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A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 119
A revision of the giant Amazonian ants of the genus
Dinoponera (Hymenoptera, Formicidae)
Paul A. Lenhart1,†, Shawn T. Dash2,‡, William P. Mackay2,§
1 Department of Entomology, Texas A&M University, 2475 TAMU, College Station, Texas USA 2 Department
of Biological Sciences, University of Texas at El Paso, 500 West University Avenue, El Paso, Texas 79968
† urn:lsid:zoobank.org:author:6CA6E3C7-14AA-4D6B-82BF-5577F431A51A
‡ urn:lsid:zoobank.org:author:9416A527-CCE2-433E-A55F-9D51B5DCEE8A
§ urn:lsid:zoobank.org:author:70401E27-0F2E-46B5-BC2B-016837AF3035
Corresponding author: Paul A. Lenhart (palenhart@neo.tamu.edu)
Academic editor: Wojciech Pulawski|Received 16 November 2012|Accepted 7 January2013|Published 20 March2013
urn:lsid:zoobank.org:pub:10404A9C-126A-44C8-BD48-5DB72CD3E3FF
Citation: Lenhart PA, Dash ST, Mackay WP (2013) A revision of the giant Amazonian ants of the genus Dinoponera
(Hymenoptera, Formicidae). Journal of Hymenoptera Research 31: 119–164. doi: 10.3897/JHR.31.4335
Abstract
Dinoponera Roger 1861 has been revised several times. However, species limits remain questionable due
to limited collection and undescribed males. We re-evaluate the species boundaries based on workers and
known males. We describe the new species Dinoponera hispida from Tucuruí, Pará, Brazil and Dinoponera
snellingi from Campo Grande, Mato Grosso do Sul, Brazil and describe the male of Dinoponera longipes
Emery 1901. Additionally, we report numerous range extensions with updated distribution maps and
provide keys in English, Spanish and Portuguese for workers and known males of Dinoponera.
Resumen
El género Dinoponera Roger 1861 ha sido revisado varias veces. Sin embargo, la distribución geográca
de las especies de este género todavía es cuestionable debido a colecciones limitadas, y a que en general los
machos continúan sin descripción. Reevaluamos los límites geográcos de las especies utilizando carac-
teres merísticos y morfométricos basados en obreras y machos conocidos. Describimos las nuevas especies
Dinoponera hispida de Tucuruí, Pará, Brasil, y Dinoponera snellingi de Campo Grande, Mato Grosso do
Sul, Brasil, y describimos el macho de Dinoponera longipes Emery 1901. Además, reportamos numerosas
extensiones nuevas de las distribuciones geográcas con sus respectivos mapas actualizados, y proporcio-
namos claves de identicación en inglés, español, y portugués para las obreras y los machos conocidos del
género Dinoponera.
JHR 31: 119–164 (2013)
doi: 10.3897/JHR.31.4335
www.pensoft.net/journals/jhr
Copyright P. A. Lenhart et al. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY),
which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
RESEARCH ARTICLE
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
120
Resumo
O género Dinoponera Roger1861, é tem revisados varios vezes. No entanto, a claricação das especies
ainda segue questionável devido a um coleção limitado, e a que em geral os representantes do sexo mascu-
lino continuam sem descrição. Reavaliamos os limitas de espécies utilizado caráteres merísticos e morfo-
métricos, baseado em operários e machos conhecidos. Descrevemos novas espécies Dinoponera hispida de
Tucuruí, Para, Brasil e Dinoponera snellingi de Campo Grande, Mato Grosso do Sul, Brasil, e descrevemos
o macho de Dinoponera longipes Emery 1901. Además, reportamos distribuições geográcas com seus
respetivos mapas atualizados, e proporcionamos chaves de identicação em inglês, espanhol, e português
para operários e machos conhecidos do género Dinoponera.
Keywords
South America, Brazil, taxonomy, gamergate, ponerinae
Introduction
Dinoponera Roger 1861 is a strictly South American genus in the subfamily Ponerinae,
tribe Ponerini, commonly called tocandiras or giant Amazonian ants (Zahl 1959, Four-
cassié and Oliviera 2002, Haddad et al. 2005). ese ants are generally less well known
than Paraponera clavata (Fabricius 1775), the bullet ant or hormiga bala (Haddad et al.
2005), yet Dinoponera workers may surpass 3 cm in total body length, making them
the largest in the world. e genus has been found from montane rainforest on the
eastern slope of the Andes in Perú, Ecuador and Colombia to savannah and lowland
rainforest in Brazil, Guyana, south through Bolivia, Paraguay and Argentina.
Roger (1861) described Dinoponera based on Ponera gigantea (Perty 1833) and
the synonym Ponera grandis (Guérin-Méneville 1838). Several authors described ad-
ditional species (Santschi 1921, Borgmeier 1937, Kempf 1971), subspecies (Emery
1901) and varieties (Santschi 1921, Santschi 1928), resulting in several tetranomials
and taxonomic confusion. Kempf (1971) provided a revision of the genus and with
additional corrections (Kempf 1975), that formed the basis for the current taxonomy
of the genus (Table 1).
Species limits among Dinoponera are dicult to dene (Kempf 1971). Further-
more, revisionary studies have been hampered by availability of specimens and col-
lection bias. Previous works (Perty 1833, Guérin 1838, Roger 1861, Emery 1901,
Santschi 1921, Santschi 1928, Borgmeier 1937) were based on limited numbers of
specimens. Most specimens have been collected only at the fringes of the Amazon
basin, or along major rivers (Fig. 13). Currently, six species are recognized in the
genus Dinoponera (Table 1), as well as two questionable subspecies of Dinoponera
australis based on male coloration (Borgmeier 1937). Kempf (1971) had doubts
as to the status of several species largely because uncollected areas suggested that
geographic variation was poorly understood. Character integration between what
appeared to be distinct species was a possibility, based on the lack of collections in
key areas. In particular, Kempf (1971) recognized that because of gaps in distribu-
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 121
Table 1. Current taxonomy of Dinoponera as suggested by Kempf (1971, 1975) and modied by this
publication, including known castes of each taxa. 1 is taxa or caste was described by this publication.
Female is used instead of the caste designation of gyne and worker because they are morphologically
indistinguishable. 2Validity of these taxa is questionable and could not be addressed in this study as types
were unavailable.
Taxa Described castes
Dinoponera australis Emery, 1901 Male/Female
D. a. bucki Borgmeier, 1937 2Male/Female
D. a. nigricolor Borgmeier, 1937 2Male/Female
Dinoponera gigantea (Perty, 1833) Male/Female
Dinoponera hispida sp. n.1Female1
Dinoponera longipes Emery, 1901 Male1/Female
Dinoponera lucida Emery, 1901 Female
Dinoponera mutica Emery, 1901 Female
Dinoponera quadriceps Santschi, 1921 Male/Female
Dinoponera snellingi sp. n.1Male1
tional data, character integration was a possibility in the uncollected areas between
D. longipes, D. mutica, D. quadriceps and D. gigantea where sympatry could exist. If
sympatry is demonstrated between dierent Dinoponera species and character inte-
gration is not found, this lends support to the recognition of taxa as distinct species.
Kempf (1971) suggested that a study incorporating more specimens and quantitative
characters could be more eective at dening species limits. Current analyses such
as DNA bar-coding or other molecular investigation could solve this dilemma. is
study seeks to determine whether the current taxonomy of Dinoponera reects the
actual species richness. We re-evaluate the alpha taxonomy of Dinoponera using mor-
phological characters (including male genitalia), and interpretation of distribution
and sympatry for workers and males.
Specimens and methods
We examined 345 workers and 11 males of Dinoponera, including type specimens for
D. australis, D. longipes, D. lucida, D. mutica and D. quadriceps. We were unable to ac-
quire the type specimens of Dinoponera gigantea, D. australis nigricolor and D. australis
bucki. In the material examined section for each species, female workers including pos-
sible gamergates are designated with a “w”, and male specimens with a “m” following
the number of specimens for each. Collections cited in this study are abbreviated as
follows:
AMNH American Museum of Natural History
CASC California Academy of Sciences Entomological Collection
CUIC Cornell University Insect Collection
FMNH Field Museum of Natural History
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
122
FSCA Florida State Collection of Arthropods
LACM Los Angeles County Museum of Natural History
UNAMB Museo Entomológico Facultad de Agronomía, Universidad Nacional de
Colombia
MCSN Museo Civico di Storia Naturale ‘Giacomo Doria’
MZSP Museo de Zoologia da Universidade de São Paulo
MCZC Museum of Comparative Zoology, Harvard University
NHMB Naturhistorisches Museum Basel
PALC Paul Alvarado Lenhart personal collection
QCAZ Pontica Universidad Católica del Ecuador, Catholic Zoology Museum
USNM Smithsonian Institution’s National Museum of Natural History
CWEM William and Emma Mackay Collection at the University of Texas at El Paso
Morphological characters were selected after extensive examination of the material.
Previous authors (Roger 1861, Emery 1901, Santschi 1921, 1928, Borgmeier 1937,
Kempf 1971) had identied several characters that in combination have been used to
distinguish workers. ese characters include gular striations, tooth-like antero-ventral
pronotal process, pilosity type and color, microsculpturing, petiole prole and stridula-
tory le shape. e known males possess more discernable character states including
compound eye shape, ocelli placement, pygidial spine, volsella and aedeagal shape.
Our own examination provided congruence for these characters.
Measurements were made with an ocular micrometer using a Wild stereomicro-
scope at 64×. A subset of 91 worker specimens including available types were measured
totaling 21 D. australis, 16 D. longipes, 15 D. gigantea, 17 D. quadriceps, 5 D. lucida,
12 D. mutica and 5 D. hispida. Specimen numbers of D. lucida, D. mutica and D. his-
pida were limited by availability in museum holdings. In descriptions measurements
include the range and in parenthesis the mean. Observations on the D. longipes males
were made using two specimens and therefore no mean is provided.
A measure of gaster length is not as consistent as other measurements because of
dierential expansion or contraction of the gastral segments in individual specimens,
resulting in a larger range of variation. However, gaster length is useful for a measure
of the approximate overall body length which is helpful when identifying D. australis
which diers notably in size from other species.
Photomicrographs (Fig. 12) were taken using a Carbeco ZDM1 Digital Video
Microscope.
Standard myrmecological morphometric parameters were generally selected to best
characterize the observed dierences and included:
MDL Mandibular length. e straight-line length of the mandible from the man-
dibular apex to the articulation with the clypeus.
SL Scape length. Maximum length of rst antennal segment not including the
articular boss and condyle.
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 123
FL1 First funicular segment length. Maximum length of second antennal segment,
for males only.
FL2 Second funicular segment length. Maximum length of third antennal segment,
for males only.
HL Head length. Midline measured from the distal edge of the clypeus at the
median (not including clypeal teeth) to the occipital margin of the head at its
median (not including ocelli in males).
HW Head width. Maximum width of the head in full-face view including eyes.
EL Eye length. Maximum length of the eye measured along its longer vertical axis.
EW Eye width. Maximum width of the eye measured along its shorter horizontal
axis.
OD Ocelli diameter. Maximum diameter of the medial ocellus, for males only.
WL Weber’s length. Distance measured between the anterior margin of the prono-
tum to the posterior margin of the metapleural bulla in lateral view.
FWL Forewing length. Maximum length of the forewing measured from the base of
the costal vein to the wing apex, for males only.
HWL Hindwing length. Maximum length of the hindwing measured from the base
of the costal vein to the wing apex, for males only.
PL Petiole length. e maximum measurable longitudinal distance between the
anterior and posterior extensions of the petiolar node in lateral view.
PH Petiole height. Height of the petiole measured laterally from the median of the
subpetiolar process viewed laterally to the median of the dorsum of the petiole.
PW Petiole width. Maximum width of the petiole measured in dorsal view.
GL Gaster length. Maximum longitudinal distance from articulation with petiolar
helcium to distal edge of hypopygidium (subgenital plate in males) measured
in lateral view.
HFL Hind femur length. Maximum length of the posterior femur measured from
its basal articulation with the trochanter to its apex at the articulation with the
tibia.
TBL Total body length (sum of MDL, HL, WL, PL, GL).
Label data were used from all specimens to plot distributions. ese data were
combined with localities derived from literature (Kempf 1971, Kempf 1975, Araujo et
al. 1990, Peeters et al. 1999, Monnin and Peeters 1999, Fourcassié and Oliviera 2002,
Monnin et al. 2003, Mariano et al. 2004, Araújo and Rodriques 2006, Marques-Silva
et al. 2006) to supplement specimen distribution data. GPS coordinates were recorded
from labels, or estimated from Google Earth (http://earth.google.com/) or the Global
Gazetteer Version 2.1 Directory of cities and towns in the world (http://www.fall-
ingrain.com/world/index.html). Maps were constructed using the software package
ArcGIS version 9.2 (ESRI 2007). A complete table of localities including estimated
geographic coordinates is provided in supplementary le 1.
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
124
Results
Genus Dinoponera Roger, 1861
http://species-id.net/wiki/Dinoponera
Family Formicidae, subfamily Ponerinae, tribe Ponerini. Described by Perty (1833)
as Ponera gigantea. Dened as a genus by Roger (1861) (Type species: Dinoponera
gigantea)
Diagnosis. Size (TBL > 2.5cm) can easily distinguish Dinoponera from other worker
ants. Two laterally projecting clypeal teeth (Fig. 1A) and rows of spines on the py-
gidium and hypopygidium will further distinguish this genus. e gamergates of Dino-
ponera are not distinct from workers in their external morphology (Haskins and Zahl
1971, Araujo et al. 1990, Paiva and Brandão 1995, Monnin and Peeters 1998). True
gynes have not been found in this genus.
