. Here
we report the first records of P. epinotalis for the United States. Collections were made in three parishes across southern Louisiana
in cypress-tupelo swamps using floating pitfall traps placed in floating vegetation and arboreal pitfall traps placed on trunks and
limbs of three wetland tree species. One additional specimen of this species was collected in Highlands County, Florida. Based
on collections of specimens in Louisiana, including multiple dealate females at different localities, P. epinotalis appears to be well
established in this state. We discuss the design and implementation of modified arboreal pitfall traps that were instrumental in this
discovery.
1. Introduction
The tribe Dacetini (Hymenoptera: Formicidae) is composed
of small, cryptic, predatory ants that typically occur in soil
and/or leaf litter where they feed on various minute arthropods [1, 2]. Species in this group show great diversity in
predatory strategies, which is reflected in the marked
differentiation between species groups. With their uniquelooking body types and head shapes that are variously
adorned with bizarre station, elongate mandibles with
uniquely arrayed dentition, and as-yet-unexplained cuticular
outgrowths called spongiform tissue, members of this group
are among the most unusual in the ant world. This large
and diverse tribe includes more than 900 described species
worldwide, of which 327 are in the genus Pyramica [3].
Primarily considered a tropical group, only 41 species of
Pyramica have been reported from the USA. Thirty-seven of
these species occur in the southeastern United States [4]. Five
species of the related Strumigenys are known from the same
region [5]. The relatively high density of dacetine species in
the Southeast is likely due to the humid, subtropical climate
and mild winters typical of the region and the availability of
large continuous tracts of forested habitats, which appear to
facilitate establishment of these species’ colonies.
Currently, nine introduced dacetine species are known
from the southeastern USA including Pyramica eggersi (Emery) (Florida), P. gundlachi (Roger) (Florida), P. hexamera
(Brown) (Alabama, Florida, Louisiana, and Mississippi), P.
margaritae (Emery) (Alabama, Florida, Georgia, Louisiana,
and Mississippi), P. membranifera (Emery) (Alabama,
Arkansas, Florida, Georgia, Louisiana, Mississippi, North
Carolina, and South Carolina), Strumigenys emmae (Emery)
(Florida), S. lanuginosa Wheeler (Florida), S. rogeri Emery
(Florida), and S. silvestrii Emery (Alabama, Florida, Georgia,
Louisiana, and Mississippi) [6]. Pyramica subnuda MacGown and Hill, which is in the schulzi species group, may
also be introduced despite the fact that it was described from
a specimen collected in Mississippi as other members of this
group are tropical [4]. Here, we present records of another
introduced dacetine ant from South America, P. epinotalis
(Weber), which also belongs to the schulzi species group.
2
2. Taxonomy and Identification
Weber described Strumigenys (Cephaloxys) studiosi subsp.
epinotalis (= P. epinotalis) in 1934 from specimens collected
by G. C. Wheeler from Costa Rica, Prov. Limon, Estrella
Valley, Talia Farm, 18.vi.1924 [7]. Strumigenys (Cephaloxys)
skwarrae, a synonym of P. epinotalis, was described by W. M.
Wheeler in 1934 from specimens collected in Tillandsia streptophylla Scheidweiler (Bromeliaceae) in Tlacocintla, Mexico
by E. Skwarra in 1929 [8]. In 1953, Brown synonymized S.
skwarrae with S. epinotalis and transferred it to Smithistruma
[9]. Bolton later moved the species to Pyramica in 1999 [10],
Baroni Urbani and De Andrade synonymized Pyramica with
Strumigenys in 2007 [11], and Bolton and Alpert reconfirmed
Pyramica as a valid genus in 2011 [12].
2.1. Worker (Figures 1(a)–1(c)) (Description Modified from
Bolton [13]). Total length 1.9–2.1 mm. Head wedge-shaped.
Color yellowish-brown, appendages only slightly paler.
Entire head including clypeus with reticulate-punctate sculpture. Mandibles subtriangular, lacking diastema; nine acute
teeth present following basal lamella; third tooth from basal
lamella spiniform, elongate and distinctly longer than other
teeth, with subsequent teeth alternating in length with the
fifth being longer than the fourth and the seventh being
longer than the sixth, and the remaining two teeth smaller
and blunter. Clypeus somewhat pentagonal shaped, narrowing anteriorly, and with anterior margin slightly convex.