Description of the worker. Abundant setae; black integument, ranges from
smooth and shiny with no microsculpturing, to nely micropunctate or scaled de-
pending on species (Fig. 12). Head: Mandibles long and curved posteriorly in side
view; seven large teeth; erect setae on dorsum. Ventral surface of head with sparse de-
cumbent and subdecumbent setae; may have ne striations depending on species; Pa-
pal formula 4, 4; large bilobed labrum. Clypeus with two laterally projecting teeth on
anterior edge, clypeus bulging medially, extending posteriorly between frontal lobes,
anterior edge with row of long setae; sparse appressed setae from distal edges to medial
area of clypeus. Area posterior to clypeus with varying amounts of striation. Tento-
rial pits apparent. Frontal lobes raised and conspicuous, with striations at posterior
constriction. Antennae: geniculate, 12 segments, all with agellate setae; scape long,
extending past posterior border of head; funiculus covered in minute appressed pu-
bescence. Gena depressed medially of eye; dense appressed setae on the antero-lateral
sides of the head; covered in conected punctulate sculpturing. Eyes large, elliptical
with slight depression (ocular ring) around circumference. Frons with large pads of
long agellate pubescence (lost in older or poorly curated specimens). Median fur-
row running from posterior termination of clypeus, between frontal lobes to center of
frons, terminates in shallow pit in most specimens. Entire head covered in long agel-
late subdecumbent setae (Fig. 1A). Mesosoma: in lateral view weakly convex; covered
in long subdecumbent to erect agellate pilosity and dense pubescence; pronotal disc
with slight bulges; promesonotal suture distinct, suture between mesopleuron and pro-
podeum distinct; mesonotum fused with propodeum and episternum, separated by
slight furrows; basilar sclerite large, ovaloid; propodeum with broadly rounded dorsal
outline, dorsal surface gradually curves into posterior face (Fig. 2); propodeal spiracle
forms nearly vertical slit; sulcus running from center of propodeum along lower edge
of propodeal spiracle to posterior edge of propodeum at dorsal edge of bulla, patches
of short white pubescence at curved posterior border of pronotum and basilar sclerite.
Legs long, covered in long setae with short, sti pubescence. One well-developed,
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 125
antennae cleaning, comb-like spur on foreleg; one spine-like appendage and one less
developed denticular comb on mesothoracic tibia; one spine and one comb-like spur
on hind tibia. Posterior side of fore leg basitarsus with dense pads of golden setae; tar-
sal claws bidentate. Petiole: node large and tabular in lateral view, narrow attachments
at base to propodeum and gaster; in dorsal view largest width less than propodeum
and gaster, varies from ovate rectangular to ovate triangular in outline; covered in
long subdecumbent to erect agellate pilosity; pubescence on anterior face and ridges
of subpetiolar process; subpetiolar process reduced, slightly variable between species.
Gaster: typical of ponerines; covered with agellate setae with short pubescence; small
protuberance at articulation of gastric sternite III and the petiole; stridulatory le of
Figure 1. Features of Dinoponera workers. A Head, frontal view B–C Occiput of head, oblique antero-
lateral view D–E Pronotum, lateral view F–H Petiole, lateral view. A–B D. longipes C D. hispida D D.
gigantea. E–F D. mutica G D. hispida H D. lucida.
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
126
varying size on acrotergite of gastral tergum II; posterior edges of the pygidium and
hypopygidium with characteristic rows of minute spines.
Description of the male. Integument: smooth and nitid; reddish to dark brown/
black. Head: Mandibles greatly reduced, rounded, spoon shaped, lacking teeth; palps
elongated, maxillary palps 4 segmented, labial palps 3 segmented; labrum reduced,
rounded to truncate, emarginated distal margin in D. snellingi and D. longipes covered
with setae. Clypeus large, triangular, bulging medially; anterior tentorial pits large;
frontal lobes absent; antennal sockets almost touching, located at posterior apex of
clypeus. Antennae: geniculate, 13–segmented, pilosity varies from ne pubescence to
long setae in dierent species; scape shorter than second funicular segment, but shorter
than 1st, 1st funicular segment reduced. Compound eyes large, along lateral side of
head, deeply emarginated medially. ree ocelli at posterior margin of head, bulging
beyond margin of head in all species except D. australis. Entire head immaculate, cov-
ered in ne pubescence and long erect setae (Fig. 3). Mesosoma: pronotum triangular,
exposed narrowly dorsally anterior to scutum; scutum large, bulging antero-dorsally,
Figure 2. Dinoponera hispida worker. Head in full frontal view; body in lateral view.
Figure 3. Dinoponera longipes male. Head in full frontal view; body in lateral view with wings not shown.
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 127
with 3 longitudinal carina; small tegula over insertion of forewing; scutellum domed,
side with vertical carina, dorsal surface smooth; basilar sclerite under hind wing re-
duced; fused mesopleuron, separated by furrow into anepisternum and katepisternum;
metanotum exposed between scutellum and propodeum, reduced; dorsal face of pro-
podeum shorter than posterior face, rounded into posterior face; coxa large, conical
(Fig. 3). Wings: covered in minute pubescence, venation as shown in Figure 5. Legs:
one well-developed, antennae cleaning, pectinate spur on foreleg; one spine-like and
one less developed denticular comb on mesothoracic tibia; one spine and one comb-
like spur on hind tibia. Posterior side of fore basitarsus with dense pads of golden setae;
tarsal claws bidentate. Petiole: narrow attachments at base to propodeum and gaster;
petiolar node humped dorsally, subpetiolar process anteriorly triangular. Gaster: large,
cylindrical, covered in ne silvery pubescence; pygidium terminating in spine poste-
riorly, with short cerci; hypopygidium with long ne erect setae, tabular subgenital
plate with posterior end truncated, often emarginated. Genitalia (Figs 6–11): basal
ring with dorso-anterior loop structures; parameres long, rounded, with emarginated
ventro-basal edge (Fig. 9); volsella articulated with basiparamere along ventral edge,
lateral nger-like cuspis volsellaris, medial digitus volsellaris with distal wide toothed
cusp, basal medial lobe with tooth-like structures varying with species (Fig. 10); penis
valve of aedeagus roughly triangular and rounded, aedeagal apodeme curved horn-like
antero-lateral arm structure arising from mid-valve ridge, terminating at interior sur-
face of basiparamere (Fig. 11).
Description of the larvae. A basic description of the larva of D. quadriceps (cited
as D. grandis mutica) is present in Mann (1916). A detailed description of the egg and
all larval stages of Dinoponera gigantea are present in Wheeler and Wheeler (1985). e
following generic description of Dinoponera larvae is from their work:
"Prole pogonomyrmecoid (i.e., diameter greatest near the middle of abdomen, de-
creasing gradually toward anterior end and more rapidly toward posterior end, which is
rounded; thorax more slender than abdomen and forming a neck, which is curved ven-
trally). Body with numerous (114–160) mammiform tubercles, each with 2–25 short
simple hairs; body hairs lacking elsewhere. Cranial hairs lacking. Mandible dinoponeroid
(i.e. narrowly subtriangular in anterior view; anterior portion curved posteriorly; with or
without medial teeth.)"
Discussion. Dinoponera’s status as a genus is validated as several characters dif-
ferentiate it from other genera. Size (TBL>2.3cm) is the most obvious character dis-
tinguishing Dinoponera. e only other ants with a worker caste approaching this size
are Paraponera clavata (Fabricius) and the larger Pachycondyla such as P. crassinoda (La-
treille 1802), P. impressa Roger 1861 and P. villosa (Fabricius 1804). Paraponera clavata
is easily separated by its anvil shaped petiole with a spine on the ventral surface, highly
sculptured body and deep antennal scrobes. Pachycondyla is regarded as the sister taxa
to Dinoponera (Kempf 1971, Schmidt 2010). Dinoponera, in addition to their size, are
distinguishable from Pachycondyla by the presence of two laterally projecting clypeal
teeth (Fig. 1A) and rows of spines on the pygidium and hypopygidium. Several (n=6)
specimens have been observed to have a single ocelli in the pit at the termination of
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
128
the median furrow. ese anomalous specimens were previously thought to be queens
(Borgmeier 1937) but as it has been shown that Dinoponera lacks queens, the presence
of the ocelli is hypothesized to be the result of a Mermis Dujardin 1842 nematode
parasite (Kempf 1971).
Dinoponera biology. Dinoponera is one of the roughly 10 ponerine genera in
which some species have secondarily lost the typical morphologically specialized queen
caste for a reproductive worker known as a gamergate (Haskins and Zahl 1971, Araujo
et al. 1990, Paiva and Brandão 1995, Monnin and Peeters 1998, Peixoto et al. 2008).
Conict over dominance is intense in colonies with younger workers usually joining a
linear hierarchy of one to ve workers depending on colony size. e gamergate, or al-
pha female has the highest ranking (Monnin and Ratnieks 1999, Monnin et al. 2003).
e alpha female mates with non-nestmate males at night at the entrance of the nest
(Monnin and Peeters 1998, Monnin and Peeters 1999). After copulation the female
bites through the male’s gaster to release herself and pulls out the genital capsule which
acts as a temporary sperm plug (Monnin and Peeters 1998). After mating the female is
unreceptive to other males and remains monandrous (Monnin and Peeters 1998). e
gamergate maintains dominance with ritualized behaviors such as antennal boxing and
biting, ‘blocking’, as well as gaster rubbing and curling (Monnin and Peeters 1999).
Lipid stores within Dinoponera australis females were found to be strongly related to
foraging activity and reproductive status within the colony, ranging from 1–39% of an
individual’s dry mass (Smith et al. 2011). It is uncertain, however, whether nutritional
dierences between females is a cause or consequence of rank. Gamergate females
possess a higher concentration of a cuticular hydrocarbon (9-hentriacontene, 9-C31:1)
that indicates rank and is passed onto gamergate-laid egg cuticles (Monnin and Peeters
1997, Monnin et al. 1998, Peeters et al. 1999). Additionally, alpha females may ‘sting
smear’ a competing female with secretions from the Dufour’s gland, triggering the
lower ranking workers to immobilize the marked female (Monnin and Ratnieks 2001).
Subordinate females (beta, gamma, or delta) may produce unfertilized eggs but these
are usually consumed by the alpha female in a form of “queen policing” (Monnin and
Peeters 1997). Egg recognition in D. quadriceps was found to be due to dierences
in cuticular hydrocarbons, and only workers engaged in brood care could distinguish
non-nestmate eggs (Tannure-Nascimento et al. 2009). Cuticular hydrocarbons are also
used to distinguish adult nestmates from non-nestmates, however, this is only eec-
tive with non-nestmate foragers (Nascimento et al. 2012). Nascimento et al. (2012)
found that brood-caring workers from dierent colonies had very similar hydrocarbon
proles and were more often accepted into alien colonies.
Males are born throughout most of the year in tropical species (Araujo and Jais-
son 1994, Monnin and Peeters 1998), however D. australis which lives in the more
temperate south was found to only produce males in May-July (Paiva and Brandão
1995). When the alpha declines reproductively or dies, she is replaced by a high-rank-
ing worker (Monnin and Peeters 1999).
New colonies are founded by ssion, a process in which a beta female is fertilized
in the natal nest (Monnin and Peeters 1998). is new alpha female then leaves the
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 129
nest with a cohort of workers to found an incipient colony, sometimes employing tan-
dem running (Overal 1980).
Colonies vary in size depending upon species. Dinoponera australis colonies have
an average of 14 workers (range 3–37) (Paiva and Brandão 1995, Monnin et al. 2003),
D. gigantea average 41 workers (range ~30–96) (Overal 1980, Fourcassié and Oliviera
2002, Monnin et al. 2003) and D. quadriceps has the largest colonies with an average of
80 workers (range 26–238) (Monnin and Peeters 1999, Monnin and Ratnieks 2001).
Morgan (1993) excavated two D. longipes nests, a possible incipient colony with 7
workers and another mature colony of 120 workers.
e nest consists of large chambers and tunnels in the soil possibly with an earthen
mound and can be 0.10–1.2m deep (Araujo et al. 1990, Morgan 1993, Fourcassié and
Oliviera 2002, Vasconcellos et al. 2004). Nests are deeper in D. australis and D. quadri-
ceps than in D. gigantea, Monnin et al. (2003) suggests that deeper nests are a possible
adaptation to seasons and aridity. Dinoponera gigantea nests may have up to eight en-
trances and can be weakly polydomous (Fourcassié and Oliviera 2002), whereas 1–30
openings with an average of 11 were recorded for D. longipes (Morgan 1993). Nesting
density and spatial distribution varies depending on habitat (Fowler 1985, Vasconcel-
los et al. 2004). Density ranges from 15–40 nests per ha-1 (Vasconcellos et al. 2004)
to 80 nests per ha-1 (Paiva and Brandão 1995). Morgan (1993) measured a spacing
between nests for D. longipes with a median of 35m (n=22, range 14–69.5m). Dinopo-
nera australis and D. gigantea usually nest at the base of trees (Paiva and Brandão 1995,
Fourcassié and Oliviera 2002). Observations of D. quadriceps nests show that in more
arid Caatinga and Cerrado habitats, nests are predominantly constructed under trees,
whereas in Atlantic forest 60% of nests were 3m away from any tree (Vasconcellos et
al. 2004).
Workers lower in the hierarchy forage individually for food items on the sub-
strate and do not recruit other nestmates to assist with food transport (Fowler 1985,
Fourcassié et al. 1999, Fourcassié and Oliviera 2002, Araújo and Rodrigues 2006).
Although foraging workers do not recruit nestmates, Nascimento et al. (2012) found
a positive feedback between incoming food and stimulation of new foragers as well
as task partitioning once food was brought into the nest. Lower ranking females pro-
cessed protein resources while higher ranking females handled small food pieces and
distributed them to the larvae. Fourcassié and Oliviera (2002) found D. gigantea for-
aging to be concentrated in the early morning and afternoon but did not sample at
night. Morgan (1993) observed the highest activity at night in D. longipes. Dinoponera
quadriceps has a marked seasonal pattern in activity. It is most active in May-August
, the late rainy season to early dry season in the semiarid Caatinga (Medeiros et al.
2012). Activity was strongly negatively correlated to temperature and positively cor-
related to prey abundance (Medeiros et al. 2012). e diets of both D. gigantea and
D. quadriceps have been shown to be predominantly scavenged invertebrates, but in-
clude live prey, seeds and fruits (Zahl 1959, Fourcassié and Oliviera 2002, Araújo and
Rodrigues 2006). Araújo and Rodrigues (2006) state that the taxonomic diversity of
prey is comparable to other tropical ponerines, but has an optimal prey size of 2–3
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
130
cm in Dinoponera. Diet seems to be very similar across the genus, regardless of habitat
(Araújo and Rodrigues 2006).
Despite their large size and strong venom, Dinoponera are likely preyed on by
many vertebrate and invertebrate species across South America. Like many other ant
species, Dinoponera can be infected by the entomopathogenic fungi Codyceps sp. (Ev-
ans 1982). Buys et al. (2010) discovered a Kapala sp. eucharitid wasp emerging from
the puparia of Dinoponera lucida.
Anatomy has been described several times. Marques-Silva et al. (2006) studied of
the sensilla and glands of the antennae. Anatomy of the venom apparatus and mandib-
ular glands of Dinoponera gigantea is presented in Hermann et al. (1984). Further stud-
ies of the mandibular glands and its contents were presented by Oldham and Morgan
(1993) and Oldham et al. (1994). Oldham et al. (1994) found that the mandibular
gland secretions of workers diered greatly from those of gamergates, which were 98%
dimethylalkylpyrazine and lacked the four other pyrazines and 50 times more volatiles
than those found in worker secretions. e post-pharyngeal gland morphology was ex-
amined by Schoeters and Billen (1997). e cuticular hydrocarbons used in nestmate
recognition may be produced by epidermal glands which Serrão et al. (2009) found in
the epidermis of abdominal sternites in D. lucida.
For subduing large live prey and defense (Morgan, 1993), workers possess a
sting that has been known to cause severe pain lasting up to 48 hours, lymphaede-
nopathy, edema, tachycardia and fresh blood to appear in human victim feces are
common symptoms (Haddad et al. 2005). In gamergates the venom sac is empty
(Monnin et al. 2002). Workers may have 60–75 unique proteinaceous components
in the venom (Morgan et al. 2003, Johnson et al. 2010). e convoluted gland
within the venom system of Dinoponera australis has been found to possess close
similarities to those of vespine wasps (Schoeters and Billen 1995). e contents
of D. australis venom have been found to be similar to those of Pachycondyla spp.
(Cruz López 1994, Johnson et al. 2010). Billen et al. (1995) studied the morphol-
ogy and ultrastructure of the pygidial gland of D. australis. Due to the high diversity
of compounds and systemic eects found by Haddad et al. (2005), venom of Di-
noponera could be of use to the pharmaceutical industry. For instance, Sousa et al.
(2012) demonstrated in mice that venom from D. quadriceps had antinociceptive
properties. e authors note that the local population of northeast Brazil uses dry
crushed D. quadriceps ants to treat earaches, and the stings of live ants are adminis-
tered for back pain and rheumatism.
Several studies of the cytogenetics of Dinoponera species have been conducted. Di-
noponera lucida may have the highest number of chromosomes within the Hymenop-
tera however the karyotype is variable between populations (2n=106–120) (Mariano
et al. 2004, Mariano et al. 2008, Barros et al. 2009). Mariano et al. (2008) interpreted
the karyotype dierences between populations as being due to a division of the species
into allopatric populations during the Quaternary. Variability in the karyotype within
a described species has been found in the Pachycondyla as well, and may represent cryp-
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 131
tic species (Mariano et al. 2012). Descriptions of the banding patterns on Dinoponera
chromosomes are provided by Barros et al. (2009) and de Aguiar et al. (2011).
Dinoponera belongs to the tribe Ponerini in the subfamily Ponerinae. e evo-
lutionary position of the genus within Ponerinae was resolved by Schmidt (2010).
Based on the phylogenetic analysis of Schmidt (2010) and karyotype analysis by Mari-
ano et al. (2012), Dinoponera’s closest living relatives are in the Pachycondyla species
group consisting of P. crassinoda, P. harpax (Fabricius 1804), P. impressa, P. metanotalis
Luederwaldt 1918, and P. striata Smith 1858. Prior to the generation of well support-
ed phylogenies other associations had been proposed. Carpenter (1930) suggested that
the fossil Archiponera wheeleri Carpenter from the Miocene Florissant shale of Colo-
rado may be an ancestor of Dinoponera and Streblognathus aethiopicus Smith 1858.
Molecular data has shown that Carpenter’s (1930) hypothesis is false (Schmidt 2010).
Streblognathus is not closely related to Dinoponera, and its morphological similarity is
purely convergence. e placement of Archiponera wheeleri is still questionable.
Key to the workers of Dinoponera
1 Antero-inferior corner of pronotum with distinct tooth-like process
(Fig.1D).....................................................................................................2
– Antero-inferior corner of pronotum without tooth-like process (Fig. 1E) ....4
2 Head (Fig. 12B), sides of petiole and terga 1 and 2 of gaster smooth and pol-
ished, integument with bluish luster (Fig. 12A); southeastern coast of Brazil ..
..................................................................................................lucida Emery
– Head, lateral sides of the petiole and terga 1 and 2 of gaster nely micro-
punctate/scaled (Fig. 12B) ..........................................................................3
3 Total body length under 3 cm; Bolivia, Paraguay, northern Argentina and
southern Brazil .................................................................... australis Emery
– Total body length over 3 cm; Brazil, Peru, and Guyana ....... gigantea (Perty)
4 Body covered in bristle-like setae which are not agellate (Fig. 1C); Pará,
Brazil ...................................................................................... hispida sp. n.
– Body covered in ne and agellate setae (Fig. 1B) .......................................5
5 Appressed golden yellow pubescence present on frons (Fig. 1A); Colombia
south to Perú to northern Brazil ........................................... longipes Emery
– Pubescence absent from frons or not golden yellow ....................................6
6 Sides of head, lateral sides of the petiole and terga 1 and 2 of gaster nely
micro-punctate/scaled (Fig. 12B); in prole antero-dorsal edge of petiole
bulging (Fig. 1G); northeast Brazil ................................quadriceps Santschi
– Sides of head, sides of the petiole and terga 1 and 2 of gaster smooth, shiny and
polished with no microsculpturing, integument with bluish luster (Fig.12A);
in prole petiole with even dorsal corners (Fig. 1F); eastern Bolivia, northern
Paraguay and southwestern Brazil ..........................................mutica Emery
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
132
Key to the known males of Dinoponera
(couplets 1 and 2 are included to easily separate males of other genera which are likely
confused with Dinoponera)
1 Total body length less than 15 mm ......... Pachycondyla and other poneroids
– Total body length greater than 15 mm ........................................................ 2
2 Subpetiolar process in form of spine; pronotum heavily sculptured; palp for-
mula 5:3 .................................................................................... Paraponera
– Subpetiolar process without spine; pronotum shiny, microsculptured; palp
formula 4:3 ................................................................................................. 3
3 Ocelli protruding on occipital margin of head (Fig. 4A–D) ........................4
– Ocelli not protruding on occipital margin of head (Fig. 4E); Bolivia, Para-
guay, northern Argentina and southern Brazil ..................... australis Emery
4 Setae on funiculus long and erect (Fig. 4F, G) ............................................. 5
– Setae on funiculus short, sti and subdecumbent, or minute pubescence pre-
sent (Fig. 4H, I, J) ......................................................................................6
5 Digitus volsellaris with toothless lobe at distal end (Fig. 10C); Brazil, Peru,
and Guyana ........................................................................ gigantea (Perty)
– Digitus volsellaris without lobe at distal end (Fig. 10A); northeast Brazil ......
.....................................................................................quadriceps Santschi
6 Pygidium terminating in narrow, elongate spine (Fig. 4M); penis valve of
aedeagus wedge-shaped in lateral view (Fig. 11E); Colombia south to Perú to
north western Brazil ............................................................. longipes Emery
– Pygidium terminating in short, broad, triangular angle (Fig. 4N); penis valve
of aedeagus with distal ange and triangular ventral lobe (Fig. 11B); Mato
Grosso do Sul, Brazil ..............................................................snellingi sp. n.
Clave para la identicación de las obreras de Dinoponera
1 Esquina antero-inferior del pronoto con proceso en forma de diente distinto
(Fig.1D).....................................................................................................2
– Esquina antero-inferior del pronoto sin proceso en forma de diente
(Fig.1E) .....................................................................................................4
2 Cabeza, lados laterales del pecíolo, y tergos 1 y 2 del gáster lisos y brillantes,
con reexiones azules (Fig. 12A) ............................................. lucida Emery
– Cabeza, lados laterales del pecíolo, y las tergos 1 y 2 del gáster namente
punteados o con escamas nas (Fig. 12B), a veces con reexiones azules .....3
3 Largo total menos de 3 cms ................................................ australis Emery
– Largo total más de 3cms .....................................................gigantea (Perty)
4 Cuerpo cubierto con pelos gruesos, no agelados (Fig. 1C)..... hispida sp. n.
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 133
– Cuerpo cubierto con pelos nos y agelados (Fig. 1B) ................................5
5 Frente con pubescencia recostada y amarillo-dorada (Fig. 1A) .......................
............................................................................................ longipes Emery
– Frente sin pubescencia recostada y amarilla .................................................6
6 Lados de la cabeza, lados del pecíolo y tergos 1 y 2 del gáster namente punt-
eados o con escamas nas (Fig. 12B); pecíolo (en perl) con el borde anterior-
dorsal hinchado (Fig. 1G) .............................................quadriceps Santschi
– Lados de la cabeza, lados del pecíolo y tergos 1 y 2 del gáster lisos, y brillantes,
sin escultura na, con reexiones azules (Fig. 12A); pecíolo (en perl) con
esquinas dorsales al mismo nivel (Fig. 1F) .............................. mutica Emery
Clave para la identicación de los machos conocidos de Dinoponera
1 Largo total menos de 15 mm .....................Pachycondyla y otros poneroides
– Largo total más de 15 mm .......................................................................... 2
2 Proceso subpetiolar en forma de espina; pronoto fuertemente esculturado;
formula palpal 5:3......................................................................Paraponera
– Proceso subpetiolar no en forma de espina; pronoto liso, microesculturado;
formula palpal 4:3.......................................................................................3
3 Ocelos muy hinchados en el margen occipital de la cabeza (Fig. 4A–D) ......4
– Ocelos no muy hinchados en el margen occipital de la cabeza (Fig. 4E) ........
........................................................................................... australis Emery
4 Funículo con pelos alargados y rectos (Fig. 4F, G) ....................................... 5
– Funículo con pelos cortos, rígidos, y subdecumbentes, o con pubescencia
diminuta (Fig. 4H, I, J) ..............................................................................6
5 Lóbulo del digito del volsela sin dientes en el ápice (Fig. 10C) ......................
........................................................................................... gigantea (Perty)
– Digito del volsela sin lóbulo en el ápice (Fig. 10A) ........quadriceps Santschi
6 Pigidio terminando en un espina, alargada y delgada (Fig. 4M); válvula peneal
del aedeago en forma de cuña (Visto en perl) (Fig. 11E) .....longipes Emery
– Pigidio terminando en una espina corta y ancha, en forma de triángulo
(Fig.4N); válvula peneal del aedeago con una reborde distal y un lóbulo tri-
angular ventral (Fig. 11B) ......................................................snellingi sp. n.
Chave para identicação de operários de Dinoponera
1 Esquina antero-inferior do pronoto com processo em forma de dente distinto
(Fig. 1D).....................................................................................................2
– Esquina antero-inferior do pronoto sem processo em forma de dente
(Fig.1E) .....................................................................................................4
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134
2 Cabeça, lados laterais do pecíolo, e tergas 1 e 2 do gáster lisos e brilhantes,
com reexões azuis (Fig. 12A) ................................................. lucida Emery
– Cabeza, lados laterais do pecíolo, e tergas 1 e 2 do gáster namente ponteado
ou com escamas nas (Fig. 12B) .................................................................3
3 Comprido total menos 3 cms .............................................. australis Emery
– Comprido total mais de 3 cms ............................................ gigantea (Perty)
4 Corpo cobrido com cogumelos em forma de cerdas, que não são agelados
(Fig. 1C) ................................................................................. hispida sp. n.
– Corpo cobrido com cogumelos nos e agelados (Fig. 1B) .........................5
5 Frons com pubescencia prendida e amarela-dourada (Fig. 1A) ......................
............................................................................................ longipes Emery
– Frons sem pubescencia amarelo prendido ....................................................6
6 Lados da cabeça, lados do pecíolo e tergas 1 e 2 do gáster namente pon-
teados ou com escamas nas (Fig. 12B); pecíolo (em perl) com o margem
anterior-dorsal hinchado (Fig. 1G)................................quadriceps Santschi
– Lados da cabeça, lados do pecíolo e tergas 1 e 2 do gáster lisos e brilhantes,
sem escultura na, com reexões azuis (Fig. 12A); pecíolo (em perl) com
esquinas dorsais ao mesmo nível (Fig. 1F) .............................. mutica Emery
Chave para identicação de machos de Dinoponera
1 Comprido total menos de 15mm ......... Pachycondyla e outro poneromorfos
– Comprido total mais de 15 mm .................................................................. 2
2 Proceso subpetiolar em forma de espinha; pronoto fortemente esculturado;
formula palpular 5:3 .................................................................. Paraponera
– Processo subpetiolar não em forma de espinha; pronoto liso, microescultura-
do; formula para palpular 4:3 ......................................................................3
3 Ocelos na margem occipital da cabeça muito inchados (Fig. 4A–D) ...........4
– Ocelos na margem occipital da cabeça não muito inchados (Fig. 4E) ............