Dorsum of clypeus with numerous clavate hairs directed
anteriorly or away from midline of clypeus; clypeal margin
with similarly shaped clavate hairs all curving anteriorly
toward midline of clypeus; remainder of head with slightly
larger clavate to spoon-shaped hairs that curve toward
midline of head; elongate flagelliform cephalic hairs absent;
and leading edge of scape with a row of elongate, curved
hairs, all of which curve toward the base of the scape or are
directed downward. Eye large with 5–7 ommatidia in longest
diameter.
Mesosoma, including sides, and petiole with distinct
reticulate-punctate sculpture; postpetiole disc lacking sculpture, smooth. Pronotum with an arched transverse ridge with
rowed, erect spoon-shaped hairs; pronotal humeral hairs
absent; mesonotum with appressed spoon-shaped hairs. Propodeal spines somewhat short and dentiform, directed upward; propodeal declivity bordered by a high arched lamella
on each side. Petiole with longitudinal spongiform crest
ventrally and fan-shaped, spongiform bodies present posteriorly; elongate, spoon-shaped hairs present dorso-posteriorly.
Postpetiole with large, spongiform mass ventrally, but
becoming a lamina-like structure posteriorly; with elongate,
spoon-shaped hairs present dorsoposteriorly. First gastral
tergite smooth and shining except for basigastral costulae,
which are distinct and extend to at least the basal quarter
to third of the length of the tergite. Several to numerous
suberect to erect, elongate, thickened hairs present on first
tergite.
2.2. Alate Female (Figures 1(d) and 1(e)). Similar to worker,
but larger (total length approximately 2.5 mm), ocelli
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present, mesosoma enlarged with typical modifications for
flight muscles, wings present, and katepisternum mostly
lacking sculpture.
This species is easily differentiated from other species
known from the USA by the combination of having the
third tooth (from basal lamella) on mandible longer than the
other teeth, the mesosoma of the worker being completely
reticulate-punctate, having a curved row of spoon-shaped
hairs on the pronotal dorsum, a distinct propodeal lamella, a
ventral spongiform crest beneath the petiole, and fan-shaped
patches of spongiform tissue on the petiole and postpetiole.
Currently, the only other species reported from the USA
with which P. epinotalis might be confused is P. margaritae,
another introduced species in the schulzi group. Pyramica
margaritae is the only other species known to occur in
the United States that has sculpture on the entire side of
the mesosoma; however, P. margaritae lacks a curved row
of spoon-shaped hairs on the pronotal dorsum, has much
longer propodeal spines, lacks a propodeal lamella, lacks
spongiform bodies beneath the petiole, and has reduced
spongiform tissue present beneath the postpetiole.
3. Natural History
Although the vast majority of dacetine ants nest in soil and
leaf litter, members of the schulzi species group are typically associated with plants, and several species have been
recorded from epiphytes or plant cavities [13]. Many species
in this group also differ in that workers have enlarged
compound eyes, as compared with their epigeic and hypogeic
relatives. Similar to most members of this group, P. epinotalis
also has enlarged eyes and is thought to be an arboreal
species. Weber described this species in 1934 from specimens
collected by George Wheeler in 1924 in an Atlantic slope wet
forest in Costa Rica, but he did not indicate whether the
ants were collected arboreally or in litter [7]. Collecting in
the same region years later, Longino reported that multiple
litter samples from near the type locality did not yield
specimens of this species, which suggests that perhaps this
species might be arboreal [14]. In 1934, Wheeler reported
that Dr. Skwarra discovered four colonies of this species
(reported as Strumigenys skwarrae Wheeler) in Tillandsia
streptophylla at two localities in Mexico in 1929 [8]. Bolton
[13] reported that collections of this species were made in
Mexico by Dressler and by Dejean, both of whom worked
with epiphytes, which implies that their specimens were also
from epiphytes. More recently, Rider reported collections of
this species in the canopy in Ecuador, which further validates
its status as an arboreal species [15].
4. Methods
A single alate female was collected by Mark Deyrup on 14
August 2009 using a Townes Malaise trap placed in Florida
scrub habitat in Highlands County. Scrub habitat was located
near a “bayhead,” a periodically flooded forest dominated by
magnolia (Magnoliaceae) and gordonia (Theaceae) trees.
Deyrup compared his specimen to specimens identified by
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3
(a)
(b)
(c)
(e)
(d)
Figure 1: Pyramica epinotalis: (a) full face view of worker, (b) lateral view of worker, (c) dorsal view of worker, (d) full face view of female,
and (e) lateral view of alate female. Scale bar equals 200 µm.
Figure 2: Cypress-tupelo swamp north of Gramercy, Ascension
Parish, Louisiana.