........................................................................................... australis Emery
4 Funículo com cogumelos alongados e retos (Fig. 4F, G) .............................. 5
– Funículo com cogumelos curtos, rígidos, e subdecumbentes, ou com pubes-
cencia minuta (Fig. 4H, I, J) .......................................................................6
5 Lóbulo do dígito do volsela sem dentes no ápice (Fig. 10C) ...gigantea (Perty)
– Dígito do volsela sem lóbulo no ápice (Fig. 10A) ..........quadriceps Santschi
6 Pigidio acabado numa espinha, alongada e delgada (Fig. 4M); válvula penal do
aedeago em forma de cunha (visto em perl) (Fig. 11 E) ........longipes Emery
– Pigidio acabado numa espinha apara e alarga, em forma de triângulo (Fig.4N);
válvula penal do aedeago com um reborde distal e um lóbulo triangular ven-
tral (Fig. 11B) ........................................................................snellingi sp. n.
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 135
Dinoponera australis Emery
http://species-id.net/wiki/Dinoponera_australis
Figs 4E, J, O, 5D, 9D, 10D, 11D, 13
Dinoponera grandis subsp. australis Emery, 1901: 48, worker, BRAZIL: S. Paulo: Avan-
handava [5 syntypes workers examined, MCSN]; additional syntypes from PARA-
GUAY: Rio Apa, (leg.) Balzan, (leg.); ARGENTINA: Missiones, 1881, Berg (leg.),
Giabibiri, Misiones, marzo 1884 G. Bove (leg.); Santschi, 1921: 85 (male); raised
to species, Borgmeier, 1937: 227.
Dinoponera grandis subsp. australis var. brevis Santschi, 1928: 416 , PARAGUAY: Re-
ichensperger (leg.) [type worker examined, NHMB]. Unavailable name, junior
synonym of D. brevis: Kempf, 1971: 387.
Subspecies. Dinoponera australis bucki Borgmeier, 1937:228. BRAZIL: Rio Grande
do Sul: Palmeira [types not available].
Dinoponera australis nigricolor Borgmeier, 1937:228. BRAZIL: Goyaz: Campinas
[types not available].
Worker diagnosis. is species is most easily recognized by the antero-inferior
corner of pronotum having a distinct tooth-like process (Fig. 1D), the pilosity being
short and relatively sparse and the integument being nely micro-sculptured and dull
(Fig 12B). In addition the scape length is shorter than the head width and the total
body length is under 30 mm. Dinoponera lucida could be confused with D. australis
in that it also has a tooth-like process on the pronotum and can have a TBL under 30
mm, but diers in having the smooth and shiny integument, long agellate hairs on
lobe and forward slanting dorsal edge of petiole.
Description of the worker. Measurements (mm) (n=21) TBL: 23.42–29.31
(26.21); MDL: 3.59 – 4.31 (3.88); HL: 4.51–5.64 (4.99); HW: 4.31–5.74 (4.89);
SL: 4.31–5.02 (4.73); WL: 6.25–7.69 (7.12); PL: 1.79–2.26 (2.03); PH: 2.56–3.28
(2.90); PW: 1.59–1.95 (1.75); GL: 7.28–9.64 (8.20); HFL: 5.54–6.66 (6.16). A de-
scription of the external morphology of the worker is given by Kempf (1971):
"Antennal scape length equal to, or shorter than head width. Pubescence on front of
head short and inconspicuous. Gular face of head subopaque, nely reticulate-punctate
throughout; the ne, arcuate striae variably developed from completely covering the un-
dersurface of head to only vestigially shown antero-laterally or nearly absent. Sides of head
reticulate-punctate, subopaque. Antero-inferior corner of pronotum dentate. Pronotal disc
supercially reticulate and quite shining; paired swellings either feeble or distinct. Length of
hind tibia equal to or less than head length. Petiole, in dorsal view, subquadrate, width over
length proportion always more than 0.80, notably shorter and broader than in the other
species; its shape…resembling that of mutica, with the upper anterior and posterior corners
equally rounded; nely reticulate, somewhat shining; vertical sulcus on posterior surface
either absent of more rarely vestigial to feebly developed. Terga I and II of gaster either
reticulate-punctate or more supercially reticulate (in the southern range of the territory)
and accordingly either subopaque or somewhat shining: ne appressed pubescence lacking
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
136
completely on disc of the terga, present on the sides. Stridulatory le on acrotergite of tergum
II of gaster well developed, broad and triangular, extending back to the acrotergite for about
one half to two thirds of its length."
Male diagnosis. Dinoponera australis males are recognized by their rounded head,
with compound eyes, reduced ocelli and the posterior margin around the ocelli not
protruding as in other species (Fig. 4E). is species is also characterized by the short,
broad pygidial spine (Fig. 4O), volsella with tear-drop shaped basal lobe covered in
minute teeth (Fig. 10D) and aedeagus with a latero-apical fold, notches and teeth
along ventral edge as shown in Fig. 11D.
Description of the male. A description of the external morphology of the male is
given in Kempf (1971):
"Head…with smaller eyes, the maximum interocular width being greater than their
diameter; with smaller ocelli not protruding above the posterior border of head when seen in
full-face view; antennal scape very short, less than twice as long as broad; funiculi without
standing hairs; petiole distinctly shorter although variable in outline…; pygidium with a
very short spine, not projecting beyond the long cerci; hypopygidium apically broadly trun-
cate, the truncation either straight, or convex, or concave."
Figure 4. Features of Dinoponera males. A–E Head, frontal view F–J Right scape, rst and second fu-
nicular segments, frontal view K–O Pygidial spine, dorsal view A, F, K D. gigantea B , G , L D. quadriceps
C , H , M D. longipes D, I, N D. snellingi E , J , O D. australis.
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 137
Description of the male genitalia. Basal ring with thick dorso-anterior loop
structures, reduced; parameres short, broad, rounded, small lobe on dorsal edge,
emarginated ventro-basal edge (Fig. 9D); cuspis volsellaris with few bumps or teeth,
digitus volsellaris with numerous small circular bumps at distal lateral face, tuft of
setae on ventro-distal side of broad cusp, large tear-shaped lobe on basal ventral
corner, covered in minute teeth (Fig. 10D); penis valve of aedeagus with lateral arm
of apodeme at anterior border, no ventral concavity under ridge at base of apodeme,
dorsal edge rounded, sloping posteriorly, ventro-anterior triangular projection fol-
lowed by circular notch, ventral projecting tooth, smaller hemispherical notch with
sclerotized border, thin, nely serrated distal edge, noticeable lateral apical fold with
slight serration ending ventrally in serrated ridge, rounded un-serrated lobe at distal
apex of valve.
Figure 5. Wings of known males. A D. quadriceps B D. snellingi C D. gigantea D D. australis E D.
longipes.
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
138
Distribution. Dinoponera australis has the widest known range of the Dinoponera.
is species is found in the department of Santa Cruz in Bolivia, southern Brazil in the
states of Mato Grosso, Goias, Minas Gerais, São Paulo, Mato Grosso do Sul, Paraná,
Santa Catarina and Rio Grande do Sul, eastern Paraguay in the departments of Itapúa,
Alto Paraná and Guairá, as well as the province of Misiones in Argentina (Fig. 13).
Discussion. Dinoponera australis is the most aberrant of the Dinoponera species
because of its relatively small size, sparse non-agellate pubescence, as well as the male
characters stated above which distinguish this species. e male coloration dierence
is the basis for the designation of the subspecies D. australis bucki and D. a. nigricolor.
ese may be separate species or the product of intra-specic variation, but this cannot
be diagnosed here as the types designated by Borgmeier (1937) were not available to
us and the extent of intra-specic variation could not be determined from the limited
sample size available.
Material examined. ARGENTINA, MISIONES: Iguazú (1 w, 4–10.x.1927,
RC and EM Shannon, USNM); Iguazú Falls (2 w, 20–22.i.1920, CUIC); Loreto (1
w, N Kusnezov, USNM); Loreto malaise trap in subtropical wet forest (3 w, i.2001,
P Fidalgo, FSCA); Parque Nacional Iguazú (3 w, 24.xii.1988, DH and AC Kistner,
LACM); Parque Nacional Iguazú Cantera old gravel pit at forest edge 200m (18 w,
8.xii.1990–6.i.1991, S and J Peck, FMNH); Parque Nacional Iguazú Puerto Canoas
hill forest 200m (30 w, 8.xii.1990–6.i.1991, S and J Peck, FMNH); Parque Nacional
Iguazú Puerto Canoas river forest 180m (35 w, 2 m, 8.xii.1990–6.i.1991, S and J Peck,
FMNH); Puerto Iguazú 100m (1 w, 25.xi–8.xii.1983, A Bordón, MCZC); Santo Pipó
(1 w, N. Kusnezov, MCZC); locality not specied (1 w, USNM). BOLIVIA, SANTA
CRUZ: Ayacucho (1 w, 13.x.1987, P Bettella, LACM); Buena Vista (1 w, 8.iv.1950,
LE Pena, CUIC); Buena Vista (1 w, 20.ii.1999, L Stange, FSCA); Lomas de Arena
Biol. Park (4 w, 10.ii.1999, LA Stange, FSCA); Velasco, Santa Cruz de la Sierra (1 w,
J Steinbach, MCZC); Santa Cruz de la Sierra (1 w, 23.iv.1989, G Morales, LACM).
BRAZIL, GOIAS: Anápolis (1 w, 12.ii.1936, G Fairchild, MCZC); 24 km E Formoso
(1 w, 29.v.1956, FS Truzal, LACM); (1 w, 30.iv.1956, FS Truxal, LACM); Parque Na-
cional da Chapada dos Veadeiros (1 w, 29.xi.1989, J Cuellar, LACM); Parque Nacional
da Chapada dos Veadeiros 18–24km N of Alto Paraiso 1400–1500m (1 w, 2–5.x.1985,
SE Miller, LACM), Parque Nacional da Chapada dos Veadeiros, 18–24km N of Alto
Paraiso 1400–1500m (1 w, 22.iv.1956, FS Truxal, LACM); MATO GROSSO: Rio
das Mortes nr. São Felix do Araguaia (1 w, 1944, JV Ca, MCZC); MATO GROSSO
DO SUL: 24 km W Campo Grande loose on ground (1 w, 7.xi.1989, WP Mackay,
CWEM); 6 km SE Campo Grande nest in soil (2 w, 8.xi.1989, WP Mackay, CWEM);
8 km SE Ponta Purá, loose on ground (2 w, 15.xi.1989, WP Mackay, CWEM); 3 km
NW Taunay, loose on ground (1 w, 17.xi.1989, WP Mackay, CWEM); Urucum, Co-
rumbá (1 w,23–29.xii.1919, CUIC); SÃO PAULO: Corumbataí, loose on ground(1
w, 6.xi.1989, WP Mackay, CWEM); RIO GRANDE DO SUL: Passo Fundo (1 w,10.
iii.1939, PA Berry, USNM). PARAGUAY, ALTO PARANÁ: Villa Encarnación (1 w,
10.i.1905, CASC); GUAIRÁ: Rogue González (1 w, 14.i.1995, B. Garcete and Alex
Wild, LACM).
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 139
Dinoponera gigantea (Perty)
http://species-id.net/wiki/Dinoponera_gigantea
Figs 1D, 4A, F, K, 5C, 9C, 10C, 11B, C
Ponera gigantea Perty, 1833: 135, pl. 27, Fig. 3. (worker) BRAZIL: Amazonas, Rio
Negro [type not found]; Kempf, 1971: 372 (male); combination in Dinoponera,
Roger, 1861:38.
Ponera grandis Guérin-Méneville, 1838: 206 (worker) [type not found]; combina-
tion in Dinoponera, Roger, 1861:38; junior synonym of gigantea: Roger, 1861:
38; Kempf, 1971:371. Emery, 1911: 219 (male); Wheeler, G.C. and Wheeler, J.
1985: 387 (larvae).
Worker diagnosis. Dinoponera gigantea can be distinguished from other Dinoponera
species by the following combination of character states: antero-inferior corner of pro-
notum with distinct tooth-like process (Fig. 1D); integument nely micro-sculptured
and not shiny (Fig. 12B); drab pilosity notably dense, long and agellate; scape length
longer than head width; total body length over 30 mm. Dinoponera gigantea is the larg-
est species in the genus reaching up to 3.6 cm total body length. Dinoponera gigantea
can be separated from all but two species by the presence of a tooth-like process on
the antero-inferior corner of the pronotum. Dinoponera lucida and D. australis have
a tooth-like process on the pronotum, but are smaller (usually less than 30 mm). In
addition D. lucida has a shiny integument and D. australis lacks the long, agellate
pubescence.
Description of the worker. Measurements (mm) (n=15) TBL: 31.62–36.02
(34.34); MDL: 4.59–5.35 (4.92); HL: 5.89–6.65 (6.31); HW: 5.74–6.27 (6.00);
SL: 5.95–6.43 (6.30); WL: 8.71–9.94 (9.35); PL: 2.72–3.06 (2.81); PH: 3.08–3.67
(3.59); PW: 1.85–2.07 (1.98); GL: 9.43–12.24 (10.94); HFL: 8.10–9.3 (8.74). A de-
scription of the external morphology of the worker is given in Kempf (1971):
"Length of scape exceeding maximum width of head. Pubescence on front of head quite
dense yet inconspicuous, not concealing the integument. Gular (ventral) surface of head
reticulate-punctate throughout, very nely striate in front, the striae strongly converging
mesad toward the anterior border. Sides of head reticulate-punctate, subopaque. Antero-in-
ferior corner of pronotum dentate. Pronotal disc reticulate-punctate, subopaque, the paired
swellings rather inconspicuous, but the median impression between swellings distinct, integ-
ument irregularly wrinkled. Tarsus I of hind leg longer than maximum head length. Petiole
reticulate-punctate and subopaque, rectangular in prole, the anterior surface straight to
slightly concave; the anterior upper corner more narrowly, the posterior corner more broadly
rounded; posterior surface with the vertical sulcus always distinct; in dorsal view the peti-
ole is decidedly longer than broad, width-length proportion below 0.80. Terga I and II of
gaster opaque, sharply reticulate-punctate, densely foveolate (from each foveola arises a short
decumbent hair), with scattered, bristle bearing, larger pints. e appressed pubescence,
although inconspicuous, is densely spread over the entire terga, stridulatory le on acroter-
gite (portion of tergum that is normally concealed under the overlapping preceding tergum)
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
140
of tergum II short, narrow, inconspicuous, not crossing beyond anterior half of acrotergite
(hence not easily seen if entire acrotergite is not exposed)."
Male diagnosis. Males of this species are easily recognized by their funiculus which
is covered in long standing setae (Fig. 4F), shiny dark reddish brown integument and
the combination of a long pygidial spine (Fig. 4K), volsella with two basal teeth, lobed
end of digitus volsellaris (Fig. 10C) and deep concavity on the side of the penis valve
of the aedeagus (Fig. 11C).
Description of the male. A description of the external morphology of the male
is given in Kempf (1971). Measurements done by Kempf (1971) are included as only
one male D. gigantea was examined by us while the measurements of Kempf (1971)
most likely represent the means of the four males examined in that study:
"Measurements in millimeters: total length 22.0; maximum length of head capsule
2.48; maximum width of head (eyes included) 3.10; maximum diameter of eyes 1.86;
scape length 0.93; length of funicular articles I: 0.21, II: 1.86; Weber’s length of thorax
7.12; hind femur length 5.57; hind tarsus I length 5.38; petiole length 2.16, width 1.24,
height 1.76; tergum I of gaster length 3.09, width 2.88; fore wing length 15.6; hind
wing length 12.15. Chestnut brown, smooth and shining except funiculi, clypeus, front,
tibiae and tarsi which are nely punctuate to reticulate-punctate; terga III and following of
gaster weakly, supercially and nely reticulate. e entire insect covered with long, subde-
cumbent, silky pubescence, except funiculi where the pubescence is minute. Standing hairs
long and abundant on body, lacking on mid-dorsum of terga II-V of gaster; long hairs on
scapes rather numerous, length not much longer than twice the diameter of scape…Anterior
border of labrum rounded, not visibly excised… Pygidial spine long and well developed.
Parameres (gonostyli) of genitalia in side-view narrow and spear-pointed… Hypopygidium
(subgenital plate of subandrium) apically rounded…"
Description of male genitalia. Basal ring with wide dorso-anterior loop struc-
tures, dorsal depression on basal ring posterior to cleft between dorso-anterior loops,
ridge extending from anterior of depression to center; parameres long, narrow, round-
ed spade-shape, emarginated ventro-basal edge (Fig. 9C); volsella with nger-like cus-
pis volsellaris and broad cusp-like digitus volsellaris, cuspis volsellaris with pointed
end, ange extending on dorsal edge, digitus volsellaris with numerous small circular
bumps, lobe on postero-dorsal edge, 2 teeth on medial ventro-basal corner of volsella,
posterior tooth with lobe on posterior edge; penis valve of aedeagus with lateral arm
of apodeme at anterior border, deep, wide, ventral concavity under ridge at base of
apodeme, distal edge of valve wedge-shaped, proximal ventral edge of valve ending in
tooth descending at roughly 45°, ventral edge with large laterally curved lip, serrated
edge (Fig. 11C), serrations facing laterally on either side of aedeagus in dorsal view
(similar to penis valve in Fig. 8A).
Distribution. Dinoponera gigantea has been found on the coast of Guyana, in the
Brazilian states of Amazonas, Pará including Marajo Island, Mato Grosso and Ma-
ranhão as well as the Loreto Province in Perú. Dinoponera gigantea is reported to be
common in un-ooded forests in the vicinity of Belém, Pará (Kempf 1971, Overal
1980) (Fig. 13). It is probable that D. gigantea is found in French Guyana, Surinam,
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 141
Venezuela and southeastern Colombia because these regions are adjacent to known D.
gigantea localities and have similar lowland rainforest habitat. However, no specimens
from these nations are known to us, perhaps as a result of a lack of sampling or the
range is absent from Colombia and southwestern Venezuela.
A record from Rio de Janeiro (from the CASC) is puzzling as it is disjunct from
the known range of D. gigantea. e most southeastern locality for D. gigantea is over
1,480 km to the nearest portion of the state of Rio de Janeiro. In addition, Rio de
Janeiro is in a well collected area where no other Dinoponera have been found. e
label reads ‘R.d.Janeiro, Brazil, D. Davis’ and the specimen agrees in all morphological
characters with D. gigantea. is locality is omitted from the species’ range map (Fig.
13) because of its questionable nature. If other collections can validate this locality it
would mean a signicant range extension for D. gigantea.
Discussion. Dinoponera gigantea is a valid species with a distinct suite of morpho-
logical characters listed in the diagnosis above. e validity of D. gigantea is strength-
ened by range overlaps with D. longipes and actual sympatry with D. hispida, both with
no integration of morphological characters.
Material examined. BRAZIL, PARÁ: Belém (6 w, v.1924, FX Williams, LACM, 7
w, vi.1924, FX Williams, LACM, USNM, 1 w, i.1938, GN Wolcott and LF Martorell,
USNM, 1 w, 19.ix.1943, MCZC, 1 w, vii.1961, WA Burk, LACM, 1 w, 17.iii.1964,
CE and ES Ross, CASC, 1 w, 16.ii.1975, ES Ross, CASC, 1 m, 21.vii.1979, JO
Schmidt, LACM); Belém APEG Forest ight trap (1 w, 29.vii-6.viii.1974, DG
Young, FSCA); Braganza (1 w, HB Merrill, USNM); Jabaty (1 w, v.1924, JF Illig-
worth, LACM); Marajo Island (3 w, viii–x.1907, HB Merril, FMNH); Mocajuba (1 w,
9.xii.1926, EG Holt, USNM); Peixe Boi (1 w, viii–x.1907, HB Merrill, FMNH); Río
Guamá nr. Belém (20 w, 10.xii.1976, CL Hogue, LACM); Tucuruí Margem esq. (1
w, 16.iii.1979, WL Overal, LACM); Utinga tract nr. Belém (1 w, 2.viii.1962, PF Dar-
lington, MCZC); Souza (2 w, but the 16.ix.1920, LACM, AMNH); locality not speci-
ed (1 w, HB Merrill, LACM); locality not specied (2 w, 1954, WM Mann, LACM,
USNM), 4 w, Baker, MCZC, CUIC, USNM, 4 w, ayer Expedition, AMNH, 1 w,
G Franch, AMNH); RIO DE JANEIRO: locality not specied (1 w, D Davis, CASC);
State and locality not specied (1 w, HH Smith and T Pergande, USNM). GUYANA,
CUYUNI-MAZARUNI: Dist. Bartica Kalacoon (1 w, 1916, AMNH).
Dinoponera hispida sp. n.
urn:lsid:zoobank.org:act:83696C4B-912A-4833-ACFD-A59E9CCD4D11
http://species-id.net/wiki/Dinoponera_hispida
Figs 1C, G, 2, 13
Worker diagnosis. Distinguished from other species by the following combination of
character states: conspicuous bristle-like setae covering the entire body but most pro-
nounced on the dorsum of the head, mesosoma, petiole and gaster (Fig. 1C); ne stria-
tions on dorsum of the head; integument smooth and shiny with bluish luster most
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
142
visible on sides of the head (Fig. 12A); antero–inferior corner of pronotum without
tooth-like process (Fig. 1E); petiole bulging at antero-dorsal corner; insertions of setae
on dorsum of petiole raised, papillate (Fig. 1G).
Description of the worker. Measurements (mm) (n=5) TBL: 30.39–31.83
(31.08); MDL: 4.20–4.51 (4.38); HL: 5.64–6.05 (5.86); HW: 5.02–5.33 (5.19); SL:
6.05–6.36 (6.22); EL: 1.23–1.33 (1.27); EL: 0.72–0.97 (0.84); WL: 7.89–8.71 (8.36);
PL: 2.5–2.56 (2.52); PH: 2.87–3.18 (3.05); PW: 1.33–1.54 (1.47); GL: 9.69–10.15
(9.95); HFL: 7.89–8.41 (8.14). (See Fig. 2) Entire body with short, thick, sti, subde-
cumbent to erect setae (Fig. 1C); integument black, smooth, shiny, appearing polished
with bluish luster (Fig. 12A). Head: mandibles long, linear, 7-toothed, large diastema
between basal tooth and six apical teeth; clypeus with two laterally projecting teeth
on anterior edge, bulges medially, extends posteriorly between frontal lobes, sparse
appressed setae from distal edges to disc of clypeus, short sti setae on anterior edge;
large bilobed labrum; ventral surface of head with varying amounts of ne striation,
erect bristle-like setae; gena with ne striations running from eye into clypeus; median
furrow running from termination of clypeus between frontal lobes to shallow pit in
middle of frons (with ocelli in two individuals of type series); frontal lobes raised and
conspicuous, with striations at posterior constriction; antennae all with erect bristle-
like setae, funiculus covered in minute appressed pubescence; scape long, extending
past posterior border of the head, covered in erect bristle-like setae; frons with sparse
pads of short appressed setae; entire head covered with erect to subdecumbent bristle-
like setae. Mesosoma: antero-inferior corner of pronotum rounded, without tooth-
like process; pronotal disc with slight bulges; mesonotum fused with propodeum and
episternum, separated by slight furrows; basilar sclerite large, ovaloid; propodeum with
broadly rounded dorsal outline; propodeal spiracle nearly vertical slit; sulcus running
from center of propodeum along lower edge of propodeal spiracle to posterior edge of
propodeum at dorsal edge of bulla; mesosoma and coxae with white pubescence, espe-
cially dense on basilar sclerite, appressed white pubescence along dorsal surface facing
medially, middle posterior dorsum of pronotum lacking appressed pubescence found
on mesonotum and propodeum. Legs: long, femur and tibia with sparse erect bristle-
like setae. One well-developed, antennae cleaning, comb-like spur on fore leg; spine-
like and less developed denticular comb on mesothoracic leg; spine and comb-like spur
on hind leg, posterior side of fore leg basitarsus with dense pads of golden setae; tarsus
or mesothoracic and hind leg with short, sti setae, tarsal claws bidentate. Petiole:
large and tabular with narrow attachments at base to the propodeum and gaster, nar-
row in dorsal view; ne erect setae on anterior surface above articulation with meso-
soma; bulging at antero-dorsal corner as in D. longipes; integument nitid, papillate at
insertion of setae on dorsal surface; keel-like subpetiolar process, anterior triangular
projection (Fig. 1G). Gaster: small protuberance at articulation of gastral sternite III
and the petiole, covered in erect setae; stridulatory le of varying size on acrotergite of
gastric tergum II; covered in bristle-like setae, sparse short appressed setae on terga I
and II; polished integument; posterior edges of the pygidium and hypopygidium with
characteristic rows of spines.
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 143
Male. Unknown.
Distribution. Known only from the type locality (Fig. 13).
Discussion. Dinoponera hispida is considered a valid species based on the above
mentioned morphological characters, as well as its sympatry with D. gigantea without
any visible character integration. Based on the morphological character states D. his-
pida is most similar to D. longipes and D. mutica. Dinoponera longipes diers greatly in
the setae which are agellate and golden, completely lacks gular striations and lacks the
raised insertions of setae on the petiole. Dinoponera mutica also diers in pilosity, pos-
sessing drab-colored agellate pubescence and lacks the bulging antero-dorsal corner
and papillate dorsum of the petiole.
Etymology. hispida, from the Latin hispidus: bristle, referring to the conspicuous
bristle-like setae covering the friend and body.
Type series. Holotype worker (MCZC) BRAZIL, Pará: Tucuruí, I.1979, Coll.
M. Alvarenga; Paratypes (CASC, CWEM, LACM, MZSP) BRAZIL, Pará, Tucuruí,
I.1979, Coll. M. Alvarenga” (1 worker). “Brasil Pará Tucuruí Margem esq. 21.II.1979,
Brasil Pará, WL Overal, Várzea” (1 worker). “Pará Tucuruí, 20.VIII.1979, Brazil PA,
R B Neto” (1 worker). “Pará Tucuruí, 19.VIII.1979, km 28, Brasil Pará WL Overal”
(1 worker).
Dinoponera longipes Emery
http://species-id.net/wiki/Dinoponera_longipes
Figs 1A, B, 3, 4C, H, M, 5E, 8, 9E, 10E, 11E, 13
Dinoponera grandis subsp. longipes Emery, 1901: 48 Holotype worker PERÚ: Cumbase
(MCSN) [examined]. Raised to species: Kempf, 1971: 375.
Worker diagnosis. is species can easily be recognized by the golden luster of its con-
spicuous long, agellate hairs especially on the frons. In addition this species has the
following combination of character states: pronotal corner rounded without tooth-like
process (Fig. 1E), no gular striations, a reective, smooth and shiny integument (Fig.
12A). All specimens have a petiole which bulges on the dorso-anterior edge except for
those from the Rio Madeira and Rio Negro in Brazil.