W. L. Brown, which were collected by R. L. Dressler in
Ocosingo, Chiapas, Mexico in 1954 (M. Deyrup, pers.
comm.).
Collections of ants in Louisiana were made in cypresstupelo swamps from spring through fall of 2009 and 2010
on a privately owned tract of land north of Gramercy in
Figure 3: Cypress-tupelo swamp in Jean Lafitte National Historical
Park and Preserve, Jefferson Parish, Louisiana.
Ascension Parish (30◦ 09′ 48′′ N 90◦ 48′ 39′′ W) (Figure 2) and
during the late spring and summer of 2011 (May to September) in Jean Lafitte National Historical Park and Preserve
in Jefferson Parish (29◦ 47′ 38′′ N 90◦ 06′ 17′′ W) (Figure 3)
and Maurepas Swamp Wildlife Management Area (Western
Tract) in Saint James Parish (30◦ 06′ 56′′ N 90◦ 40′ 47′′ W)
4
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Figure 5: Floating pitfall trap shown floating on surface of water in
cypress-tupelo-blackgum freshwater swamp.
Figure 4: Cypress-tupelo swamp in Maurepas Swamp Wildlife
Management Area, Saint James Parish, Louisiana.
(Figure 4). All locations are within the Mississippi River
deltaic plain in coastal Louisiana.
Cypress-tupelo swamps in Louisiana are characterized
and dominated by the presence of bald cypress, Taxodium
distichum (L.) Rich (Cupressaceae), water tupelo, Nyssa
aquatica L. (Cornaceae), and red maple, Acer rubrum L. var.
drummondii (Hook. and Arn. Ex Nutt.) Sarg. (Aceraceae)
[16]. Collections from floating vegetation were made using
a floating pitfall trap in Gramercy, LA, as described by
Parys and Johnson [17]. These collections were made as
part of a larger study to examine the biodiversity of insects
associated with invasive aquatic vegetation. In addition to the
characteristic tree species, this site has dense mats of invasive
aquatic vegetation formed from common salvinia (Salvinia
minima Baker (Salviniaceae)), water hyacinth (Eichornia
crassipes (Mart.) Solms (Pontederiaceae)), and water pennywort (Hydrocotyle sp. (Araliaceae)). Traps were filled with
ethylene glycol as a preservative and emptied at two-week
intervals.
Arboreal collections were made on bald cypress (T. distichum), water tupelo (N. aquatica), and red maple (A.
rubrum var. drummondii) because they were the most common tree species observed. We chose three to six of each of
the aforementioned tree species spaced >50 meters apart for
trap deployment. We placed both a cup trap and a bottle trap
in each selected tree’s canopy and tied a trunk trap at breast
height onto the trunk of each tree. Each of the traps was
filled with approximately 50 mL ethylene glycol as a specimen
preservative.
4.1. Trap Designs
4.1.1. Floating Pitfall Trap (Figure 5). Floating pitfall trap
designs were described in detail by Parys and Johnson [17].
4.1.2. Cup Trap (Figures 6(a)–6(e)). We modified a trap that
was originally described by Oliveira-Santos et al. [18]. Cup
traps were constructed from a single 400 mL tricorner plastic
beaker (Figure 6(a)). Three holes were bored into a flange
on the rim of the cup (Figure 6(b)). A 6.35 mm cotton rope
strand was then threaded through each of the three holes, and
hot glue was used to secure each rope (Figure 6(c)). Braided
nylon rope was used to link the 6.35 mm cotton rope together
at a common point above the trap (Figure 6(d)). The excess
cotton rope below the flange on the cup was then taped to
the sides of the cup (Figure 6(e)).
4.1.3. Bottle Trap (Figures 7(a)–7(h)). We modified the bottle
trap design as described in Kaspari [19]. The traps were
created using an inverted 600 mL drink bottle with the base
removed (Figure 7(a)). Three holes were bored into the edge
of the base (now top) of the container (Figure 7(b)). A foam
square (10 cm × 10 cm) was fitted around the base opening
of the bottle allowing for at least 2 cm between the opening
and the outer edge of the foam square (Figure 7(c)). Fishing
line (40 lb test) was tied through the holes in the base of the
bottle and around the foam square to connect the square
to the bottle (Figure 7(d)). We attached two plastic dowels
(6.35 mm dia) to the 10 cm long ends of a length of canvas
(40 cm × 10 cm) using hot glue. A hole was burned through
the canvas and dowels using a soldering iron; a zip-tie was
then attached to the dowel through this hole. A 6 cm hole
was also cut into the middle of the canvas 5 cm from one of
the dowels. Eight 2 cm slits were then cut 4 cm apart into the
canvas around the edge of the hole (Figure 7(e)). The canvas
was attached to the base of the bottle using hot glue applied
to the inside of the bottle (Figure 7(f)). We tied a 28.3 g
fishing weight to the bottle using fishing line (Figure 7(g)).