Description of the worker. Measurements (mm) (n=16) TBL: 30.85–34.75
(32.83); MDL: 4.61–5.33 (4.89); HL: 5.48–6.87 (6.12); HW: 5.23–5.84 (5.57);
SL: 5.54–6.56 (6.23); WL: 7.84–9.33 (8.51); PL: 2.46–2.82 (2.64); PH: 2.77–3.59
(3.28); PW: 1.44–1.85 (1.67); GL: 9.02–12.20 (10.67); HFL: 7.48–8.87 (8.36). A
description of the external morphology of the worker is given in Kempf (1971):
"Antennal scape from slightly shorter to slightly longer than maximum head width (index:
scape L/head W × 100 = 94–103). Pubescence on front of head (as well as on thorax and dor-
sum of gaster) golden brown, very dense and rather long. Gular face of head smooth and shining,
without vestiges of striae antero-laterally. Antero-inferior corner of pronotum obtusely angulate,
not dentate. Pronotal disc smooth and shining, but densely covered with piligerous punctulae;
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
144
paired swellings from faint to distinct; integument not wrinkled. Tarsus I of hind leg decidedly
longer than maximum length of head capsule. Petiole smooth and shining; shape resembling
that of quadriceps…, dorsal surface faintly to distinctly slanted backwards; width-length pro-
portion distinctly lower than 0.80: vertical sulcus on posterior surface present in specimens from
Acre Territory, Brasil, absent in specimens seen from Perú. Terga I and II of gaster smooth and
shining but densely covered with punctulae from which arises the long and dense pubescence
that covers the entire segments. Stridulatory le on acrotergite of Tergum II of gaster very short
but broadly triangular, not extending backwards beyond the anterior half of acrotergite."
Male diagnosis. Distinguished from other Dinoponera by the following combina-
tion of character states: funiculus of antennae with short, thick decumbent setae (Fig.
4H); pygidial spine (Fig. 4M) shorter than in D. gigantea and D. quadriceps but longer
and narrower than in D. australis and D. snellingi, volsella with broad basal lobe cov-
ered in minute teeth (Fig. 10E).
Description of the male. Previously undescribed. Measurements (mm) (n=2)
TBL: 19.78, 21.12; HL: 2.10, 2.26; HW: 2.67, 2.77; SL: 0.92, 0.92; EL: 1.49, 1.59;
EW: 0.923, 0.923; WL: 6.66, 6.66; FWL: 17.43, 15.38; HWL: 13.12, 11.48; PL:
1.90, 2.05; PH: 1.38, 1.54; PW: 0.97, 1.03; GL: 9.12, 10.15; HFL:5.23, 5.54. (See
Fig. 3) Integument: smooth and shining reddish brown, mesosoma slightly darker
than head, petiole, gaster. Head: Mandibles reduced, rounded and broad, lacking
teeth; palps elongated; labrum reduced, emarginated on distal margin, covered with
setae. Clypeus large, triangular, bulging medially, covered in appressed to subdecum-
bent setae; anterior tentorial pits large; frontal carinae reduced to slight ridge along
antennal socket; antennal sockets close, located at posterior apex of clypeus. Anten-
nae reddish brown; funiculus covered in minute appressed pubescence with thicker
bristle-like decumbent setae; scape shorter than second funicular segment, 1st funicular
segment reduced. Compound eyes large, along lateral side of head, deeply emargin-
ated border medially, ocelli bulging beyond margin of head, depressed area between
posterior ocelli. Entire head covered in short decumbent to erect setae, longer hairs on
clypeus and around ocelli (Fig. 4C). Mesosoma: covered in long dense suberect to de-
cumbent setae; pronotum triangular, exposed narrowly dorsally below scutum; scutum
of mesonotum large, bulging antero-dorsally, with 3 longitudinal ridges; small tegula
over insertion of forewing; scutellum domed, sides with vertical ridges, dorsal surface
smooth; basilar sclerite under hind wing, reduced; fused mesopleuron, separated by
furrow with mesosternite; metanotum, exposed between scutellum and propodeum,
reduced; mesoepimera, mesoepisternite and propodeum fused, rounded; coxa large,
conical, covered in long, dense subdecumbent to decumbent setae. Wings: covered in
dense minute setae, venation as shown in Fig. 5E. Legs: dark reddish brown integu-
ment; covered in minute subdecumbent to decumbent sti setae; one well-developed,
antennae cleaning, pectinate spur on fore leg; spine-like and less developed denticular
comb on meso-thoracic leg; spine and comb-like spur on hind leg; tarsal claws biden-
tate. Petiole: dark brown, narrow attachments at base to propodeum and gaster; petiole
humped dorso-posteriorly; subpetiolar process reduced, bulging slightly posteriorly,
covered in long erect setae. Gaster: large, cylindrical; covered in ne silvery suberect to
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 145
subdecumbent setae, longer on rst tergite and all sternites; rst gastric tergite round-
ed, elongated; pygidium terminating in short spine with a broad base, narrow tip,
shorter than in D. gigantea; cerci short, covered in erect setae; tabular subgenital plate
with posterior truncate and slightly emarginated with rounded corners. Genitalia: (Fig.
8) basal ring with wide, thin dorso-anterior loop structures; parameres long, narrow,
rounded distally, emarginated ventro-basal edge (Fig. 9E); cuspis volsellaris nger-like
with slight raised rounded bumps, digitus volsellaris broad cusp with numerous small
circular bumps, roughly triangular lobe at ventro-basal corner of volsella covered in
minute teeth (Fig. 10E); penis valve of aedeagus with lateral arm apodeme at anterior
border, ventral concavity under ridge at base of apodeme, ridge recurving dorsally near
distal edge of penis valve, distal edge wedge-shaped, proximal ventral edge of valve
ending in downward facing tooth, ventral edge serrated with large dorsally curved lip
(Fig. 11E), serrations facing laterally on either side of aedeagus in dorsal view (Fig. 8A).
Distribution. Dinoponera longipes have been collected in eastern Perú in the de-
partments of Loreto, Amazonas, Huánuco, San Martin and Pasco, as well as Ecuador
in the province of Pastaza. In Colombia it has been recorded near the Peruvian border
in the department of Amazonas. In Brazil, D. longipes has been found in Acre, Ama-
zonas as far east as Manaus, as well as along the Rio Madeira in Rondônia (Fig. 13).
Discussion. Doubt was raised by Kempf (1971) as to whether D. longipes was
a valid species. Since few specimens have been collected from western Brazil a clinal
variation in character form with D. gigantea or D. mutica was a possibility (Kempf
1971). e specimens we have examined from Brazil show no such integration. Ad-
ditionally, evidence of species validity comes from the unique morphology of the male.
Among the holdings at the CASC, males were located with 20 worker specimens from
Tingo Maria, Departamento de Huánuco, Perú; all D. longipes. e nearest known
locality of another species is D. gigantea 550 km away at Estirón Rio Ampiacu in the
Departamento de Loreto, Perú. ere is a possibility that these could be males of a yet
undiscovered species. However, relatively intensive collecting of Dinoponera in the area
by numerous collectors has not revealed any other form.
Material examined. BRAZIL, AMAZONAS: Uypiranga Rio Negro, 14 km from
Manaus, 81 m (1 w, x.1941, A Rabaut, AMNH); Tabatingo (2 w, MCZC, CUIC);
União Rio Madeira (1 w, iii.1921 or 1922, WM Mann, USNM); RONDÔNIA: Porto
Velho, Rio Madeira (1 w, Mann and Baker, USNM). COLOMBIA, AMAZONAS: 18
km N Leticia (1 w, 25.ii.1974, Sand J Peck, MCZC); Leticia (1 w, x.1977, F Castaño,
CWEM); Leticia, Rio Tacana, loose on ground (1 w, 3.viii.2002, L Mejia, UNAB); 5 km
N Zaragoza (1 w, 18.ix.1988, F. Fernández, CWEM); El Encanto, (9 w, 25.viii.1920,
CUIC, LACM, MCZC, AMNH);, La Sombra to El Encanto, (2 w, 23.viii.1920, CUIC,
AMNH). ECUADOR, PASTAZA: Moretecocha Ex. Barrido plataforma (1 w, 1–7.
vi.1996, J Naranjo,QCAZ). PERÚ, AMAZONAS: Km 292–296 E of Montenegro Ol-
mos-Maranón Hwy 700–800m (3 w, 21.i.1964, PC Hutchison and JK Wright, CASC);
Montenegro, Bagua 350m (3 w, 29.ix–2.x.1963, Wygodzinsky, AMNH); MADRE DE
DIOS: Cueva de Castillo nr. Tingo María 600 m (3 w, 7.viii.1982, JM Wilson, LACM,
1 w, 31.x.1970, J Schuster, LACM); Tingo María 670 m (1 w, Weyrauch, MCZC);
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
146
Tingo María 2200 ft. (1 w, 8.x.1946, JC Pallister, AMNH) (3 w, 12.x.1946, JC Pallister,
AMNH, 1 w, 28.x.1946, JC Pallister, AMNH, 1 w, 23.v.1947, JC Pallister, AMNH, 1 w,
1.vi.1947, JC Pallister, AMNH, 1 w, xii.1949, HA Allard, USNM, 2 w, 27.ix.1952, NA
Wells, CASC); Tingo María, Monson Valley (1 w, 18.ix.1954, EI Schlinger and ES Ross,
CASC, 3 w, 23.ix.1954, EI Schlinger and ES Ross, CASC, 1 w, 8.x.1954, EI Schlinger
and ES Ross, CASC, 2 w, 10.x.1954, EI Schlinger and ES Ross, CASC, 1 w, 19.x.1954,
EI Schlinger and ES Ross, CASC, 1 w,21.x.1954, EI Schlinger and ES Ross, CASC, 1
m, 2.xi.1954, EI Schlinger and ES Ross, CASC, 1 w, 3.xi.1954, EI Schlinger and ES
Ross, CASC, 1 w, 9.xii.1954, EI Schlinger and ES Ross, CASC, 1 m, 11.xii.1954, EI
Schlinger and ES Ross, CASC, 3 w, 15.xii.1954, EI Schlinger and ES Ross, CASC, 1 w,
23.xii.1954, EI Schlinger and ES Ross, CASC, 1 w, 13–17.ix.1956, C Gregoire, USNM,
1 w, 11.viii.1960, DA Young, USNM, 1 w,16.v.1964, CE andES Ross, CASC, 1 w, 9–12.
iii.1967, WL Brown, MCZC); Parque Nacional de Tingo María, Cueva de las Lechuzas
tropical rainforest window trap (1 w, 8–16.i.1983, A Newton and M ayer, MCZC);
Parque Nacional de Tingo María, Cueva de las Lechuzas, sweeping (1 w, 11.viii.1985,
JF Cornell, LACM); 14 km N Tingo María (1 w, 7.ii.1984, WN Mathis, USNM);
Parque Nacional de Tingo María, 6 km W Tingo María (1 w, 9.ii.1984, WN Mathis,
USNM); 12km SW Tingo María (1 w, 12–15.viii.1985, JF Cornell, LACM); Parque
Nacional de Tingo María, 660 m (3 w, 11–17.iv.1987, JE Eger, FSCA); Tambello Chico
Canyon, 13km S Tingo María, 800 m (1 w, vi.1983, CM Stevens, FSCA); LORETO:
Amazon Camp Rio Momón, nr. Iquitos, 97.5 m (1 w, 1–10.xii.1982, ES Ross, CASC);
Amazon Conservatory for Tropical Studies, 70 km NE Iquitos, extracted from nest,
lowland tropical wetforest (1 w, 9.vii.2002, RC Morgan, CASC); Amazon Safari Camp
Nr. Iquitos (1 w, 25.vi.1980, CL Hogue, LACM); Aventurama Camp Rio Napo/Rio
Yagua (2 w, one infected with fungus, CL Hogue, LACM); Boquerón 500m (5 w, 7–14.
vii.1965, J Schunke, LACM); Explornapo Camp 100mi NE Iquitos (1 w, 15.vii.1990, S
Dunkle, FSCA); Rio Napo at Sucusnui (1 w, 13.vii.1985, CL Hogue, LACM); PASCO:
Río Iscozazin (1 w, 8–19.vii.1961, FS Truxal, LACM); SAN MARTIN: Tarapoto (9 w,
A Vasquez, AMNH); UCAYALI: Balta Rio Curanja (2 w, vii.1966, A Gardner, FSCA);
department not specied, Upper Rio Huallaga (1 w, 29.x.1925, H. Bassler, AMNH, 1
w, 30.x.1925, H. Bassler, AMNH, 1 w, xi.1930, H Bassler, AMNH); DEPARTMENT
NOT SPECIFIED, Upper Rio Marañon (2 w, Orton, CASC, MCZC), Rio Marañon
(1 w, 10.vii.1930, AMNH); Rio Santiago (1 w, 15.ix.1923, AMNH, 1 w, 17.xi.1923,
AMNH); Middle Rio Ucayali (3 w, 1.x.1929, H Bassler, AMNH).
Dinoponera lucida Emery
http://species-id.net/wiki/Dinoponera_lucida
Figs 1H, 13
Dinoponera grandis subsp. lucida Emery, 1901: 48. Syntype workers BRASIL: Espírito
Santo, ex coll Fruhstorfer. (MCSN) [examined]. (specic locality of Vila Velha (Cidade
do Espirito Santo) proposed by Kempf (1971). Raised to species: Kempf 1971: 376.
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 147
Worker diagnosis. is species can be recognized by the following combination of
character states: anterior inferior pronotal corner with tooth-like process (Fig. 1D),
pilosity long and agellate with white luster, integument smooth and shiny with bluish
luster (Fig. 12A), scape length longer than head width, petiole slanting obliquely on
dorsal edge (Fig. 1H). Total body length ranges from 27–30mm which is between the
lengths of D. australis and the other larger species.
Description of the worker. Measurements (mm) (n=5) TBL: 27.01–30.39
(28.64); MDL: 3.79–4.31 (3.97); HL: 4.92–5.64 (5.34); HW: 5.02–5.13 (5.07);
SL: 5.23–5.64 (5.42); WL: 7.33–8.20 (7.84); PL: 2.25–2.51 (2.39); PH: 3.18–3.28
(3.26); PW: 1.54–1.90 (1.72); GL: 8.00–10.05 (9.10); HFL: 6.87–7.28 (7.18). A de-
scription of the external morphology of the worker is given in Kempf (1971):
"Antennal scape distinctly longer than head width. Pubescence on front and vertex of head
variable, either short and inconspicuous or longer, denser and quite visible. Gular surface of head
nely striate either throughout or at least on anterior half; very seldom the striae are conned to a
narrow stripe along the anterior border and obsolescent yet still discernible. Sides of head smooth,
not quite glossy but with a silky sheen on account of the supercial reticulate microsculputure.