An optional modification to reduce friction on the branches
when setting up the traps was to cut the extra tail of canvas
to 3 cm width (Figure 7(h)).
4.1.4. Trunk Trap. Trunk trap designs were taken directly
from Pinzón and Spence [20]. The only modification was the
removal of the plastic flues used to funnel insects into the
traps.
4.1.5. Trap Placements. Placement of floating pitfall traps is
discussed in Parys and Johnson [17]. The top quarter of
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5
(a)
of Oliveira-Santos et al. [18]. For bottle trap designs, the
fishing line was connected to the zip-ties attached to the
plastic dowels. It is critical that the edge of the cup trap and
the edge of the foam on the bottle trap be in contact with a
tree branch or trunk to ensure maximum yield of specimens.
After each sampling period, the entire contents of the traps
were removed, new ethylene glycol was added, and the trap
was returned to the canopy.
(d)
(b)
5. Results and Discussion
(e)
(c)
Figure 6: Cup trap design: (a) tricorner beaker, (b) holes bored in
flange of beaker, (c) cotton rope threaded through flange hole and
secured with hot glue, (d) cotton ropes linked together using nylon
rope above trap, and (e) completed trap with excess cotton rope
below lip of flange taped to sides of beaker (cup).
(a)
(b)
(c)
(d)
(e)
(g)
(f)
(h)
Figure 7: Bottle trap design: (a) inverted 600 mL plastic bottle, (b)
hole bored into base of bottle, (c) foam square cut to bottle base, (d)
foam square place around bottle base with fishing line tied through
holes and around square, (e) canvas strip, with hole, slits around
hole, dowels at each end, and zip-ties added, (f) canvas attached to
inside of bottle with hot glue (g) fishing weight added to bottle with
fishing line, and (h) excess canvas trimmed from foam square.
each arboreal trap was brushed with liquid Teflon (Dupont
Polymers, Wilmington, Delaware) to prevent ants from
escaping. Canopy traps were set in the trees using the same
slingshot method as Kaspari [19]. However for ease of
sampling and returning the traps to the canopy, the tie-down
lines were tied together to make a loop similar to the methods
Here we report the first records of P. epinotalis for the United
States. A single alate female was collected by Mark Deyrup
in Highlands County, Florida, on 14 August 2009 using a
Townes Malaise trap (M. Deyrup, pers. comm.). We collected
five females and 14 workers of P. epinotalis in Ascension,
Jefferson, and Saint James Parishes in Louisiana on various
dates from 8 to 21 September 2009 and from 30 May to
23 September 2011. Louisiana collections were made using
floating pitfall traps placed directly upon the surface of the
water and with cup traps, bottle traps, and trunk traps placed
on trunks and branches of three species of trees as described
in the methods section. Louisiana collections were made by
Katherine Parys, Xuan Chen, and Benjamin Adams. Other
collection data are as follows: Florida: Highlands County,
Highlands Park Estates, N27.53864, W081.35071, 14 August
2009, M. Deyrup, Townes trap in scrub habitat (near bayhead). Louisiana: Ascension Parish, Gramercy, N of 61 and
I-10, 30◦ 09′ 48′′ N 90◦ 48′ 39′′ W, 8–21 Sept; 2009, K. A. Parys,
floating pitfall trap in cypress-tupelo freshwater swamp with
dense mats of Salvinia minima on water surface (1 worker).
Jefferson Parish, Jean Lafitte National Historical Park and
Preserve, 29◦ 47′ 38′′ N 90◦ 06′ 17′′ W, 30 May–13 June 2011,
cup trap on Nyssa aquatica branch (1 female); same data
except, trunk trap on trunk of Acer rubrum var. drummondii
(1 worker); same data except, 13 June–18 August 2011,
bottle trap on Taxodium distichum branch (1 female); same
data except, trunk trap on trunk of Taxodium distichum (2
females); same data except, trunk trap on trunk of Nyssa
aquatica (2 workers); and same data except, bottle trap on
Acer rubrum var. drummondii branch (1 worker); same data
except, cup trap on Acer rubrum var. drummondii branch (1
female); same data except, 18 August–23 September 2011,
bottle trap on Nyssa aquatica branch (1 worker); same data
except, trunk trap on trunk of Acer rubrum var. drummondii
(2 workers). Saint James Parish, Maurepas Swamp Wildlife
Management Area (Western Tract), 30◦ 06′ 56′′ N 90◦ 40′ 47′′
W, 9 June–17 September 2011, trunk trap on trunk of Nyssa
aquatica (4 workers); same data except, cup trap on Nyssa
aquatica branch (2 workers).