Antero-inferior corner of pronotum dentate. Pronotal disc smooth and shining, lacking wrinkles
and dense, ne punctulae; paired swellings quite distinct. Hind tarsus I decidedly longer than
head length. Petiole…smooth and polished, its anterior face not excavate, its dorsal face slanted
forward; vertical sulcus on posterior face either present or absent; width-length proportion well
under 0.80; anterior face lacking dense pubescence. Terga I and II of gaster smooth, highly shin-
ing, lacking dense, ne punctulae; pubescence loosely scattered on sides, entirely absent on disc.
Stridulatory le on acrotergite of tergum II well developed, nearly crossing the entire tergite."
Male. Unknown.
Distribution. is species inhabits fragments of Atlantic rainforest in the Brazil-
ian state of Espirito Santo, across the border into Minas Gerais, the southern portion
of Bahia and São Paulo (Fig. 13). It is possible that D. lucida exists in Rio de Janeiro
but we are not aware of any specimens from this area. Refer to Mariano et al. (2008)
for information on the biogeography of this species. With the locality data available D.
lucida is the only species with no known range overlaps with other Dinoponera species.
Discussion. Dinoponera lucida appears to be a valid species based on the unique
suite of characters including a tooth-like process on the pronotum, smooth and shiny
integument, long and agellate pilosity and petiole slanting forward on the dorsal
edge. However, the limited sample size (n=5) restricts the certainty with which we
can assert that D. lucida is a separate species because a broad spectrum of intraspecic
variation may not be represented. ere may be a possibility of character integration
with D. australis in the area between the states of São Paulo and Rio de Janeiro. Di-
noponera lucida is only slightly larger than D. australis but diers in its integument
micro-sculpturing and pilosity type (see the diagnosis above). Dinoponera lucida can
be confused with D. australis but is distinguished by its shiny integument and whitish
setae, as opposed to the micro-sculptured integument and dull tan setae of D. australis.
Dinoponera lucida has been classied as endangered in Brazil by the Ministério do
Meio Ambiente (Campiolo et al. 2003) due to habitat destruction in the Atlantic forest.
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
148
Material examined. BRAZIL, BAHIA: Itabuna, Itapebi 177 m (1 w, ii.1999,
JRM Santos, CASC); ESPIRTIO SANTO: Santa Tereza (1 w, 24.xi.1954, A Ruschi,
MCZC); MINAS GERAIS: Ataléia (1 w, 27.i.1994, I Cardoso, LACM).
Dinoponera mutica Emery
http://species-id.net/wiki/Dinoponera_mutica
Figs 1E, F, 12B, 13
Dinoponera grandis subsp. mutica Emery, 1901: 48 Syntype workers, BRAZIL: Mato
Grosso, Germain (leg.) (MCSN) [examined] (specic locality of Rondonópolis
proposed by Kempf (1971); Wheeler, G.C. and Wheeler, J. 1952: 607 (larvae.);
raised to species Kempf, 1971: 378.
Worker diagnosis. Dinoponera mutica is identied by its smooth and shiny integu-
ment with a bluish luster (Fig. 12A), a rounded pronotal corner lacking a tooth-like
process (Fig. 1E), gular striations on the ventral surface of the head, long and agellate
pubescence, scape length longer than head width and petiole with even dorsal corners
(Fig. 1F).
Description of the worker. Measurements (mm) (n=12) TBL: 29.42–32.34
(30.99); MDL: 4.10–5.48 (4.71); HL: 5.13–6.30 (5.65); HW: 5.13–5.64 (5.39);
SL: 5.43–6.05 (5.72); WL: 7.53–8.61 (8.20); PL: 2.26–2.67 (2.41); PH: 2.82–3.38
(3.17); PW: 1.54–1.90 (1.76); GL: 8.61–11.99 (10.06); HFL: 7.18–8.00 (7.60). A
description of the morphology of the worker is given in Kempf (1971):
"Antennal scape remarkably longer than head width. Pubescence on front and vertex
generally longer and denser than in gigantea, but lacking the golden luster on longipes.
Gular face smooth and shiny, with ne, more or less distinct striation antero-laterally
and antero-mesially (sometimes nearly eaced). Sides of head smooth and shining in spite
of the very ne, supercial microsculpture which is reticulate-punctate. Antero-inferior
corner of pronotum obtusely angulate or rounded. Pronotal disc smooth and shiny, lack-
ing irregular fossae and wrinkles; the paired swellings rather weakly expressed. Tarsus I
of hind leg decidedly longer than head width. Petiole of distinctive shape…, shorter than
that of gigantea and longipes, but width-length proportion still under 0.08; anterior
and posterior upper corners subequally rounded; smooth and shining; vertical sulcus on
posterior face usually obsolete, present only in one Bolivian specimen. Terga I and II of
gaster very indistinctly, supercially and nely reticulate-punctate yet quite smooth and
shining, lacking the dense foviolae of longipes on disc where the pubescence is likewise
scarce. Stridulatory le well-developed, triangular but short, visible only when acrotergite
of tergum II is fully exposed."
Male. Unknown.
Distribution. Dinoponera mutica is found in central South America in the Brazil-
ian states of Rondônia, Mato Grosso, Goias and Mato Grosso do Sul, in eastern Bolivia
and northwest Paraguay (Fig. 12).
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 149
Discussion. Dinoponera mutica is a valid species based on our study. Dinoponera
quadriceps is the closest to D. mutica in terms of morphological characters and is not
synonymized in this work because of the dierences stated in the diagnosis above.
Males of D. mutica may provide further support for separation from D. quadriceps.
Dinoponera quadriceps has a nely micro-sculptured integument which is not shiny
(Fig. 12B), lacks gular striations and has a petiole which bulges on the dorso-anterior
edge. Dinoponera longipes and D. hispida may also be confused with D. mutica but this
species lacks the dense golden pubescence of the former, or the short, sti setae and
forward bulging petiole of the latter.
Material examined. BRAZIL, MATO GROSSO DO SUL: Corumbá (1 w,
MCZC); Urucum, Corumbá (3 w, 23–29.xii.1919, LACM, CUIC); RONDÔNIA:
7 km NW Costa Marques (1 w, 16.xi.1986, R Wilkerson, FSCA); Schmitt Ranch
(1 w, ix.1996, R. Rogers, PALC). BOLIVIA, SANTA CRUZ: Perseverancia (1 w,
18.iii.1990, P Bettella, LACM). PARAGUAY, BOQUERÓN: Enciso (1 w, T. del
Sinne, CWEM).
Dinoponera quadriceps Kempf
http://species-id.net/wiki/Dinoponera_quadriceps
Figs 4B, G, L, 5A, 6A, 10A, 11A, 12B, 13
Dinoponera quadriceps Kempf, 1971: 380, rst available use of Dinoponera grandis st.
mutica var. quadriceps Santschi, 1921: 84; unavailable name, BRAZIL: Pernam-
buco: São Lourenço da Mata, Tapera (NHMB, examined).
Dinoponera mutica var. Mann, 1916, male
Dinoponera gigantean mutica var. quadriceps Borgmeier 1937 male designated BRA-
SIL: Pernambuca, Tapera.
Dinoponera opaca Santschi, 1921. Holotype worker BRAZIL: Rio Janeiro (Goeldi) (1
w NHMB, examined); junior synonym of D. quadriceps Kempf 1975: 344
Worker diagnosis. is species is recognized by its nely micro-sculptured integu-
ment which is not shiny (Fig. 12B), rounded anterior inferior pronotal corner lacking
a tooth-like process (Fig. 1E), ventral side of the head lacking any gular striations and
long/agellate pilosity.
Description of the worker. Measurements (mm) (n=17) TBL: 28.09–33.73
(30.60); MDL: 4.10–5.05 (4.53); HL: 5.23–6.04 (5.58); HW: 5.33–5.97 (5.56);
SL: 5.54–6.12 (5.80); WL: 7.38–9.03 (8.20); PL: 2.26–2.68 (2.50); PH: 3.06–3.52
(3.26); PW: 1.64–1.99 (1.80); GL: 8.20–11.93 (9.80); HFL: 7.18–8.11 (7.65). A de-
scription of the worker is given in Emery 1911, Mann 1916, Borgmeier 1937, Kempf
1971. Presented below is that of Kempf (1971):
"Antennal scape notably longer than head width. Pubescence on front and vertex of
head short and inconspicuous. Gular surface of head reticulate-punctate, subopaque, but
lacking arcuate striae except for some cases when a few short and vestigial striae appear
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
150
antero-laterally, just behind the mandibular insertion. Sides of head reticulate-punctate,
subopaque. Antero-lateral corner of pronotum obtusely angulate (very seldom subdentate).
Pronotal disc reticulate-punctate, subopaque, occasionally slightly wrinkled, bristle pits ir-
regular in outline; paired swellings very faint and obsolete. Hind tarsus I longer than head
length. Petiole… of distinctive shape, the anterior surface being slightly inclined forward
and often a bit excavate; anterior upper corner narrowly, the posterior corner very broad-
ly rounded; integument minutely reticulate-punctate and subopaque; sulcus on posterior
surface always distinct. Terga I and II of gaster reticulate-punctate and opaque; piliger-
ous pits for pubescence discally greatly scattered (in a few southern specimens from Bahia
State, these pits are stronger and denser, almost as in gigantea); coarse bristle-bearing pits
greatly scattered: pubescence rather scarce on dorsum, denser and more conspicuous on sides.
Stridulatory le on tergum II of gaster weakly developed, arising from the anterior border
of acrotergite and running streak-like across the anterior half of the same (visible only when
acrotergite is fully exposed; observed in ve specimens)."
Male diagnosis. Males of this species are distinguished by the long ne setae of the
second funicular segment (Fig. 4G), light brown coloration, long narrow parameres
(Fig. 9A), volsella with two small basal teeth and lacking a lobe on the distal edge of
digitus volsellaris (Fig. 10A).
Figure 6. Schematic drawing of generalized Dinoponera male genitalia in situ; genitalia capsule and dis-
articulated aedeagus, volsella and parameres.
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 151
Description of the male. Total length 21mm (Mann 1916) 22 mm (Borgmeier
1937). A description of the male is given in Emery (1911), Mann (1916), Borgmeier
(1937), and Kempf (1971). Mann (1916) described the male as follows:
"Head, including the mandibles, as broad as long, very convex behind. Eyes very large
and long occupying the entire sides of head, the inner border deeply emarginate; ocelli very
large and convex. Clypeus convex, the anterior border truncate. Mandibles small, pointed at
apex, with a small tooth at middle of inner border. Antennae a little shorter than the body;
rst funicular joint twice as broad as long; joints 2–11 very long, cylindrical, each slightly
shorter and more slender than the preceding. orax [= mesosoma] robust; scutellum short,
triangular, broadly rounded at apex. Epinotum [= propodeum] evenly rounded, without
distinct base or declivity, unarmed. Petiole nearly twice as long as broad, narrowed in front,
with nearly straight sides; in prole longer than high, attened above…the anterior slop
gradual, more abruptly sloping behind, the antero-ventral surface with a broad, triangular
projection. Gaster long and slender, the three times the breadth. Genitalia prominent; the
valves board, rounded at apex; cerci long and slender…Wings large extending almost to
the tip of gaster … Legs very long and slender…Body and legs shining. Antennae opaque,
coarsely, densely punctured; sparsely pubescent, and having much very long, ne erect hairs,
which on the apical joints are shorter and conned to the tips; pubescence of apical joint
more dense than the rest. orax (=mesosoma ) with long silky pubescence, most abundant
on the pleurae, and very ne re erect hairs sparsely distributed node without pubescence, but
with abundant erect hairs. Gaster with a thick mat of silky pubescence, shorter and ner
than that of the thorax (=mesosoma); lateral and apical portions with ne erect hairs…
Color rufous, the antennal scape and the rst ve funicular joints fucous. Wings lightly
infuscated, veins and stigma reddish brown. Pubescence yellowish white, exempt the long
antennal hairs which are black."
To this Borgmeier (1937) added that the petiole was “rounded on top”, “the sting
of the pygidium [=pygidial spine] long; subgential plate with apex slightly concave”,
and that the wings were 16mm long and “slightly yellowish”. Kempf (1971) noted the
dorsum of the gaster lacked standing hairs.
Description of the male genitalia. Basal ring with wide, thin dorso-anterior loop
structures; parameres distinctly long, narrow, rounded end, emarginated ventro-basal
edge (Fig. 9A); cuspis volsellaris nger-like with few rounded bumps on medial face,
digitus volsellaris broad cusp-like with numerous small circular bumps, 2 teeth at
ventro-basal corner of volsella (Fig. 10A); penis valve of aedeagus with lateral arm of
apodeme at anterior border, slight ventral concavity under ridge at base of apodeme,
distal edge wedge-shaped, proximal ventral edge of penal valve ending in anterior
facing tooth, ventral edge with large dorso-laterally curved lip with serrated edge,
serrations facing laterally on either side of dorsally curved lip (similar to penis valve
in Fig. 8A).
Distribution. Dinoponera quadriceps is found in the Caatingas, Cerrados, upland
humid forest and Atlantic forest (Kempf 1971, Paiva and Brandão 1995) in the north-
eastern Brazilian states of Alagoas, Bahia, Ceará, Paraiba, Pernambuco and Rio Grande
do Norte (Fig. 13).
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
152
Discussion. Dinoponera quadriceps as characterized by Kempf (1971) is main-
tained as a valid species by our analysis. Dinoponera quadriceps may be confused with
D. mutica, but has a nely micro-sculptured integument which is not shiny (Fig. 12B),
lacks gular striations and has a petiole which bulges on the dorso-anterior edge in con-
trast to D. mutica’s roughly microsculptured integument, striated gula and petiole with
even, non-bulging corners (Fig. 1F).
We also agree with the synonymy of Dinoponera opaca by Kempf (1975) after
examination of the type. Dinoponera quadriceps and D. mutica dier in micro-sculp-
turing, gular striations and petiole shape. Distribution records show a distance of over
900 km between the two species, but if specimens are found with an integration of
characters in the area of Tocantins and northern Goias than these species should be
synonymized.
Material examined. BRAZIL, ALAGOAS: Pedra (1 w, viii.1939, A Muller,
AMNH); CEARÁ: Tianguá (1 w, 6.iv.1972, JS Bowman, MCZC); PARÁ: Óbidos
(1 w, ii.1981, CWEM); Rio Tapajoz region (1 w, viii.1983, CWEM); Santarém (1 w,
20.v.1984, CWEM); PARAIBA: Independencia (1 w, 1 m, Mann and Heath, USNM,
2 w, LACM); João Pessoa (4 w, 21.iv.1975, J Kesselring, CASC, 1 w, i.1976, BA
Bkaul, CWEM); João Pessoa forest of Gargau primary forest on ground 45m (1 w,
22.i.1981, G Ekis, MCZC); RIO GRANDE DO NORTE: Baixa Verde (2 w, WM
Mann, USNM, 1 w, gift of Wheeler, MCZC); Ceara-Mirim (1 w, 1 m, WM Mann,
USNM); Natal (6 w, WM Mann, AMNH, LACM, MCZC, USNM); São José do
Bonm (1 w, 22.iii.1945, HT Dalinat, LACM); state not specied, North Piari (1 w,
vi–vii.1944, L Parker, MCZC).
Dinoponera snellingi sp. n.
urn:lsid:zoobank.org:act:791CAB8B-6A94-47FD-B379-5CC85D1A9947
http://species-id.net/wiki/Dinoponera_snellingi
Figs 4D, 4I, 4N, 5B, 7, 9B, 10B, 11B, 13
Worker. Unknown.
Male diagnosis. Specimens of this species are distinct in several respects. e com-
bination of a bicolored body and head possessing bulging compound eyes and ocelli
(Fig. 4D) is unique to this species. More denitive is the shape of the aedeagus which
possesses a large ventral lobe and nger-like serrated ange (Fig. 11B). e short broad
digitus volsellaris with nely toothed basal lobe (Fig. 10B) is distinctive, as well as the
paramere shape (Fig. 9B).
Description of the male. Measurements (mm) (n=3) TBL: 16.14–17.09 (16.58);
HL: 1.90–2.05 (1.98); HW: 2.36–2.51 (2.44); SL: 0.62–0.72 (0.65); EL: 1.23–1.38
(1.32); EW: 0.72–0.82 (0.79); WL: 5.54–6.05 (5.77); FWL: 13.33–13.63 (13.43);
HWL: 9.93–10.46 (10.25); PL: 1.44–1.54 (1.49); PH: 1.13–1.23 (1.16); PW: 0.92–
1.13 (1.04); GL: 6.66–7.18 (6.94); HFL: 4.20–4.92 (4.54). Integument: smooth and
shining; head, mesosoma and petiole dark brown to black; gaster light brown. Head:
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 153
Mandibles reduced, rounded, lacking teeth, rounded lobe on ventro-basal edge, high
lateral ridge running along axis; palps elongated; labrum reduced, deeply emargin-
ated on distal margin, covered with setae. Clypeus large, triangular, bulging medi-
ally, covered in appressed to subdecumbent setae; anterior tentorial pits large; frontal
carinae reduced to slight ridge along antennal socket; antennal sockets close, located
at posterior apex of clypeus. Antennae: black; funiculus covered in minute, dense,
sti subdecumbent setae (Fig. 4I); scape shorter than second funicular segment, 1st
funicular segment reduced. Compound eyes large, along lateral side of head, deeply
emarginated border medially. 3 ocelli at posterior margin of head, bulging beyond
margin of head, depressed area between posterior ocelli. Entire head covered in short
decumbent to erect setae (Fig. 3). Mesosoma: covered in short suberect to decumbent
white setae; pronotum triangular, exposed narrowly dorsally anterior to scutum; scu-
tum large, bulging antero-dorsally, with 3 longitudinal carinae; small tegula over inser-
tion of forewing; scutellum domed, with sparse erect setae, sides with vertical carina,
dorsal surface smooth; basilar sclerite under hind wing reduced; fused mesopleuron,
separated by furrow with mesosternite; metanotum exposed between scutellum and
propodeum, reduced; mesoepimera, mesoepisternite and propodeum fused, rounded;
Figure 7. Dinoponera snellingi male genitalia. A dorsal view B lateral view C ventral view.
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
154
coxa large, conical, covered in dense subdecumbent to decumbent setae. Wings: cov-
ered in dense minute setae, venation as shown in Fig. 5B. Legs: black, covered in
minute subdecumbent to decumbent sti setae; one well-developed, antennae clean-
ing, pectinate spur on the fore tibia; spine-like and less developed denticular comb on
meso-thoracic tibia; spine and comb-like spur on hind tibia; tarsal claws bidentate.
Petiole: narrow attachments at base to the propodeum and gaster; petiole humped dor-
so-posteriorly; subpetiolar process reduced, bulging slightly posteriorly. Gaster: large,
cylindrical; covered in ne silvery suberect to subdecumbent setae; rst gastric tergite
broadly rounded; pygidium terminating in short, broad, triangular, spine (Fig. 4N);
cerci short, as long as pygidial spine, covered in erect setae; tabular subgenital plate
with posterior end rounded. Genitalia: (Fig. 7) basal ring with thick dorso-anterior
loop structures, reduced; parameres short, broad, rounded, large lobe on dorsal edge,
emarginated ventro-basal edge (Fig. 9B); volsella with rounded cuspis volsellaris with
raised rounded bumps on medial-ventral surface, digitus volsellaris with numerous
small circular bumps on lateral distal face, tuft of setae on ventro-distal edge, lobe on
basal ventral corner, covered in minute teeth (Fig.10B); penis valve of aedeagus with
Figure 8. Dinoponera longipes male genitalia. A dorsal view B lateral view C ventral view.
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 155
Figure 9. Dinoponera right basiparamere/paramere of known males, lateral view. A D. quadriceps B D.
snellingi C D. gigantea D D. australis E D. longipes.
Figure 10. Dinoponera right volsella of known males, lateral view. A D. quadriceps B D. snellingi C D.
gigantea D D. australis E D. longipes.
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
156
long lateral arm of aedeagal apodeme at anterior border, ventral concavity under ridge
at base of apodeme, dorsal edge broadly rounded, ventral tooth projecting into thin
anteriorly folded ange with heavy serration, rounded notch at base, large triangular
ventral lobe with nely serrated edge and vertical ridge running through middle of
lobe, edge of lobe continuing into lateral apical fold with serrated edge (Fig. 11B).
Distribution. Known only from type locality; Campo Grande, Brazil (Fig. 13).
Discussion. Dinoponera snellingi is a new species based on the suite of morpho-
logical characters presented in the diagnosis above. Most important are the shape of the
aedeagus, volsella and parameres all of which we consider apomorphic characters. e
type specimen males were unassociated with workers. Initially D. snellingi specimens
were considered males of D. australis; as workers of this species were collected at the
same location and at the same date (see D. australis materials examined). Additionally
the specimens shared the same character states of bicoloration and short pygidial spine
that Kempf (1971) used to designate D. australis. However, the size of the compound
Figure 11. Dinoponera right penis valves from the aedeagus of known males. A D. quadriceps B D. snel-
lingi C D. gigantea D D. australis E D. longipes.
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 157
Figure 12. Worker head, oblique antero-lateral view illustrating microsculpturing dierence. A Dino-
ponera mutica (this smooth integument type is also found in D. lucida, D. longipes and D. hispida) B D.
quadriceps (this rough integument type is also found in D. gigantea and D. australis).
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
158
eyes (compare Fig. 4D and 4E), bulging ocelli at the posterior of the head (compare
Fig. 4D and 4E), short broad volsella with large tear drop-shaped basal lobe (Fig. 10B)
and penis valve of the aedeagus with disto-lateral process, disto-ventral lobe and ser-
rated ange on the ventral edge (Fig. 11B) provide strong evidence supporting that
these male specimens represent a novel species.
We have compared male specimens of D. snellingi with those of D. australis col-
lected in nest series and found they dier in the characters listed above. Campo Grande
is within the range of Dinoponera mutica and there is a possibility that these specimens
represent the currently unknown males of D. mutica. However, the males of D. snellingi
Figure 13. Distribution map of Dinoponera species. Symbols in black are records added by this study;
open symbols are from literature sources (Kempf 1971, 1975, Araujo et al. 1990, Peeters et al. 1999,
Monnin and Peeters 1999, Fourcassié and Oliviera 2002, Monnin et al. 2003, Mariano et al. 2004,
Araújo and Rodriques 2006, Marques-Silva et al. 2006).
A revision of the giant Amazonian ants of the genus Dinoponera (Hymenoptera, Formicidae) 159
are closest in character states to the male of D. australis, the worker caste of which dif-
fers greatly in many characters from the other known Dinoponera workers including D.
mutica (see the Dinoponera australis discussion). erefore we hypothesize that the male
of D. mutica will most likely be similar to D. quadriceps or D. longipes, based on the
similar worker morphology, and the unknown worker of D. snellingi will be similar to
the worker of D. australis. Species groupings based on worker and male character states
overlap; leaving D. australis with D. snellingi allied and separate from the other Dino-
ponera species. Until associated workers are discovered, we contend that it is better to
describe these unique males rather than allow them to remain misidentied and unstud-
ied or describe them as males of D. mutica with only anecdotal evidence as justication.
Etymology. Named in honor of the late Roy Snelling who made considerable
contributions to the eld and spirit of myrmecology.
Type series. Holotype deposited in MZSP, BRAZIL, Mato Grosso do Sul, Campo
Grande, 12 Oct 1989, W.P. Mackay #12404, 2 paratypes, same locality, 8 Oct 1989,
#12359 collected at house light (deposited in the CWEM and MCZC).
Discussion
A synthesis of our understanding of Dinoponera morphological characters and geograph-
ic distribution supports the six species designations of Kempf (1971, 1975) as well as es-
tablishes the two new species Dinoponera hispida and Dinoponera snellingi. e collection
records from material loaned to us, in conjunction with those found in previous works
(Kempf 1971, 1975, Araujo et al. 1990, Peeters et al. 1999, Monnin and Peeters 1999,
Fourcassié and Oliviera 2002, Monnin et al. 2003, Mariano et al. 2004, Araújo and
Rodriques 2006, Marques-Silva et al. 2006) establishes range overlaps between 7 species,
including sympatry between D. longipes-D. gigantea, D. gigantea-D. quadriceps-D. his-
pida and D. australis-D. snellingi-D. mutica. Dinoponera lucida is the only species which
has no sympatric records nevertheless, a record in the state of Bahia, Brazil comes within
62 km of D. quadriceps. In all cases there is no perceptible integration of characters.
Worker characters, though seemingly indistinct upon rst inspection, allow rela-
tively easy identication of Dinoponera species. e most important characters are the
tooth on the antero-ventral corner of the pronotum, in conjunction with pilosity, micro-
sculpturing and body size. In males the pilosity of the funiculus, paramere shape, lobes
of the volsella and shape of penis lobe of the aedeagus are the dierentiating characters.
Kempf (1971) questioned the stability of characters between species’ ranges then
thought of as being mutually exclusive. e areas between D. longipes, D. mutica, D.
quadriceps and D. gigantea were specically questioned. With the exception of D. mu-
tica, our study has examined material that shows overlap between these ranges with no
integration of characters used to dene these species.
Despite the work presented here, many questions still remain in terms of Dinopon-
era taxonomy. e male caste is still undescribed in D. mutica, D. lucida and D. hispida
and the worker caste is unknown for D. snellingi. ese unknown castes likely exist
Paul A. Lenhart et al. / Journal of Hymenoptera Research 31: 119–164 (2013)
160
in the collections of Brazil and elsewhere. Ranges are roughly dened but our study
revealed several vast range extensions and country records. As evident in Fig.13, large
areas of South America remain uncollected.
Acknowledgments
We would like to give a special acknowledgment to the late Roy Snelling, who never
failed in providing advice and discussions on ant taxonomy. Our indebted apprecia-
tion is also given to two anonymous reviewers whose dedication and care improved
the quality of this manuscript. We would like to thank all the institutions and indi-
viduals that loaned us specimens including James Carpenter (AMNH), Brian Fisher
(CASC), E. Hoebeke (CUIC), James Boone (FMNH), Jim Wiley (FSCA), Roy Snel-
ling (LACM), C. Garcia (UNAB), Roberto Poggi (MCSN), Carlos Roberto Brandão,
(MZSP), Stefan Cover (MCZC), Daniel Burckhardt and Lisabelle Zürcher (NHMB),
Juan Vieira and David Donoso, (QCAZ) and Ted Schultz and David Furth (USNM),
as well as ibaut Delsinne who donated the rst Paraguayan specimen of D. mutica to
our collection, Becky Marin, Walker Johnson, and Cleat Zeiler for her help mapping
specimen localities in ArcGIS, Vicente Mata-Silva and Francisco Serna for their help
translating the Spanish abstract and helping with the Spanish key, Ana Yoshi Harada
for checking the Portuguese abstract and key, as well as Chris Schmidt, Dominic Lan-
nutti, Jose Pacheco, Israel del Toro, Cindy Morgan, Francisco Serna and Robert Len-
hart for all their advice. is revision was made possible by support from a National
Science Foundation grant for Undergraduate Mentoring in Environmental Biology
#26-1006-60, the American Museum of Natural History collection study grant award-
ed to S. T. Dash, the Ernst Mayr fund of the Museum of Comparative Zoology, and
the University Honors Program at the University of Texas at El Paso.
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