This is a significant contribution to the distributional
record for this species as previously it had only been
reported from southern Mexico (Veracruz, Chiapas, Quintana Roo, Tlacocintla), Ecuador (Tiputini), Costa Rica
(Limon Province), and Brazil (Mina Gerais) [6, 8, 10].
According to Longino [14], in Costa Rica P. epinotalis is
known only from the southern Atlantic lowlands, south of
Limon. As mentioned in the Natural History section, this
6
species is thought to be primarily an arboreal species. Similarly, collections in Louisiana were all made from arboreal
traps except for a single worker that was collected using a
floating pitfall trap, which was in the same habitat type.
Xuan Chen also collected ants in leaf litter and quadrats
on ground cover in both locations, yet P. epinotalis was not
found. Although the single Florida collection was made in
scrub habitat, the alate female could have flown there from
nearby bayhead habitat. Based on the collections made by
Longino and other records [8, 13–15], this species may prefer
wetland forest habitats. This also appears to be the case with
the Louisiana collections, all of which were made in swampy,
wetland habitats.
It seems likely that P. epinotalis is an introduced species. Evidence for this includes the large geographical gap
between the known distribution and the new records of
this species from Florida and Louisiana. Other exotic ants
(i.e., Brachymyrmex patagonicus Mayr, Pheidole obscurithorax
Naves, Solenopsis invicta Buren, and S. richteri Forel) from
South America have been introduced to the southeastern
states [21–24]. However, these species are native to southern
South America whereas P. epinotalis is more likely native
to northern South America, Central America, or southern
Mexico making this record more unique. Given the relative
novelty of trapping methods used here, it is possible that the
range of this species actually may be much more extensive.
For example, the range of another introduced arboreal
species, Pseudomyrmex gracilis (F.), is almost continuous
from South and Central America to southern Texas and is
now found in Florida, Louisiana, and Mississippi [25, 26].
Dacetine ants native to the USA are not known to nest
arboreally but rather have only been reported to nest in
rotting wood, soil, or litter [5]. Consequently, most ant
collectors in the USA would not consider searching trees for
dacetines, given that only a few species, which are primarily
tropical, nest arboreally [13].
Typical methods for arboreal collections in USA such
as baiting or beating vegetation (unless they are present on
outer limbs) would not likely yield dacetines even if they
were present. Dacetines are specialized predators [2] and
likely would not be attracted to standard baits used to attract
other generalist species of ants. Due to their coloration
and cryptic habits, these ants could easily be overlooked
during visual searches on tree trunks. Furthermore, few ant
collectors spend time in wetlands, as most ants in this
region are terrestrial. If P. epinotalis is truly associated with
wetland forests, this species could easily have been overlooked. Collections along the eastern edge of Mexico and into
Texas could greatly enhance our knowledge of this species’
distribution and provide information on whether or not
it is truly an introduced species. Until such time as these
collections can be made, we tentatively consider this species
to be an exotic species to the US.
Acknowledgments
The authors thank Dr. Julie L. Witbeck with Jean Lafitte
National Historical Park and Preserve for facilitating work
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at that location. They also thank Mr. James Boyce for access
to his property in Gramercy as well as Anna Mészáros and
Jordan Fryoux for assistance in the field. Thanks to Mark
Deyrup for his comments and specimen data, this project
was supported by the Northern Gulf Institute (L. HooperBui, Principle Investigator), the Gulf of Mexico Research
Initiative (year 1 funds through LSU to L. Hooper-Bui), Mississippi Agricultural and Forestry Experiment Station State
Project MIS-311080, the USDA-ARS Areawide Management
of Imported Fire Ant Project (Richard L. Brown, Principal
Investigator), and Louisiana Department of Wildlife and
Fisheries Contracts nos. 673252, 686647, and 696303 (Seth
J. Johnson, Principal Investigator). The funders had no role
in study design, data collection and analysis, decision to
publish, or preparation of this manuscript. This paper was
approved for publication by the Louisiana State University
Agricultural Experiment Station and approved for publication as journal article number. J-12146 of the Mississippi
Agricultural and Forestry Experiment Station, Mississippi
State University.
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